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[Reprinted from STUDIES IN THE BIOLOGICAL SciENCEs, No. 4.|

A REVISION OF THE NORTH AMERICAN SPECIES
OF THE SECTION BORAPHILA ENGLER
OF THE GENUS SAXIFRAGA
(TOURN) L.

A THESIS
SUBMITTED TO THE FACULTY OF THE GRADUATE SCHOOL

OF THE UNIVERSITY OF MINNESOTA

Lie ARY
BY A
% Reale SK

ARTHUR MONRAD JOHNSON

IN PARTIAL FULFILLMENT OF THE REQUIREMENTS
FOR THE DEGREE OF

DOCTOR OF PHILOSOPHY

JUNE 1919

1g9 2% [
bee 3 1%

A REVISION OF THE NORTH AMERICAN SPECIES OF THE
SECTION BORAPHILA ENGLER OF THE GENUS
SAXIFRAGA (TOURN) L.*

ARTHUR MonraAD JOHNSON
(With Plates I-XIX)

The section Boraphila was established by ENGLER (4) in his Index
Criticus Specierum atque Synonymorum Generis Saxifraga and was founded
on those species of the genus Saxifraga (Tourn) L. which Tauscu (30)
and later writers of the early part of the nineteenth century had grouped
under the sections Arabisa (Arabidia Seringe), Micranthes, and Hydatica.
In his later Monographie der Gattung Sagifraga ENGLER (5) retains the
classification as first set forth in the Index, and except for the enlargement
of the scope of the section the fundamental principles of this classification
are adopted in the more recent treatment in ENGLER and PrantL (6) Die
Natiirlichen Pflanzenfamilien.

In these studies the attempt will be made to show that the section
Boraphila as ENGLER persisted in construing it is a heterogeneous one, con-
sisting of species of widely different affinities, but which, however, can be
realigned into natural groups on the basis of certain fundamental characters.
Encuer himself was not unconscious of a certain amount of heterogeneity
in his section, however, for in his Monographie (p. 127) he makes this
statement: “Allerdings weichen die Artengruppen, welche ich unter der sec-
tion Boraphila zussammenfasse, mehr von einander ab, als die Artengruppen
anderer Sectionen....” Nevertheless he grouped them all in the one sec-
tion rather than make a number of new sections based on minor characters
(“geringfiigige Merkmale”).

In the Monographie ENGLER separates the section Boraphila on the shape
wp of the capsules and on their mode of dehiscence. But as far as these charac-
ts! ters are concerned his section Robertsonia is scarcely to be distinguished
~~ from certain species of Boraphila (namely those of the section Arabisa).
> And again the “capsule” of Peltiphyllum is scarcely distinguishable from

capsules of certain other species of Boraphila (for example, in Saxtfraga
oregana of the section Micranthes).
' © In Die Natiirlichen Pflanzenfamilien ENGLER includes in the section
es “= Boraphila the genera Spatularia, Micranthes, Dermasea, and Aulaxis of
cc >
)

>- HawortuH, Dermasea of Hoppe, and the sections Arabisa and Micranthes of

aes

~ _-Tauscn. No attention is given in the classification to the characters of the

i 3A thesis submitted to the faculty of the Graduate School of the University of Minnesota in
. partial fulfillment of the requirements for the degree of Doctor of Philosophy.

2 MINNESOTA STUDIES IN PLANT SCIENCE

carpels and the gland (or disc). The result is a more or less unnatural
grouping based on minor characters. We thus find a collection of species
of widely different affinities, as for example where Sawifraga erosa Pursh
(Saxifraga micranthidifolia Haw.) is grouped with Saxifraga Forbesu
Vasey, and where Saxifraga hieracifolia W. and K. is grouped with Sax-
ifraga pennsylvanica L. and Saxifraga integrifolia Hook. (See sections
Aulaxis, Micranthes, and Chionophila® respectively, in this paper.)

Turning now to the early work of Haworru (13) we find in that
writer’s Synopsis Plantarum Succulentarwm the first attempt to break up
the Linnaean genus Saxifraga, a task which he carried out more fully in
his later Saxifragearum Enumeratio. In the latter work Hawortu (14)
made sixteen genera out of the old one, retaining the name Saxifraga for
one of the genera. Without dwelling on the details of HAwortu’s work it
may suffice to state that he pointed out, a century ago, the fundamental
taxonomic distinctions between certain species which ENGLER many years
later embodied in the section Boraphila. Waworty first clearly rec-
ognized the significance of epigyny and hypogyny in the classification of
the species then known, and on these characters he based his generic dis-
tinctions. To illustrate, in establishing the new genus Aulaxis on the species
Aulaxis (Saxifraga) micranthidifolia Haw. (Saxifraga erosa Pursh)
Haworth clearly recognized the value of the characters of the carpels, as
witness his description: “capsulae superae alte bipartiiae basi solum coalitae,
in restora acuta subdivaricato progredientes.” The accuracy of his observa-
tion will be appreciated by referring to the figure of Aulaxis in Plate I. It
is strange that these characters should have escaped the notice of subsequent
investigators.

Of Hawortn’s other genera, including species with which we are here
concerned, Micranthes was reduced to sectional rank by D. Don (3),
TauscH (30), and SERINGE (26), and later by Torrey and Gray (31),
and by Hooker (16). The genus Robertsonia was reduced to sectional rank
by ENGLER (4). In a recent work of Smatt (27) Hawortnu’s genera
Micranthes and Spatularia are restored.

D. Don (3) grouped the species in two sections, Gymnopera and
Micranthes, basing the distinctions on the characters of the calyx. Thus
the calyx of the former section was described as “reflexed,” that of the
latter as “spreading.” Along with these he included descriptions of the
stamens and the carpels. Since the “reflexed” and the “spreading” condi-
tions of the calyx are characteristic of hypogynous and epigynous species

3 CHIONOPHILA sect. nov. Receptaculum ovario alte adnatum; folliculi elliptico-ovati plus
minusve inflati superne conspicue contracti, stylis plus minusve elongatis saepe crassiusculis, stig-
matibus globosis vel discoideis; glandula discoidea, margine inferiore calycis basi approximata; fila-
menta subulata plana ad aspicem incurva antheris oblongis; sepala patentia semina angusta ovata
margine minime alata plus minusve striata.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 3

respectively (but are distinctly characters of minor importance) the classi-
fication automatically tended towards a natural arrangement, the significance
of which Don seems not to have fully appreciated.

TauscH (30) grouped the species in three sections, Micranthes,
Hydatica, and Arabisa. He, too, based his distinctions on the characters of
the calyx. Thus the calyx of Micranthes was described as adhering to the
ovary and reflexed (“calyx adhaerens, ore reflexo’”), that of Hydatica and
of Arabisa as free and reflexed (“calyx liber, reflexus”). With the excep-
tion of the section Arabisa, where the species are well defined, Tauscu’s
reliance on minor characters resulted in the grouping together of species
of widely different affinities. The characters of the carpels were entirely
overlooked.

SERINGE (26). followed the classification of TauscH, with its attendant
confusion, in the relationships of the species. Neither SERINGE nor TAUSCH
appreciated the importance of epigyny and hypogyny, and HAwortH’s
accurate characterizations of the genera Robertsonia and Aulaxis seem to
have been completely ignored by them. Torrey and Gray (31) adhered
to the system of SERINGE, although they somewhat stressed the epigeous
and hypogeous character of the caudex, a character which is subject to
variation under different ecological conditions and unreliable as a character
of major taxonomic consideration. Hooxer (16) followed the same system
in Flora Boreali Americana.

The system of classification proposed by TAuscu and followed more or
less closely by the subsequent writers above mentioned was considerably
modified by LeprBour (18). He combined the sections Arabisa and

* Micranthes of Tauscu, giving the name Arabidia to the combination. Like

his predecessors, however, LEDEBOUR based his classification wholly on char-
acters of minor importance, which, together with the combination he intro-
duced, added still more to the confusion in the classification.

This first attempt at combination thus initiated by LEpEBourR reached
its culmination in the work of ENGLER, who combined into a single group
the sections that had been separated for nearly fifty years.

ROSENDAHL (25) considered the sections Micranthes, Arabidia, Spatu-
laria, and in part Hydatica, as subsections of ENGLER’s section Boraphila.
He regarded the structure of the floral axis and of the carpels as the
principal diagnostic characters of ENGLER’S section.

Contemporaneously with RosENDAHL, SMALL (27) published a revision
of the group, in which he digressed from the arrangement of previous
writers by restoring the genera Micranthes and Spatulartia of HawortH, and
at the same time adopting RAFINESQUE’s (24) genus Heterisia for the
monotypic Heterista (Saxifraga) Mertensiana Bongard. Sma. here rec-
ognized the hypoegynous character of the species included in these two

4 MINNESOTA STUDIES IN PLANT SCIENCE

genera, but his arrangement of the genus Micranthes is less fortunate since
it is made to include both epigynous and hypogvnous species. In a later
treatment of these species SMALI. (28) substitutes NECKER’s earlier generic
name Hvydatica for the Spatularia of HawortH. In other respects the
arrangement of the species described is essentially the same as that in North
American Flora (l.c.).

DISCUSSION OF THE PROPOSED CLASSIFICATION

The classification here proposed is founded on the characters of the
hairs, the receptacle, the carpels, and the gland.* All structures concerned
are shown in Plates I, II, and III. While the present work materially
increases the number of sections, the characters on which these are based
are believed to be of sufficient taxonomic importance to justify the separa-
tions.

The hairs were first made use of in the classification by Gopron (12),
and while ENcLER (5, p. 75) based his primary distinctions on seed charac-
ters he offers a substitute classification, based on the characters of the hairs,
for the larger categories.

Two types of hairs are to be found in the groups to be described, namely.
the wniseriate type (“ein-reihig” or “articulati” of ENGLER) and the multi-
seriate (“mehrreihig”’ of ENGLER). The former type is composed of a
single row, the latter of several rows of cells. The sections to be described
thus fall into two divisions, the uniseriate-hair type represented by the sec-
tions Arabisa and Hetertsia,* the multiseriate-hair type represented by the
remaining sections.

In some remarks on the use of hairs in classification ENGLER (5, p. 75)
makes the following statement: “Da die Haare soweit meine Beobachtungen
reichen, bei den Arten-der einzelnen Sectionen stets gleichartig sind, so ist
eine darauf gegriindete Uebersicht jedenfals natiirlich.” However, neither
in his Monographie nor later in Die Natiirlichen Pflantzenfamilien did he
adhere consistently to this statement inasmuch as he failed to recognize
the characters of the hairs in those species which, in the scheme of classi-
fication here proposed, are included in the sections Arabisa Tausch, and
Heterisia Rafinesque (the former being synonymous with the genus Spat-
ularia of HAworTH). ENGLER furthermore broke up the genus Kobertsonia
Haworth, setting apart in a new section, for which he retained the name,
a group of species with multiseriate hairs, but at the same time incon-
sistently failing to separate the species of HAwortu’s genus Spatularia
(Arabisa Tausch) on the same grounds.

®In this paper the disc or nectary is spoken of as the ‘‘gland.’”’
“Hererista (Raf.) sect. nov., Heterisia Raf. Fl. Tell. 2:69. 1836; as genus.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 5

Previous writers have placed considerable emphasis on caulescence in
their classifications, but since this character was found to vary considerably
it is here regarded as of little taxonomic value. Within certain limited
groups the petals and leaves are distinctive, and this is also true of the
filaments and the inflorescences. The petals of the section Arabisa, for
example, are distinctive of that section; so too are the leaves of the section
Hydatica, although very similar leaves occur in the section Calthophyllum.®
The filaments and the inflorescence of Calihophyllum are unmistakable.
Taken as a whole, however, these characters are only secondary in com-
parison with the carpels and glands. The same may be said of the calyx
segments, which soon become strongly reflexed in the hypogynous species,
but in the others the degree of reflexion may be greater or less, dependent
upon the degree of adherence of the receptacle to the ovary. In the latter
case they often become reflexed with the growth and expansion of the
ovary, and are therefore not very reliable for taxonomic purposes. Seed
characters are valuable criteria of relationship within certain groups, but
appear to have received little attention. Unfortunately most collections
are deficient in seed material. The distinctive character of the seeds is
well shown in the section Arabisa. And in the difficult section Micranthes
the seed characters sometimes proved to be the last resort in establishing
affinities.

To turn now more particularly to a discussion of the classification pro-
posed in this paper: as stated on a preceding page the sections are grouped
into two major divisions on the characters of the hairs; the sections of the
first division having “uniseriate,” those of the second “multiseriate,” hairs.
Two lines of development.are to be noted in the sections of the first division.
On the one hand we have a series of three sections in which the ovary is
wholly superior, namely Hydatica, Calthophyllum, and Aulaxis. In addition
the carpels are more or less united, two lines of evolution being indicated,
one in which the carpels are more or less bottle-shaped and erect, or spread-
ing from the base at maturity, the other in which the upper free portions
of the carpels are elongated and widely spreading—in V-shaped fashion—at
maturity, while the lower capsular portion becomes somewhat constricted
toward the base. The former condition is found in Hydatica and Cal-
thophyllum, the latter in Aulaxis, in which also the gland surrounds the
carpels as a thick girdle from the base to the middle. Marked differences
in the glands serve to distinguish Hydatica and Calthophyllum. In Hydatica
the gland lies close to the base of the carpels as a thick fleshy collar, and the

8 CALTHOPHYLLUM, sect. nov. Ovarium librum; carpidia conica ad basin minime contracta et
coalita singula infra medio glandula dilatata cincta; folliculi elliptico-oblongi distincti inferne et
superne maxime contracti, stylis brevibus attenuatis, stigmatibus parvis; filamenta primum clavata
demum maxime elongata filiforimia; sepala reflexa.

6 MINNESOTA STUDIES IN PLANT SCIENCE

follicles are erect at maturity and united either near the base or up to the
middle. In Calthophyllum the carpels are entirely distinct and are spread
apart as they increase in size, while the gland takes the form of a broad
zone surrounding the basal portion of the carpels. There are also outstand-
ing differences in the young carpels of these sections (Plate I).

On the other hand there is a series of sections in which the receptacle
is more or less united with the ovary. This condition has been termed
“subinferior.” Four sections comprise this series, namely, Dermasea,
Chionophila, Micranthes, and Tricarpum.' The carpels are in all cases
distinct to the base but are connected laterally by the adnate receptacle
and at least the lower portion of the gland. The sections may be grouped
into two divisions according to the number of carpels. Thus in Tricarpum
the number is uniformly three, although four are sometimes found. In the
remaining sections the number two is quite constant. The section Dermasea
possesses a very distinctive gland which is in the form of a fleshy ring around
the basal portion of the carpels in species having a very shallow receptacle.
In Chionophila and Micranthes the glands are in the form of a broad fleshy
and prominent disc, closely appressed to the base of the calyx in Chionophila,
but considerably elevated in Micranthes. In Tricarpum the gland is very
similar to that of Micranthes but is only slightly elevated. In both Micran-
thes and Tricarpum the margin of the gland persists as a conspicuous flange
on the sides of the carpels.. The species are readily referred to their respec-
tive sections on the characters of the gland. There is considerable variation
in the depth of the receptacle in Dermasea, where in exceptional cases
(Saxifraga nivalis) it is united to the carpels for about half their length.
This variation occurs in certain species of the section Micranthes, as for
example Saxifraga pennsylvanica and S. oregana, in which this condition
often gives the appearance of distinct hypogyny. The receptacle in Chiono-
phila, Micranthes, and Tricarpum is characteristically deep.

The second division comprises only two sections, Arabisa and Heterisia.
In both, the ovary is wholly superior and the carpels united about to the
middle. The carpels and glands indicate two distinct lines of evolution.
In Arabisa the young carpels are erect and broadly conical, terminating
above in mere peglike beaks, while the gland has the form of a narrow
girdle immediately above the base of the carpels. In Heterisia the carpels
are strongly contracted above into slender, strongly divergent beaks, and
are surrounded for the greater part of their length by a thick fleshy gland.

® DermMaseA (Haw.) sect. nov., Dermasea Haw., Saxif. Enum. 45. 1821, as genus.

7 TRICARPUM sect. noy., receptaculum ovario alte adnatum; carpidia tria distincta, receptaculo
et supra glandula conjuncta; folliculi crassi inflati stylis crassis divergentibus rigidis, stigmatibus
discoideis; glardula crassa ex calycis basi breviter elavata, margine inferiore persistente; filamenta
subulata; sepala patentia.

= aa a etal ea lala

JOHNSON: A REVISION OF THE-SECTION BORAPHILA 7

The differences between the mature follicles of these two sections are also
pronounced (Plate II).

The criteria of classification described above are therefore regarded as
fundamental, for the reason that they are fixed and quite obvious characters
that clearly define the limits of the relationships.

KEY TO THE SECTIONS

I. Hairs uniseriate
A. Ovary wholly superior; carpels more or less united
a. Carpels elliptic or oblong at maturity
1, Gland a fleshy basal cules s follicles erect, united to the middle or mereiy
UEMALMLEM RAS nano Meee es mice ULL NNER Lie ANS Sear ON ey 20 JOO a Section Hydatica
2. Gland a broad zone around the basal portion of the carpels; follicles dis-
CUTS PS) DLT SEE VEU AY 278 2 IAD ACAI AIAG GAIT Ue OR EN AIPA Section Calthophyllum
b. Carpels forming a V-shaped capsule at maturity; gland fleshy, surrounding
thewcanpels tonnear they middleseiecgi i.e Ue nioe ce weiss Section Aulaxis
B. Ovary subinferior; carpels distinct, connected by the adnate receptacle
a. Carpels two :
1. Gland a fleshy ring close to the base of the carpels and the calyx; recep-
tacle shallow to deeper, flabellate................0000. Section Dermasea
2. Gland a broad fleshy disc; receptacle deep
(a) Gland closely appressed to the base of the calyx................4..

BT eee oat nt oyalel wa tei clahatalls arate aterasle cle’cfeisl eielaiverat ates Section Chionophila

(b) Gland considerably elevated, thick, convoluted, its margin persisting
ASMA CASLITICLUL LATE! as ciecabe iar sievarciete\ sil elev Menerereateroe Section Micranthes

b. Carpels three, sometimes four; gland a broad fleshy disc slightly elevated
above the base of the calyx, and its margin persisting as a flange...........
Pr tet recuse eiag misters alladaisr anal valay aisles shareis el adel aiaiucel arene Section Tricarpum

Il. Hairs multiseriate; ovary superior; carpels united to the middle
A. Gland a narrow girdle above the base of the carpels; carpels elliptic at matu-

FitWeiin es. Ra SO HMR GLI ARN Aco eNO AD) CELLS MA 2 UU Va Section Arabisa
B. Gland a broad fleshy zone surrounding the ovary; carpels (capsule) pyriforin

atimatunity, strongly contracted abOve.)c. 24.26. oo. eres ce Section Heterisia

Section Hypatica Tausch, Hort. Canal. Fasc. 1. 1823.

Robertsonia Haw., Syn. Pl. Succ. 321. 1812, as genus; in part.
Gymnopera D. Don, Trans. Linn. Soc. 13 :343. 1822; in part.
Micranthes D. Don. l.c.

Arabidia, Seringe, DC. Prodr. 4:40. 1830; in part.

Boraphila Engler, Verh. Zool.-Bot. Ver., Wien. 19:522. 1869; in part.

Ovary wholly superior; carpels conical, at length bottle-shaped, at matu-
rity erect, stout to slender, united up to the middle or merely as high as the
gland; beaks short and stout with thick stigmas, or elongated and some-
what recurved, with small stigmas; fruit a capsule; gland a fleshy collar

surrounding the base of the carpels; filaments clavate; sepals reflexed
(Plate I).

8 MINNESOTA STUDIES IN PLANT SCIENCE

Subsection I. Rotundifoliatae. Leaves orbicular or reniform-cordate,
short and stout, except in one species where they are long and slender;
crenate, long-petioled; carpels united beyond the gland; follicles erect,
greenish or purplish; beaks stout or slender; stigmas globose or somewhat
flattened.

Subsection II. Cuneifoliatae. Leaves cuneate at the base, tapering to
short or elongate petiole-like bases, rounded and deeply incised-toothed at
apex; carpels united merely at the base or to the height of the gland; cap-
sules stout and thick, except in one species where they are elongated; beaks
stigmas thick.

Like the section Arabisa this is one of the most interesting and most
clearly defined of the series. While the characters of the leaves in the
above described subsections are decidedly distinctive, other strong and unmis-
takable characters are to be found in the carpels, the gland, and the mature
capsules. The very young carpels are conical but soon develop into stout
bottle-shaped organs surrounded at the base by the fleshy, but not deep,
gland.

KerY To THE Species oF SECTION HypaTIca

I. Leaves orbicular or reniform-cordate, crenate....... Subsection I. Rotundtfoliatae
A. Inflorescence open
a. Inflorescence elongate, paniculate-cymose..........cecceessscccrces S. arguta
b: Inflorescence'short, .cymose;) delicate. 2.0 .:..c2c+ cc eee ne eee S. aestivalis
Bi Inflorescencel‘contracted) or capitate. ac.. dues tas ccee eek eneieeed S. Nelsoniana
II. Leaves cuneate at the base, deeply incised-toothed at apex.......... Seas eae
STARNALS BS EA) SLi 2 PUN UR SPA A aS EOE ARE Subsection II. Cunetfoliatae
A. Inflorescence open, with slender branches, many-flowered............ S. Lyallié
B. Inflorescence few-flowered, the peduncles stout
a. Follicles jelongates)contracted ss... sisc eetis dele Cece oes eee S. davurica
b= Follicles stout, beaks short, and thick. von coe iaeen aes S. unalaschensis

The species of the Rotundifoliatae are most readily recognized by the
characters of the inflorescence as described in the key, especially in the
case of Saxifraga arguta. In S. aestivalis the inflorescence is very delicate,
in which respect it differs most obviously from S. Nelsoniana. The same
characters hold true in regard to the species of the Cuneifoliatae to distin-
guish S. Lyallii from the other members of the group, which do not differ
to any extent in their inflorescences. S. davurica and S. wnalaschensis can
be distinguished from each other by the mature follicles, as characterized
in the key.

The young carpels are very similar in the Rotundifoliatae, and those of
S. aestivalis and S. Nelsoniana (Plate IV) are difficult to distinguish from
each other except that those of the latter are larger and stouter, with con-
siderably thicker beaks and stigmas. The gland of S. aestivalis is much

JOHNSON: A REVISION OF THE SECTION BORAPHILA 9

thinner, and at first slightly elevated in the form of a projecting ring, while
that of S. Nelsoniana is fleshier and crowds the bases of the stamens
closely. Besides, the stigmas of the latter are somewhat flattened, while
those of the former are large in proportion to the size of the carpels, and
globose. Both species are stoloniferous, with a tendency to branch more
in S. Nelsoniana.

The carpels of S. arguta (Plate IV) have elongated bodies and beaks,
terminating in very small stigmas, and are united up to the middle. Fine
lines run lengthwise along the sides. The species is much less stoloniferous
than its allies. ©

The seeds of the Rotundifoliatae are distinctive in their markings in
comparison with those of the Cunetfoliatae. The distinctions are shown in
Plate IV. In this case the markings have the form of thin, septate, ribbon-
like structures running along the-surface and projecting conspicuously. In
the Cuneifoliatae, on the other hand, the markings are plainly pectinate.

Three types of carpels occur in the Cuneifoliatae, which are made clear
in Plate IV. The stout, erect, purple carpels of S. unalaschensis are un-
mistakable. In S. davurica (Plate IV) they are elongated and not a little
contracted, with much more slender and divergent beaks, and at the same
time somewhat separated. In S. Lyalli (Plate IV) the beaks are greatly
elongated and slender, and the stigmas oblong. The rich, dark-purple color
of the carpels of S. wnalaschensis is absent in these species.

The Rotundifoliatae are species of the high western mountain ranges.
S. arguia is the most widely distributed and also the most abundant,
occurring throughout the length of the Rocky Mountains and also south-
ward into the Sierras. S. aestivalts occurs from the region of Yakutat Bay,
Alaska, to Montana, and south into Oregon. S. Nelsoniana follows the
coast of Alaska and thence inland along the Rocky Mountains to the
Selkirks. Their ranges thus overlap for a considerable distance and, con-
sidered as a whole, constitute a notably continuous distributional belt, which,
when coupled with the close structural similarities, further emphasizes the
close relationships of the group.

The Cuncifoliatae do not show the same continuity of range, as far as
can be determined. S. unalaschensis has been found on the islands of Bering
Sea and in a few scattered localities on the west coast of Alaska to Unalaska.
S. davurica is known to occur along the shores of Kotzebue Sound and
Bering Strait, but has an extended Asiatic range as far as Lake Baikal and
Dauria. The two species are very similar in vegetative characters. S.
Lyallui appears to be an intermediate form; it has the elongate open in-
florescence of S. arguta and its leaves are sometimes rounded distally but
always cuneate at the base; besides, the carpels are greatly elongated and
terminate in slender beaks. These similarities are further emphasized by

10 MINNESOTA STUDIES IN PLANT SCIENCE

the continuity of its range with that of S. arguta, and also of S. Nelsomiana,
but only as far south as the Selkirks. In general it may be said to occupy
the same relative position among the Cuneifoliataé as does S. arguta among
the Rotundifoliatae.

SAXIFRAGA AESTIVALIS F. & M., Ind. Sem. Hort. Petrop. 1:37. 1835.
Micranthes aestivalis (F. & M.) Small, N. Am. Fl. 22:2:147. 1905.

A low delicate plant with a thyrsoid inflorescence of delicate branches
(Plate IV).

Distribution.—From the region of Yakutat Bay, Alaska, including the islands
along the coast, to the Bitter Root Mountains of Idaho and western Montana, and to
the region of Mt. Jefferson in Oregon.

Specimens examined:

ALASKA: at base of Mt. Tabenkoff, June 22, 1892, F. Funston No. 40 (MBG*
84013; USNH 423509); summit of high mountains, Yes Bay, August 30, 1895, Thos.
Howell No. 1622; Kodiak, July 1, 1899, Wm. Trelease No. 4062.

British Cotumpia: Asulkan Valley, Glacier, July 26, 1910, F. K. Butters and
E. W. D. Holway No. 584 (UMH); swampy places, Cowichan-Alberni Trail, Van-
couver Island, June 26, 1906, C. O. Rosendahl No. 1602; stream side, Cameron River
Valley, Vancouver Island, June 27, 1907, C. O. Rosendahl No. 1999 (UMH).

ALpBerTA: Lake Louise, Rocky Mountains, July 19, 1904, Edith M. Farr (MBG
84418) ; head of Lake Louise, July 30, 1904, John Macoun No. 65312 (USNH 529169).

WasHINcToN: Olympic Mountains, Clallam County, August, 1900, A. D. E. Elmer
No. 2640 (USNH 401936, UMH); north fork of Bridge Creek, Okanogan County,
September, 1897, .4. D. E. Elmer No. 716 (MBG 83922); Mt. Rainier, alt. 5,000 ft.,
September 1, 1913, F. K. Butters (UMH); borders of brooks, Goat Mountains, August
6, 1895, O. D. Allen No. 198 (UMH, MBG 84518, USNH 314093) ; Baldy Peak, Che-
halis County, July 24, 1897, F. H. Lamb No. 1342 (MBG 84516); Mt. Rainier, 1906,
Mrs. Carpenter No. 37 (MBG 84508) ; Paradise Park, Mt. Rainier, alt.. 1,500-1,800 m.,
July 22-30, 1905, M. W. Lyon No. 116 (USNH 529458), as S. aestivalis by Engler and
Irmscher; Mt. Rainier, August 15, 1888, C. V. Piper No. 2622 (UMH); Mt. Rainier,
alt. 6,000 ft., August, 189, F. C. Smith (MBG 84510); Stevens Pass, alt. 4,000-6,000
ft., Sandberg and Leiberg (MBG 84124) ; Mt. Paddo, alt. about 2,000 m., July-October,
1896-1905, W. N. Suksdorf No. 2504 (MBG 84129, UMH).

OREGON: at base of Mt. Jefferson, alt. about 6,000 ft., August 25, 1898, F. V.
Coville and I. Applegate No. 1176 (USNH 361993), as S. aestivalis by Engler and
Irmscher.

SAXIFRAGA ARGUTA D. Don, Trans. Linn. Soc. 13 356, 1822.

Saxifraga denudata Nutt., mss. (Unverified. )

Saxifraga punctata acutidentata I-ngler, Verh. Zool.-Bot. Ver., Wien.
19:548. 1869.

Micranthes arguta (Don) Small, N. Am. FI. 22 :2:146. 1905 (Plate IV).

® The abbreviation MBG refers to the herbarium of the Missouri Botanical Garden and USNH
refers to specimen sheets in the United States National Herbarium. In like manner UMH refers
to the herbarium of the University of Minnesota.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 11

Distribution—From William’s Sound, Alaska, to New Mexico and Arizona, east
central California, and extreme western Nevada.

Specimens examined:

WasuIncTon : along alpine rivulets, Olympic Mountains, August, 1895, C. V. Piper
No. 2213 (UMH) ; wet banks, alt. 6,000 ft. Mt. Rainier, August, 1895, C. V. Piper
(UMH) ; along rivulets, alt. 6,500 ft., Mt. Rainier, August, 1895, C. Vv. Piper No. 2025
(UMH) ; Olympic Mountains, Clallam County, August, 1900, 4. D. E. Elmer No. 2639
(UMH, USNH 401962) ; Blue Mountains, July 29, 1897, Robert M. Horner (USNH
318696) ; Yakima region, 1882, R. E. Brandegee No. 304310 (MBG 84525); Mt.
Rainier, alt. 6,500 ft., August 11, 1889, E. C. Smith (MBG 84511); Paradise Park,
Mt. Rainier, July 17, 1907, H. C. Cowles No. 681 (MBG 745452) ; Stevens Pass, alt.
4,000-5,000 ft., August, 1893, Sandberg and Leiberg (UMH, MBG 84104); slopes of
Mt. Stewart, alt. 4,000-5,000 ft., July 24, 1893, Sandberg and Leiberg No. 570 (MBG
84561).

IpAHO: marshy land along streams, alt. 7,000 ft., Silver City, Owyhee County,
July 30, 1910, J. F. MacBride, No. 439 (USNH 542334, MBG 84592, UMH); sunny
brook margins, alt. 5,000 ft., Custer County, July 31, 1910, J. F. MacBride No. 1463
(MBG 84567, UMH) ; stream banks, Trinity Lake region, Elmore County, alt. 8,000
ft., August 29, 1910, J. F. MacBride No. 673 (MBG 84591, UMH); head of Bear
Creek, Bitter Root Forest Reserve, alt. 2,050 ft., August 26, 1897, J. B. Leiberg No.

2941 (USNH 306773) ; near Stevens Peak, alt. 1,600 m., August 4, 1895, J. B. Leiberg
~ No. 1463 (MBG 84079, USNH 250912) ; wet soil by a stream, Victor, July 11, 1901,
E. N. Wilcox and E. Merrill No. 1239 (USNH 581292) ; near Sawtooth, July 30-31,
1896, B. W. Evermann No. 668 (USNH 277512).

OrEGON: near Strawberry Butte, Blue Mountains, alt. 7,000 ft., July 13, 1896,
F. V. Coville No. 567 (USNH 276169) ; granitic, wet soil, source Immaha, alt. 3,000 m.,
August 11, 1908, W. C. Cusick No. 3319 (MBG 84048); 1871, Elihu Hall No. 153
(MBG 84526) ; 1871, Elihu Hall No. 153 (MBG 84251).

CauirorniA: Mt. Brocks, Farwell Gap, alt. 10,300 ft., April-September, 1897,
C. A. Purpus No. 5252 (MBG 84515) ; along cold stream, three miles above Truckee,
Nevada County, July 20, 1903, A, A. Heller No. 7005 (USNH 467405) ; south side of
Echo Lake in granite, moist places, El] Dorado County, August 11, 1916, 4. A. Heller
No. 12547 (USNH 882627); Coyote Meadows, Tulare County, July 30, 1904, C. F.
Baker No. 4311 (MBG 84069). Caulescent; looks different.

MonTANA: Stanton Lake, August 3, 1894, R. S. Williams No. 180 (USNH
288934) ; Spanish Creek, Gallatin County, July 1, 1901, J. Vogel (UMH); Red Lodge,
July 24, 1893, J. N. Rose No. 12 (MBG 84073) ; in brooks Mystic Lake, alt. 6,500 ft.,
July 11, 1905, J. W. Blankinship No. 182 (MBG 84115) ; upper temperate life zone,
spruce belt, alt. 7,000 ft., Alta, July 26, 1909, Marcus E. Jones (MBG 84058) ; Spanish
Bain, Madison Range, alt. 6,000 ft., July 11, 1896, J. H. Flodman No. 515 (MBG
84521) ; Belt Mountains, July 8, 1886, F. W. Anderson, No. 136 (UMH); Basin, July
5, 1892, F. D. Kelsey (UMH) ; Garnet, July 18, 1901, E. W. Scheurer (UMH).

Wyominc: Teton Mountains, alt. 10,000 ft. August 30, 1910, Merrit? Cary No.
602 (USNH 858939) ; Electric Peak, Yellowstone National Park, alt. 8,400 ft., July
26, 1902, EZ. C. S. No. 156 (USNH 488736) ; Teton Pass, August 22, 1900, E. D. Mer-
rill No. 359 (USNH 390769) ; Cold Springs, near Hot Springs, alt. 6,000 ft., 6,200 ft.,
July, 1893, F. H. Burglehaus (UMH); Bald Mountains, August 15, 1892, B. C. Buffum
(UMH); on the banks of cold streams, Medicine Bow Mountains, Albany County,

12 MINNESOTA STUDIES IN PLANT SCIENCE

July 28, 1900, 4. Nelson No. 7759 (UMH, MBG 84509); Union Pass, Wind River
Mountains, August 11, 1894, A, Nelson, No. 843 (MBG 84520); June 28-August 1,
1899, L. H. Pammel (MBG 84514).

UtaH: Clayton Peak, Wasatch Mountains, alt. 9,500 ft., August 12-26, 1903, S. G.
Stokes (MBG 84500); flora of Nebraska and Utah. Expedition of Capt. Jas. H.
Simpson, 1858-59, Henry Engelmann (MBG 84187); slope of Aquarius Plateau, wet
rocks, alt. 8,500 ft., July 29, 1875, L. F. Ward No. 453 (MBG 84507) ; Peterson Canon,
Peterson, common on cold, moist banks, July 19, 1902, L. H. Pammel and R. E. Black-
wood No. 3799 (MBG 84503) ; headwaters of Little Cottonwood Creek. above Alta,
July 10, 1905, P. A. Rydberg No. 6849 (USNH 506353) ; headwaters of White Clay
Creek, in moss attached to roots of trees close to the water. Expedition of Capt. Jas.
H. Simpson, August 23, 1889, Henry Engelmann (MBG 84188); Mt. Ellen, Henry
Mountains, alt. 11,000 ft., July 25, 1894, M@. E. Jones No. 5678 (MBG 84051); Mt.
Ellen, Henry Mountains, alt. 10,000 ft., July 25, 1894, M. E. Jones No. 5478 (MBG
84523).

Nevapa: head of Fall Creek, Ormsby County, alt. 2,460 m., August 6, 1902, C. F.
Baker No. 1434 (USNH 441006) ; Slide Mountain, Washoe County, alt. 7,500 ft., July
11, 1910, A. A. Heller No. 10207 (USNH 509144).

Arizona: Thomas Peak, White Mountains, July 1-15, 1910, L. N. Gooding No.
626 (USNH 661595).

CoLorapo: Virginia Gulch, south slope of Needle Mountains, alt. 11,500 ft., July
14, 1901, Whitman Cross No. 32 (USNH 434312); Ruby, region of the Gunnison
watershed, August 7, 1901, C. F. Baker No. 711 (UMH, USNH 412340) ; Hills, Lari-
mer County, alt. 9,500 ft., July 7-19, 1905, C. F. Baker No. 3179 (UMH); upper La
Plata Canyon, alt. 10,000 ft., July 13, 1898, Baker, Earle, and Tracy (UMH, MBG
84517, MBG 84102) ; Cameron Pass, alt. 10,000 ft., July 29, 1896, C. F. Baker (MBG
84522) ; Dark Canon, alt. 2,800 m., August 6, 1901, F. E., and E. S. Clements No. 392
(MBG 84512, UMH); vicinity of Mount Carbon, along creeks, alt. 3,200 m., August 8,
1910, Ivar Tidestrom No. 3988 (USNH 751491); near Cameron Pass, August, 1892,
G. E. Osterhout (UMH); Tolland, along stream, alt. 9,000 ft., June 29, 1913, L. O.
Overholts No. 10064 (MBG 743695) ; upper Mt. Ricthophen, five miles west of Cameron
Pass, alt. 9,000 ft., July 23, 1894, C. F. Baker, No. 3179 (UMH) ; Front Range, alt. 10,000
ft., July 6, 1896, C. S. Crandall (MBG 84524) ; from the headwaters of Cedar Creek,
and the alpine ridges lying east of Middle Park, 1861, C. C. Parry No. 163 (MBG
84247) ; alpine and subalpine, lat. 39°-41°, 1862, C. C. Parry (MBG 84248) ; Berthoud
Pass, rich pine woods, alt. 8,500 ft., July 15, 1884, C. S. Sheldon (UMH); near
Breckenridge, Mt. Guyot, and Mt. Baldy, alt. 9,700 ft., August 16, 1900, J. P. Anderson
(MBG 718611) wet places, Arg. Pass, July 17, 1886, and Alpine Leadville, July 10,
1886, Wm. Trelease (MBG 84528) ; creek banks, Anita Peak, Routt County, August 3,
1903, L. N. Gooding No. 1756 (MBG 84092); Breckenridge, 1897, Mrs. Wislizenus
(MBG 84573) ; Wheeler expedition of 1873, J. Wolf (MBG 84506) ; Farnham, July 10,
1891, “E. C. S”’ (MBG 84504) ; banks of torrents below Berthoud Pass, alt. 11,000 ft.,
August 10, 1874, G. Engelmann (MBG 84250); Cafion City, Rocky Mountains, 1871,
T. S. Brandegee No. 131 (MBG 84527) ; Graymont, alt. 11,000 ft., July 22, 1886, G. W.
Letterman (MBG 777211) ; Breckenridge, 1887, S. D. Bereman No. 730 (MBG 84577) ;
near Breckenridge, Summit County, alt. 9,700 ft., August 1901, Kenneth W. Mac-
Kenzie No. 205 (MBG 84513) ; Beaver Creek Camp, on Little Beaver, No. 1961, July
15, 1898, ex Herb. St. Agr. Col. (UMH); Boreas Pass to Como, Leadville, Forest

JOHNSON: A REVISION OF THE SECTION BORAPHILA 13

(USNH 768658) ; Rocky Mountain alpine flora, lat. 39°-41°, 1862, Hall and Harbour
Park County, alt. 2,860-3,400 m., September 14, 1915, W. W. Eggleston No. 11968
No. 207 (MBG 8424).

New Mexico: Santa Fe Mountains, F. H. Snow (USNH 216133); along brook
in aspen grove, vicinity of Ute Park, Colfax County, alt. 2,900 m., August 31, 1916,
Paul C. Standley No. 14134 (USNH 896966) ; Mogollon Mountains, on or near the
west fork of the Gila River, Socorro County, alt. 10,000 ft., O. B. Metcalfe No. 515
(USNH 495522, UMH, MBG 8450) ; Ponchuila Creek, Pecos River National Forest,
alt. 8,600 ft., July 4, 1908, Paul C. Standley No. 4187 (USNH 498560) ; Pecos River,
August 8, 1898, G. E. Coghill No. 157 (MBG 84505); Hermit Peak, August, 1883,
F, H. Snow (MBG 776662) ; Rocky Mountains, 1864, C. C. Parry (MBG 84016).

SAXIFRAGA DAvuRICA Willd., Sp. Pl. 2:645. 1799.

Saxifraga punctata Pallas, Reise 3:731. 1776.

Robertsonia davurica Haw., Saxifr. Enum. 56. 1821.

Micranthes davurica (Willd.) Small, N. Am. FI. 22:2:143. 1905.
(Plate IV.)

Distribution—Alaska, from Point Barrow or beyond to Kotzebue Sound and
Bering Strait; in Siberia westward to Lake Baikal and Dauria.

Specimens examined:

ALASKA: Camp 3, ten miles east of Cape Sabine, July 30, 1904, Chester Wash-
burne (USNH 421207, without name, USNH 421208, USNH 421209—all under same
date).

“Habitat in summis alpibus Davuriae sub nive,” Willd. l.c.

“Crescit in montibus Sibiriae summus, nivalibus et in angulo maxime orientali
Asiae,” Pallas 20:47. The specimens enumerated above do not have the long petioles
figured in Pallas’ drawing (Tom. 3, Table P, p. 47, Fig. 2) although the petioles of
specimen USNH 421207 are somewhat longer than those of the other two. No mature
fruit is figured in Pallas’ plate.

SAXIFRAGA Lyatiu Engler, Verh. Zool.-Bot. Ver., Wien. 19:542. 1869.

Saxifraga dahurica Lyall, in Herb. of the Royal Gardens, Kew. (Un-
verified. See Engler, Monogr. Saxifr. 141. 1872).

Saxifraga Lyalhi Var. laxa Engler, Monogr. Saxifr. 142. 1872.

Micranthes Lyallii (Engler) Small, N. Am. Fl. 22:2:143. 1905.

A taller, more slender species than the other members of the Cunetfolt-
atae. The leaves are more rounded and flabellate, having broader teeth,
and longer, narrower petioles. Five-carpelled flowers are not uncommon
(Plate IV).

Distribution—From the Malaspina Glacier and Yakutat Bay, Alaska, to Montana
and Idaho,

The southern limits of the range of this species are rather indefinite.
Sma. (27) gives the distribution as “Alaska and the Bering Sea region to
Alberta, British Columbia, and Montana.” ENGLEr’s (5) distribution reads:
“Auf den rocky mountains unter 49° n.B. um 2,000 m. Fort Colville

14 MINNESOTA STUDIES IN PLANT SCIENCE

(Lyall).” Lyall’s Fort Colville specimen seems doubtful. Piper (21) does
not mention this species as occurring in northeastern Washington.

Specimens examined:

AvasKA: Muir Glacier, Harriman Alaska expedition, July 11, 1899, Wm. Trelease,
No. 4066 (MBG 83986); Muir Glacier, Harriman Alaska expedition, July 12, 1899,
Wm. Trelease, No. 4067 (MBG 83985) ; Alpine Granite Creek, August 12, 1907, H. C.
Cowles, No. 1365 (MBG 746118) ; Sindham Bay, July 28, 1905, Baker and Culbertson,
No, 4952 (MBG 84096).

BritIsH CoLuMBIA: Prairie Hills, Selkirk Mountains, alt. 6,500 ft., July 19, 1904,
C. H, Shaw, and T. B. Snyder, No. 402 (MBG 83913, UMH) ; alpine rivulet, Carbonate
Draw, Selkirk Mountains, July 14, 1904, C. H. Shaw, No. 340 (UMH) ; alpine meadow
in the Big Bend district, about 118° 20’ W. long., 51° 45’ N. lat., alt. 6,000 ft. July
22, 1905, C. H. Shaw, No. 941 (MBG 83914 and UMH); Eagle Mountain Glacier,
August, 1902, C. O. Rosendahl, No. 998 (UMH); Streamsides, Prairie Hills, alt. 5,850
ft., Butters and Holway, No. 222 (UMH); Cougar Valley, Selkirk Mountains, August
10, 1913, Butters and Holway, No. 818 (UMH); Fish Creek Valley, August 10, 1906,
Butters and Holway, No. a25 (UMH).

ALBERTA: Lake Louise, Rocky Mountains, July 19, 1904, Edith. M. Farr, No. 30
(MBG 83912); Mt. Temple, Laggan, Alberta, alt. 7,000 ft., August 26, 1904, Butters,
Holway, and Rosendahl (UMH) ; Laggan, alt. 6,500 ft., July 19, 1910, C. O. Rosendahl,
No. 2489 (UMH); Mt. Temple, Laggan, Alberta, alt. 8,000 ft. September, 1906,
Butters, Holway, and Rosendahl (UMH).

Montana: Lambert Valley, Mission Mountains, upper temperate life zone, alt.
7,000 ft., August 17, 1909, M. E. Jones (MBG 84060).

SAXIFRAGA NELSONIANA D. Don, Trans. Linn. Soc. 13:355. 1822.
Saxifraga punctata L. var. Nelsoniana (Don) Engler, Monogr. Saxiir.

p. 39. 1872.
Micranthes Nelsoniana (Don) Small, N. Am. Fi. 22:2:147. 1905.
A species of usually much stouter habit than S. aestivalis (Plate IV).

Distribution—From Point Barrow along the coast of Alaska to Montana.
Specimens examined: ;

ALASKA: Cooper Island, September 3, 1891, as Saxifraga crassifolia, Jas. M.
Macoun (MBG 83993, USNH 249366) ; Unalaska, July 25, 1891, Jas. M. Macoun, No.
45 (USNH 249364); St. Matthew Island, August 10, 1891, Jas. M. Macoun, No. 46
(USNH 249363); St. Paul Island, July 30, 1891, Jas. M. Macoun, No. 58 (MBG
84015) ; St. Matthew Island, August 19, 1891, Jas. M. Macoun No. 59 (MBG 83994).

The following collections were made by Wm. Trelease on the Harriman Alaska
expedition, 1899; Douglas Island, July 21, 1899, No. 4047 (MBG 84009); Aqua Dulce
River, June 21, 1899, No. 4048 (MBG 84007); Port Wells, June 26, 1899, No. 4049
(MBG 84011) ; Ocra, June 24, 1899, No. 4050 (MBG 83995, 83996; St. Paui Island, July
9, 1899, No. 4051 (MBG 84003) ; St. Lawrence Island, July 13, 1899, No. 4052 (MBG
84008) ; Hall Island, July 14, 1899, No. 4053 (84001) ; Hali Island, July 14, 1899, No.
4054 (MBG 84997) ; Pt. Clarence, July 12, 1899, No. 4055 (MBG 84002) ; Pt. Clarence,
July 12, 1899, No. 4056 (MBG 83998) ; St. Matthew Island, July 15, 1899, No. 4057
(MBG 84000) ; St. Matthew Island, July 14, 1899, No. 4058 (MBG 83999) ; Juneau,
July 25, 1899, No. 4060, “Unusually stout and pubescent” (MBG 84010) ; Juneau, July
25, 1899, No. 4061 (MBG 84006).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 15

Kukak Bay, Harriman Alaska expedition, July 5, 1899, De Alton Saunders, No.

4065 (MBG 84005); Popoff Island, United States Coast Survey, Dall’s exploration,
1871-72, M. W. Harrington (USNH 424500); Shumagin Island, 1871-72, M. W.
Harrington (MBG 84014); Mucklung River, near Nushagak, July 25, 1882,
No. 35 (USNH 423510); St. Paul Island, July 30, 1891, C. H. Merriam (USNH
424511) ; Kukak Bay, Harriman Alaska expedition, July 1 to 5, 1899, F. V. Coville
and H. Kearney, Jr., No. 1663 (USNH 374093); Popoff Island, Shumagin Islands,
July 8 to 19, 1899, Trevor Kincaid (USNH 375087) ; Hubbard Glacier, Disenchant-
ment Bay, Harriman Alaska expedition, June 21, 1899, F. V. Coville and T. H.
Kearney, No. 1081 (USNH 374091) ; Kukak Bay, Harriman Alaska expedition, July 1
to 5, 1899, F. V. Coville and T. H. Kearney, No. 1624 (USNH 374092); Unalaska,
Harriman Alaska expedition July 8, 1899, F. 1’. Coville and T. H. Kearney, No. 1753
(USNH 374095) ; Halibut Cove, Cook Inlet, July 21, 1899, Harriman Alaska expedi-
tion, F. V. Coville and T. H. Kearney, No. 2432 (USNH 374094) ; Sitka, Harriman
Alaska expedition, July 14-17, 1899, F. V. Coville and T. H. Kearney, No. 867 (USNH
374090) ; Port Wills, Prince William Sound, Harriman Alaska expedition, June 26,
1899, F. V. Coville and T. H. Kearney, No. 1265 (USNH 374089); summit of high
mountains, Yes Bay, August 3, 1895, Thos. Howell, No. 1622 (MBG 84012) ; springy
slope, Bennett, July 31, 1907, H. C. Cowles, No. 002 (MBG 745046) ; Windham Bay,
May 31, 1905, as S. Nelsoniana on label by Engler and Irmscher, J. D. Culbertson
(Baker No. 4888), (USNH 666035, MBG 84098) ; St. Lawrence Island, mountain side,
spongy tundras, Gambell, July 15, 1904, as S. Nelsoniana by Engler and Irmscher,
E, P. Campbell (USNH 422067, USNH 422070) ; Herendeen Bay, July 12, 1890, The
Albatross (USNH 423501); Herendeen Bay, July 12, 1890, The Albatross (USNH
423502) ; Kodiak, July 28, 1904, C. V. Piper, No. 4273 (USNH 420725) ; 1873, Dall
(USNH 430725) ; Seal Island, 1875, Capt. Bryant (USNH 423508) ; St. George Island,
Pribiloff Islands, July 21, 1895, F. W. True and D. W. Prentiss, Jr., No. 53 (USNH
231527) ; Camp Retreat, near Hotham Inlet, June 29, 1880, Lt. G. N. Storey (USNH
139475) ; 1878, L. M. Turner, No. 2092 (USNH 423503) ; Atka Island, common in rocky
places, July 1, 1880, L. M. Turner, No. 1220 (USNH 139478) ; Glinka, Copper Island,
Commander Islands, southwest of Bering Sea, August 7, 1895, Leonard Stejneger, No. 53
(USNH 242470); Wrangle Island, Arctic America, 1881, Dr. Ross (USNH 266832) ;
Stepovak Bay, Harriman Alaska expedition, July 8 to 19, 1899, Chas. Palachr (USNH
375479) ; Tundra, Teller Reindeer Station, vicinity of Port Clarence, F. A. Walpole,
No. 1464 (USNH 348570, 378564); Herald Island, 1881, Capt. Hooper (USNH
423507) ; St. Paul Island, Bering Sea, July 2, 1890, Wm. Palmer, No. 423 (USNH
327982) ; Cape Sabine, lat. 68° 53’ N., long. 164° 43’ W., August 12, 1894, James T.
White (USNH 270187) ; Sitka, 1867, expedition to Alaska, U.S. Coast Survey (name
illegible), No. 403 (USNH 423506).

British CotumsBia: Selkirk Range, August 20, 1885, J. Macoun (USNH 219580) ;
north fork of Moose River, July 21, 1911, N. Hollister, No. 52 (USNH 622687, petals

purple).

SAXIFRAGA UNALASCHENSIS Sternb., Rev. Sax. Suppl. 2:9. 1831.

Saxifraga flabellifolia R. Br., mss., ex Torrey and Gray, Fl. N. Am.
1:569. 1838-40.

Saxifraga davurica var. unalaschensis (Sternb.) Engler, Monogr. Saxifr.
148. 1872.

Micranthes flabellifolia (R. Br.) Small, N. Am. FI. 22:2:143. 1905.

16 MINNESOTA STUDIES IN PLANT SCIENCE

The very stout, deep purple follicles of this species are distinctive, differ-
ing markedly in this respect from the elongated, more slender follicles of
S. davurica, besides, the blades of the leaves are more rounded toward the
apex and longer petioled than in that species, but the dentation is similar
(Plate IV).

Distribution—Unalaska, Alaska Peninsula, Pribiloff Islands, Hall Island, St.
Matthew Island, and probably along the Aleutian Islands.

Specimens examined:

ALASKA: Herendeen Bay, The Albatross, 1890, W. H. Hall’s exploration, U.S.
Coast Survey (USNH 423487); Herald Island, 1881, Capt. C. L. Hooper (USNH
423486, 423485) ; Popoff Island, Shumagin Islands, July 8 to 19, Harriman Alaska
expedition, Trevor Kincaid (USNH 375422); St. Paul Island, August 8, 1891, as
“Saxifraga unalaskensis’ C. Hart Merriam (USNH 423484); Cold Bay, July, 1904,
C. V, Piper, No. 4276 (USNH 420728); Hall Island, Harriman Alaska expedition
as S. davurica Pall., July 14, 1899, Wm. Trelease, No. 4070 (MBG 83965) ; St. Matthew
Island, Harriman Alaska expedition, July 15, 1899, Wm. Trelease, No. 4072 (MBG
83966) ; Popoff Island, Harriman Alaska expedition, July 18, 1899, De Alton Saunders,
No. 4069 (MBG 83968) ; St. Paul Island, August, 1891, Jas. M. Macoun (MBG 83969).

Section Calthephyllum, sect. nov.

Gymnopera D. Don, Monogr. Gen. Saxifr. in Trans. Linn. Soc. 13 :343.
1822; in part.

Hydatica (Tausch) Seringe, DC. Prodr. 4:41. 1830; in part.

Micranthes (Haw.) Small, N. Am. Fl. 22:2:146. 1905, as genus; in part.

Boraphila (Engler) Rosendahl, Beibl. Bot. Jahrb. Nr. 83:71. 1905; in
part.

Ovary free; carpels conical, slightly contracted and united at the base,
each surrounded below the middle by a dilated gland; follicles elliptic-
oblong, strongly contracted above and below; styles short and slender;
stigmas small; filaments at first clavate, then greatly elongated and filiform ;
sepals reflexed (Plates I, V).

Monotypic, including only the species described below.

The carpels of this section are at first conical but soon become char-
acteristically bottle-shaped. They are united for a very short distance
from the base. In the mature follicles the united portion is represented
by the conspicuous constriction at the base. The receptacle is strongly con-
tracted at maturity so that the follicles are spread apart considerably. The
hypogyny is complete. The gland forms a broad girdle around each carpel,
extending from the middle to within a short distance of the base. It is only
slightly dilated and seems to disappear completely as the carpels develop.

The section combines certain features of Dermasea and Hydatica. It
resembles Dermasea in the shape of the young carpels and to some extent

JOHNSON: A REVISION OF THE SECTION BORAPHILA 17

in the gland, but differs from that section in being completely hypogynous
and in having the carpels united at the base. In foliar characteristics it
resembles Saxifraga arguta of the section Hydatica.

_ The most conspicuous features of the section are the showy inflorescence
and the remarkably elongated and attenuated filaments which at maturity
attain a length of over six millimeters.

Little is known about the distribution of the species representing the
section. The specimens examined are all from Alaska, namely a few
scattered localities in the lower basin of the Yukon. According to some
writers it extends into British Columbia.

SAXIFRAGA SPICATA D. Don, Trans. Linn. Soc. 13. 1822.
Saxifraga Geum Pursh, Fl. Am. Sept. 311. 1814.
Micranthes spicata (D. Don) Small. N. Am. FI. 22:2:146. 1905.

The only mention of this species made by ENGLER (4) appears in the
following words: “S. spicata Don... .verosimiliter—punctata L.” He makes
no disposition of it in the sections (Plate V).

Distribution.—British Columbia to Alaska.
Specimens examined:

ALasKA: valley of Alatna River, near camp of July 20, about 15 miles above
the mouth, July 20, 1901, W. C. Mendenhall (USNH 377353) ; five miles below Fort
Hamlim, Yukon Valley, 1902, Arthur J. Collier, No. 153 (USNH 379839); Cape
Vancouver, 1891, British Bering Sea Commission collection, Jas. M. Macoun, No.
230 (USNH 249381) ; on the Yukon River, between Andreafski and Anvik, July 16-18,
1889, J. C. Russell (USNH 417561) ; Fort St. Michaels, Norton Sound, 1865-66, H. M.
Bannister (USNH 423413).

The following annotations from the works of previous writers shed light
on the distribution of the species:

“Alaska to British Columbia.” Small, l.c.

“Sledge Island, N. W. Coast, Mr. Nelson, Behring’s Straits, Menzies (v. sp. in
herb. Banks).” Torrey and Gray, Fl. N. Am. 1:567. 1838-490.

Section AULAXIS, Haw., Saxifr. Enum. 47. 1821; as genus.

Robertsonia Haw., Syn. Pl. Succ. 322. 1812, in part; as genus.

Hydatica Tausch, Hort. Canal. Fase. 1. 1823; in part.

Micranthes Torrey and Gray, Fl. N. Am. 1:572. 1838-40; in part.

Boraphila Engler, “Index Criticus,” Verh. Zool.-Bot. Ver., Wien. 19:522.
1869.

Ovary superior; carpels erect, conical, united below the middle, the
upper free portion becoming widely spreading at maturity. Gland, a thick
fleshy girdle extending up over the carpels nearly to the middle, disappearing
with age. Filaments clavate. Sepals reflexed. Seeds elongate, fusiform,

18 MINNESOTA STUDIES IN PLANT SCIENCE

marked with conspicuous longitudinal ridges, each surmounted by an undu-
lating septate keel or edge. Hairs uniseriate. Fruit a capsule (Plates I, V).

Monotypic, including the species described below.

On a preceding page a reference was made to the distinctions on which
HawortH founded the genus Aulaxis—namely on the ground that the
capsules are divided almost into two. He also described the capsules as
united only at the base “capsulae superae alte bipartitae basi solum coalitae.”
In this respect his description was accurate. He was mistaken, however, in
describing the corolla as irregular “corolla irregularis.’ To these distinc-
tions he added an accurate description of the seeds.

HawortH included in the genus a new species, Aulaxis (Saxifraga)
nuda which he had separated from Aulaxris (Saxifraga micranthidtfolia), on
the ground of its glabrous character, but he seems to have left no account
of its origin. SERINGE (26) embodied the species in the section Hydatica,
preceding it with a question mark. In the account of an excursion to the
mountains of North Carolina Dr. Gray (9) writes as follows concerning
this species: “Aulaxis nuda (Haworth, l.c., of unknown origin) appears to
be the more ordinary and more glabrous form of this species” (S. micran-
thidifolia Haw.). The species was not included by Torrey and Gray
(31) in the Flora of North America.

The carpels, the mature fruit, and the gland are very distinctive here.
In addition the hypogyny is complete. At maturity the upper free portions
of the carpels spread widely in a V-shaped fashion. No traces of the gland
could be detected in the mature stage. The young carpels are quite sharply
conical, the beaks soon elongating and terminating in small stigmas. The
gland is at first fleshy and envelopes the carpels to the middle.

The distribution is limited to the Appalachian Mountains, from northern
New Jersey to northeastern Georgia. The altitudinal range ascends to 2,000
feet or more.

SAXIFRAGA MICRANTHIDIFOLIA (Haw.) Britton, in Britton and Brown,
Fl. 2:174. 1897.
Robertsonia micranthidifola (Haw.), Syn. Pl. Succ. 322. 1812.
Aulaxis micranthifolia (Haw.), Saxifr. Enum. 47. 1821.

Saxifraga erosa Pursh, Fl. Am. Sept. 311. 1814.

Saxifraga micranthifolia Steud., Nom. Bot. 736. 1821.

Saxifraga Wolleana Torrey and Gray, Fl. N. Am. 1:569. 1840.
Micranthes micranthidifolia (Haw.), Small, Fl. S.E.U.S. 501. 1903.

The species has often been confused with S. pennsylvanica which it
somewhat resembles in vegetative characters, but the plants can be readily
distinguished by the carpels. There does not appear to be any close affinity
between the two (Plate V).

Til.

—

JOHNSON: A REVISION OF THE SECTION BORAPHILA 19

Pursh’s name, Saxifraga erosa, was for a long time applied to the
species until Brirron (1) restored HAwortuH’s name. Pursu (23) based
his distinctions wholly on inadequate vegetative characters. Dr. Gray (31)
referred Pursh’s species to the section Micranthes Tausch, but later altered
his opinion and referred it to the section Hydatica Tausch, though on what
grounds is not quite clear. At the same time he was led also to modify his
conclusions regarding Saxifraga Wolleana Torrey and Gray, which was
referred by him to the section Hydatica.

Distribution——In the Appalachians from northern New Jersey to northeastern
Georgia.

Specimens examined:

NortH CaroLina: cold streams and wet rocks, Biltmore, April 27, July 3, 1897,
Biltmore Herbarium No. 885 (MBG 84083, UMH); cold streams, Biltmore, May 29,
1896, Biltmore Herbarium No. 885 (UMH, MBG 84318) ; Statesville, MW. E. Hyams.
(UMH); flora of Roan Mountains, alt. 6,000 ft., June 25, 1880, J. S. Chickering, Jr.
(MBG 84316, UMH) ; Grandfather Mountain, Watauga County, June 16, 1891, Small
and Heller, No. 246 (MBG 83939, UMH 1 and 2); mountains, S. B. Buckley (MBG
84321) ; wet places, flank of Roan Mountains, June 20, 1879, Redfield, No. 11722 (MBG
84314) ; mountains of North Carolina, Mitchell County, W. W. Ashe (MBG 84121).

TENNESSEE: along Wolf Creek near the mountains, July, 1896, A. Ruth (MBG
83985); banks of mountain stream, near Thunder Head Mountains, July, 1894, A.
Ruth (UMH); banks of a mountain stream, Wolf Creek, July, 1898, A. Ruth, No.
265 (MBG 83938).

PENNSYLVANIA: no locality, 1841, S. B. Mead (MBG 84217) ; Bethlehem, June 6,

1872, T. C. Porter (MBG 84216); Easton, Redfield herbarium, 1895, Tras! Green
(MBG 84317).

West Vircinta: Pocahontas County, June 15, 1896, W. M. Pollock (MBG
84313).

No state cited: S. B. Buckley, no locality (MBG 84320).
Section Dermasea (Haw.). sect. nov.

Dermasea Haw., Saxifr. Enum. 45. 1821; as genus.
Micranthes D. Don. Trans. Linn. Soc. 13. 1822.
Boraphila Engler, Verh. Zool.-Bot. Ver., Wien. 19:522. 1869.

Ovary partly inferior; receptacle adnate merely to the base of the ovary
or deeper; carpels bottle-shaped, free to the base, connected by the adnate
receptacle and the gland; follicles erect or spreading at maturity; beaks
short or elongated and slender, divergent; stigmas globose or sometimes
oblong. Gland a fleshy ring or collar around the carpels a short distance
above the base of the calyx. Filaments clavate or subulate. Sepals spread-
ing or reflexed. Hairs uniseriate (Plates I, VI).

Haworrts founded the genus Dermasea on Saxifraga (Dermasea Haw.)

nivalis L., and included two new species of questionable status. He regarded

20 MINNESOTA STUDIES IN PLANT SCIENCE

the spreading calyx segments, the “filiform” stamens, and the “semisu-
perior” ovary as the distinctive characters of the genus.

The uniformly bottle-shaped carpels and the peculiar swollen ring-like
gland are the most distinctive characters of this section. The gland lies
a short distance above the base of the carpels in those species in which the
receptacle is very shallow; where the receptacle is deeper the gland lies a
short distance above the base of the calyx. Among some western species
the carpels and the glands are so strikingly alike that it is difficult to dis-
tinguish between them. But in such cases the type of inflorescence and
the form of the filaments serve to throw the species into categories whereby
their individual characteristics are projected and their identification thus
facilitated. The most pronounced deviation from the type appears in
Saxifraga caroliniana where the carpels and their beaks are very slender
and elongated. A deviation from the typical form of the receptacle is
found in S. nivalis where this structure is adnate to the carpels for about
half their length. In general the differences in the depth of the receptacle
are sufficiently pronounced to separate the species into two distinct natural
groups, one possessing a shallow, the other a deep, receptacle. The species
of the first category can be further separated on the characters of the
inflorescence into two nearly equal groups, one with a cymose, the other
with a paniculate, inflorescence. On comparison the species within each
group will appear to be quite closely related, so much so that on foliar
characteristics alone a number of them might easily be confused, as for
instance S. aequidentata, S. Marshall, and S. Allenit, or S. caroliniana, S.
idahoensis, and S. microcarpa.

The smaller categories are based on the characters of the filaments,
scapes, follicles, and petioles. Specific distinctions rest on the characters
of the leaves, sepals, scape, and follicles, where these are of sufficient con-
stancy and distinctiveness.

Turning to the category in which the receptacle is deeper and also
broader, the division is made on the basis of the characters of the filaments
—one with subulate, the other with clavate, filaments. The species of the
first category here also show close affinity ; for example: S. virginiensis and
S. nivalis, which have been frequently confused. The two remaining
species, which comprise the clavate division, are readily distinguished by
the differences in size of the scapes and by the characters of the leaves.

Geographically the species are confined largely to the United States.
Saxifraga nivalis, S. virginiensis, S. tennesseensis, S. Careyana, and S. caro-
liniana are species of the eastern United States. The two first named are
not clearly delimited as to their ranges. S. nivalis is circumpolar, being a
plant particularly of the Arctic regions. It is doubtful whether this species
actually occurs in the United States. No American specimens of it have

JOHNSON: A REVISION OF THE SECTION BORAPHILA 21

been available for study. The range of S. virginiensis is discussed more
at length in connection with that species. S. Careyana and S. caroliniana
are restricted to a small area in the southern Appalachians, from Tennessee
to Georgia, while S. tennesseensis is found only in a small locality in the
vicinity of Knoxville, Tennessee.

Among the western species the most widely distributed is S. saximontana,
which follows the Rocky Mountains from British Columbia to the Yellow-
stone Park, with outposts in the Black Hills of South Dakota and in the
Wallowa Mountains of eastern Oregon. The remaining species are known
only by the type specimen or by scanty specimens from scattering localities,
on which no definite statements as to their distribution can be based.

It is of interest to note that the western species may all be conveniently
inscribed within a triangle having its apex in the Black Hills and its base
coincident with a line drawn from north of Vancouver Island to the north-
east corner of California. The great bulk of the species will be included
within the lower (western) half of this triangle. When we add to this popu-
lation the species of other sections that are also natives of this region we
shall have, undoubtedly. one of the great distribution centers of the genus
Saxifraga.

Key TO THE SPECIES

I. Receptacle shallow, contracted
A. Inflorescence cymose
1, Filaments subulate
a. Scapes glabrous, at least above

(1) Leaves rusty-tomentose beneath........0.0 00.0050. e cece nseees S. rufidula
(2) Leaves glabrous, shallowly dentate towards apex.......... S. laevicaulis
b. Scapes pubescent, leaves crerate-dentate................2005. S. aequidentata
2. Filaments clavate
a. Leaves deeply and finely crenate; inflorescence branches filiform........ aie
Hit AS as Sk eRe ae CEE Ne AULT LUA NAD SUA he A A a REL EN De) EOL MAN AACE AG Madd R
b. Leaves shallowly crenate, inflorescence stout
PRMITMOGESCENCEMPANICUALes Ue @cusle eiee avataal sel lele evalele/a\eile vi oyei sje aaloteren alles a S. Allenii
1. Filaments subulate
B15 SADR) eS LS La SEN OIE ee NG ese VM ERIN HUoIRIAT ea S. Careyana
MP SEM AIS ISPLEA MIS ire Nee Namie sae euheie sional © oviate)a) sbebaeyalel shajatehe) si S. tennesseensis
2. Filaments clavate
aeollicles) slender Clongater sis. osc... cist tee ee + as) a slelejale w etabelclel S. caroliniana

b. Follicles stout
(1) Petioles much exceeding the blades

(a) Leaves cuneate at the base............. 00. ee veces ee cess S. idahoensts

(b) Leaves truncate or subcordate at the base...............-.. S. petiolata
(2) Petioles equalling or shorter than the blades

(a), Beavesi sintiate Or) Tepand sc .'<\s, .)e)s/e\e\e/elslejelele ste)-\ers|eleleleeiebel ofele a oictls S. Halli

(b) Leaves crenate-dentate toward the apex..............+.. S. microcarpa

22 MINNESOTA STUDIES IN PLANT SCIENCE

II. Receptacle deeper and broader
A. Filaments subulate; sepals spreading
1. Receptacle adnate to the basal portion of the carpels
a. Young carpels distinctly bottle-shaped; gland reduced....5. klickitatensis
b. Young carpels conical above the receptacle; mature carpels pyramidal
above the’ fléshy ;2land 2.5.25 ws cle we gee ecw eans cisteie es ee S. virginiensis
2. Receptacle deep, adnate to the lower half of the carpels, or nearly so.......
Serta kak) Ue ae ge lars ater eae tates PRIM ARS SIFU 0 Vata kat ae SO ee S. nivalis?
B. Filaments clavate; sepals spreading
1. Scapes tall; leaves serrate to sinuate-dentate..............6. S. saximontana
2. Scapes much reduced; leaves crenate, rusty-tomentose beneath..............
BE HANNE a NIAAA cy SOC UEC OOOO AC CO TOC BUDO REED bob D0 05 S. occidentalis

SAXIFRAGA ALLENIT (Small), n. comb.
Micranthes Allen Small, N. Am. FI]. 22:2:144. 1905.

From all appearances a good species, but the specimens are in the early
flowering stage so that the characteristics of the mature fruit and seeds
can not be determined and compared with those specimens referred to S.
saximontana (q.v.), and which have been confused with it. The relatively
large flowers, rather compact cymules, crenately toothed and lanuginous
leaves are distinctive. In two of the plants on the sheet the inflorescences
are conspicuously thyrsoid (Plate VII).

Distribution —Washington.

Specimens examined:

BritisH CotumBIA: alpine rivulet, small peak above timber line, about 118° 20’ W.
long., 51° 45’ N. fat., alt. 7,500 ft., July 25, 1905, C. H. Shaw, No. 1001 (UMH); Car-
bonate Draw, Selkirk Mountains, alt. 6,000 ft., July 13, 1904, C. H. Shaw, No. 302
(MBG 84110) ; gorge of Carbonate Draw, alt. 6,500 ft., July 13, 1904, C. H. Shaw, No.
257 (MBG 84089).

WASHINGTON: near Whistler Pass, east fork of Pasayton River, Okanogan Forest,
Okanogan County, alt. 2,340 m., August 4, 1916, Eggleston, No. 13416 (USNH
886307) ; Goat Mountains, alt. 5,500 ft., O. D. Allen, No. 242 (MBG 84432). Type!

SAXIFRAGA CAREYANA A. Gray, London Journ. Bot. 2:116. 1843.

Micranthes Careyana (Gray) Small, Fl. S.E.U.S. 501. 1903. —

Two specimens from the herbarium of the Missouri Botanical Garden,
No. 84203, bearing the legend ‘‘Saxifraga Careyana Gr. Coll. A. Gray, C. S.
Sargent, J. H. Redfield, and Wm. M. Canby, Roan Mt., June 23. Journey
to North Carolina, 1879” (merely a poor scape in fruit), and No. 84286,
with the legend “Saxifraga Careyana, Gray! Grandfather Mt., N.C. Col-
lected by Dr. Gray” afford the only authentic material at hand. The latter
consists, however, of the bases of two separate plants with leaves and a
single detached scape in poor condition. While the scape and inflorescence
appear characteristic of S. Careyanna, the leaves, on the other hand, are

* The detailed treatment of this species is deferred for a future paper.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 23

considerably coarser and thicker than in the other specimens cited above,
and more suggestive of S. caroliniana. Two other specimens from the same
herbarium, namely MBG 84204, “legit S. B. Buckley 1842,” from North
Carolina, and MBG 84287, “legit S. B. Buckley, . . . in montibus Carolinae
et Georgiae,” are referred by the collector to this species, but they are in too
poor condition for use. The species was recognized by RosENDAHL (25,
p. 66) and by ENcLER (5, p. 136) who, however, also referred Saxifrage
caroliniana Torrey and Gray to this species, as a variety.

Saxifraga Careyana appears to be a much smaller and more delicate
plant than either S. caroliniana or S. tennesseensis. The scape is very slen-
der and also low, and the branches of the inflorescence filiform. Harscu-
BERGER (15, p. 496), depicting the vegetation of the southern Appalachians,
found the plant growing under overhanging cliffs, and describes it thus: “the
delicate little Saxifraga Careyana.”

It is difficult to determine from the material whether the calyx-seg-
ments are consistently spreading or reflexed. In the flowers they appear
spreading, although in some cases apparently reflexed. The same difficulty
appears when in fruit, but here in most cases they are reflexed (Plate VII).

Distribution—“Mountains of North Carolina” according to SMALL (27). The

specimens examined that can with some degree of certainty be referred to this species,
are all from Roan Mountain, North Carolina, as given below.

Specimens examined :

Nortu Caro.ina: July 5, 1880, J. W. Chickering, Jr. (MBG 84284) ; alt. 6,000 ft.,
July 5, 1880, J. W. Chickering, Jr. (UMH); in shade of the overhanging rocks, June
21, 1879, J. H. Redfield, No. 11721 (MBG 8428S).

SAXIFRAGA CAROLINIANA A. Gray, Mem. Amer. Acad. 3:39. 1846.

Saxtfraga Careyana var. caroliniana Torrey and Gray, Mem. Amer.
Acad. 3:39; Engler, Monogr. Saxifr. 137. 1872.

Saxifraga Grayana Britton, Mem. Torr. Bot. Club 5:178. 1894.

Micranthes Grayana (Britton) Small, Fl. S.E.U.S. 501. 1903.

Micranthes caroliniana (A. Gray) Small, N. Am. Fl. 22:2:146. 1905.

A tall and graceful species with carpels and other floral characteristics
very similar to those of S. Careyana, except that it has clavate filaments.
The beaks of the carpels are characteristically long and slender. In foliar
characters it is strikingly similar to the western Saxifraga idahoensis Piper.
The following quotation from Smatt and Vait (29), in regard to this
species, may be of interest: “This interesting plant was first collected by
Dr. Asa Gray on Grandfather Mountain, North Carolina, growing with
S. Careyana, both plants being published under that name in Am. Journ.
Sci. XJIT, in 1842. Dr. Gray wrote that the difference between the two
plants was not detected till they bloomed in Cambridge the following spring,

24 MINNESOTA STUDIES IN PLANT SCIENCE

when they displayed characters that remained constant in cultivation.”
From descriptions of its habitat it appears to be a calciphobe species.

Dr. Gray’s (10, p. 39) original account reads: “Living plants of this
species were gathered by myself in the Allegheny Mountains of North
Carolina, and probably of Virginia, also, in the autumn of 1843, along with
those of S. Careyana, which this species so nearly resembles that the differ-
ence was not detected until both came into flower the ensuing spring.” Dr.
Gray noted and distinguished the two species on the characters of their
filaments (Plate VIII).

Distribution —Mountains of Virginia and North Carolina.
Specimens examined:

WEsT VIRGINIA: about Chatham Hill Gap, on Walker Mountain, Smyth County,
alt. 3,000 ft., June 13, 1893, John K. Small (MBG 84288, 84292). (The above two col-
lections are both labeled Saxtfraga caroliniana). UMH, as S. Grayana!

NortH CAROLINA: Negro Mountain, June 27, 1879, Gray, Sargent, Redfield, No.
11791 (Redfield herbarium), (MBG 84291).

Saxifraga Hallii, sp. nov.

Ovarium parve inferiore, carpidia conica aliquam elongata erecta dorso
plus minusve 1-costato superne in stylos gradatim contracta base constricta
glandula crassa convolutaque cincta, stigmatibus globosis, filamentis clavatis
ad maturitam evidente subulatis calyce conspicue longioribus, sepalis maxime
reflexis oblongis vel ovato-oblongis 1.5-3 mm. longis, dorso margineque
sparse pubescente, petalis ellipticis vel ovato-ellipticis apice rotundatis base
plus minusve contractis, folia basalia 7 cm. longa ovata vel elliptico-ovato
aliquam crassa inferne pallida glabra vel margine minime ciliata sinuato-
dentata in petiolum alatum cuneate contracta petiolo lamina longiore vel
aequale, scapus erectus 40 cm. altus sparse glandulo-pubescentis.

Ovary slightly inferior; carpels conical, somewhat elongate, erect, with
a more or less evident dorsal rib, gradually contracted above into the styles,
constricted at the base, and surrounded by a thick, convolute gland; stigmas
globose; filaments clavate, at maturity evidently sublate, conspicuously
longer than the calyx; sepals strongly reflexed, oblong or ovate-oblong,
1.5-3 mm. long, sparsely pubescent on back and margin; petals elliptic or
ovate-elliptic, rounded at the apex, more or less contracted at the base;
leaves basal, 7 cm. long, ovate or elliptic-ovate, rather thick, pale beneath,
glabrous or slightly ciliate on the margins, sinuate-dentate, cuneately con-
tracted into a broadened petiole, the blade exceeding or equalling the petiole;
scape erect, 40 cm. high, sparsely glandular pubescent.

The leaves of this species are very similar to those of “Saxifraga inte-
grifolia Hook.,” but the floral structures are all typical of the section Der-
masea. The bracts possess tufts of long hairs on the margins, or are

JOHNSON: A REVISION OF THE SECTION BORAPHILA 25

eciliate by the falling of the hairs. The gland becomes elevated by the
growth of the carpels and takes the form of a wrinkled girdle. The young
carpels taper gradually and somewhat pyramidally upwards into the rather
slender elongated beaks (Plate IX).

Specimens examined:

Elihu Hall 151, “Oregon” . . . “ann. 1871” (Type MBG 84380). Plantae ore-
gonensis. Legend: “Saxrifraga virginiensis Michx., a very tall form.’ MBG 84380.
(On a sheet from the Redfield herbarium, bearing also two specimens of Saxifraga
californica. The type specimen of this species is designated “a’’—the leaf-hand speci-
men).

SAXIFRAGA IDAHOENSIS Piper, Bull. Torr. Bot. Club 27 :394. 1900.

Micranthes saximontana (E. Nelson) Small, N. Am. FI. 22:2:145. 1905.

The outstanding characteristics of this species are the very shallow
receptacle and consequent slightly inferior ovary, and its habit. In foliar
characters it is strongly suggestive of Saxifraga caroliniana. It is a grace-
ful plant with an open and diffuse inflorescence, lacking the rigid character-
istics of the plants referred to as S. saximontana, whose range is contiguous.
In some respects it resembles S. Allenit.

Its limited range may probably be extended by further exploration,
since topographic and climatic conditions similar to those of its locality
extend over a wide region (Plate VIII). |

Specimens examined:

IpAHO: moist hillsides, island in Clearwater River, above Lewiston, May 2, 1892,
Sandberg, MacDougal, and Heller, No. 93; Palouse country and about Lake Coeur
d’Alene, May, 1892, J. H. Sandberg; moist hillsides, Lewiston, May, 1893, J. H. Sand-
berg.

Saxifraga klickitatensis sp. nov.

Receptaculum ovarii parte inferiore adnatum flabelliforme, carpidia conica
superne conspicue contracta, base glandula dilatata cincta stylis attenuatis,
folliculi purpurei stylis conspicue attenuatis divergentibus, stigmatibus parvis ;
filamenta subulata, plana, superne contracta; sepala reflexa; petala elliptica
1-nervia; pilorum cellulae 2-3 cylindricae in seriem unam dispositae; folia
basalia ovata serrato- vel crenato-dentata infra lanuginoso-pubescentia.

Carpels bottle-shaped, surrounded at the base or slightly above by a
dilated gland; follicles purple, inflated at maturity, contracted above into
slender elongated, strongly spreading to somewhat recurving beaks. Fila-
ments subulate, flat, tapering above to a sharp point. Sepals 1-1.5 mm.,
green, becoming reflexed; lateral nerves usually 2, wanting in smaller seg-
ments, reaching beyond the middle or coalescing with mid-nerve below apex.
Petals white, 1.2-2.7 mm. elliptic to elliptic-oblong, 1-nerved. Hairs of

26 MINNESOTA STUDIES IN PLANT SCIENCE

2-3 cylindrical cells, their glands oblong to orbicular, of 2-4 superposed
cells. Leaves ovate to oblong-ovate, 2-6 cm., glabrous above, lanuginous
beneath, shallowly serrate- to crenate-dentate, tapering to a narrow petiole-
like base about equalling or longer than the blades; ciliolate. Scapes about
15 cm. high, sparsely short glandular pubescent, branching above into a
widely spreading corymbose inflorescence, the branches rigid and slender,
arising from the axils of rather small glabrous, lanceolate bracts which are
broadest below the middle, the upper greatly reduced. Perennial from a
stout fibrous-rooted caudex which is sometimes branched.

Aspect of S. aequidentata, but differs from that species in having a
deeper receptacle, bright purple, slender-beaked follicles, shallower toothed
leaves, small petals, and orbicular rather than elongate-oblong glands on the
hairs (Plate IX).

Distribution—Known only from the type locality.
Specimens examined:

WaASHINGTON: damp rocks, west Klickitat County, April 9, May, 1883, W. N.
Suksdorf (USNH 45806); Type! Right-hand specimen designated “a.” Originally
identified as S. nivalis L. and referred to by Engler and Irmscher to S. aequidentata
Small.

Saxifraga laevicaulis sp. nov.

Receptaculum ovarii basi adnatum, carpidia conica supra basin latis-
sima, superne stylis attenuatis maxime contracta supra basin glandula dila-
tata cincta dorsale uno-costata; filamenta subulata; sepala patentia; parva
petalis multo breviora; petala elliptico-oblonga, 2.5-3.7 mm.; folia basalia
ovata repando- vel sinuato-dentata, petiolis brevibus; scapus glaberrimus
gracilis purpureus ramis corymbosis paucifloribus.

Carpels bottle-shaped, broadest at the gland, strongly contracted above
into slender beaks, stigmas subglobose. Gland an irregular lobed collar
above the base of the carpels; filaments subulate, rather slender, sharp-
pointed ; anthers short and broad to elongate, yellowish. Sepals 1.3-1.7 mm.
oblong to ovate, rounded at apex, mucronulate by the projecting mid-nerve,
dull green, thin, slightly united at the base, lateral nerves 2, reaching to
middle or uniting with mid-nerve at apex. Petals 2.5-3.7 mm., much ex-
ceeding the sepals, obtuse to rounded at apex, oblong to elliptic or some-
times ovate, sometimes slightly notched; mid-nerve with 2-3 alternate
branches arising at or below the middle, these variously branched and their
branches more or less anastomosing. Receptacle very shallow, adnate
merely to the base of the carpels. Leaves 1-4.5 cm. long, typically ovate,
the smaller sometimes elliptic, thin, green, glabrous, acute to obtuse, shal-
lowly dentate or sinuately dentate or repand, especially towards the apex,
contracted into slender petioles which mostly exceed the blades. Scapes

JOHNSON: A REVISION OF THE SECTION BORAPHILA 27

8-14 cm. high, very slender to delicate, purplish, glabrous except for an
occasional gland-tipped hair, solitary, erect or assurgent, corymbosely
branched above or from the middle into a few-branched, few-towered in-
florescence, the branches or peduncles very slender to delicate, bearing 1-3
flowers; glabrous. Perennial from a short, slender, fibrous-rooted caudex.

A species of very slender to delicate habit, with glabrous scapes which
are quite purplish in color. The inflorescence is also very delicate. In the
specimens examined the carpels are all very broad at the gland, strongly con-
tracted upwards to the beaks, and conspicuously bottle-shaped (Plate VI).

Distribution—Known only from the type locality.

/ Specimens examined:
Orecon: Mt. Grayback, June 15, 1904, C. V. Piper, No. 5061 (UMH). Type!

SAXIFRAGA AEQUIDENTATA (Small) N. Comb.
Micranthes aequidentata Small. N. Am. Fl. 22:2:145. 1905.

Ovary slightly inferior, the receptacle very shallow and flattened. Fol-
licles short to longer, stout, somewhat spreading, occasionally 3 together ;
beaks very short, spreading. Stigmas at length elongate, papillate, becoming
strongly deflexed-curved. Young carpels erect, broadened at base, with short
stout beaks. Gland a dilated collar-like ring at the very base of the carpels.
Filaments subulate, 2-3.5 mm.; anthers purple on the margins. Sepals broad,
ovate, oblong-ovate to deltoid, varying considerably, 1.5-2.5 mm., acute to
obtuse, free to base or nearly so, pubescent on the back; lateral nerves 2, of
various lengths, often coalescing with midnerve at middle or below apex;
spreading at first, later reflexed. Petals large, 3-4 mm., white, broadly ovate
to elliptic-oblong or obovate, scarcely clawed at the base; branches of the mid-
nerve 2, simple or variously branched, sometimes coalescing with midnerve
at or below apex. Hairs short, the glands elongate oblong to ovovoid, or glo-
bose, multicellular, the former consisting of a row of rounded cells. Seeds
fusiform, with longitudinal ridges of low moniliform or scarcely tuberculate
cells of “teeth.” Leaves basal, short, 7 cm. and less, ovate to elliptic, some-
times triangular-ovate or deltoid, rounded at apex, uniformly and rather
finely crenate-dentate, copiously brownish pubescent to tomentose beneath,
contracted into somewhat elongate, flattened, petiole-like bases, commonly
longer than the blades; leaves usually crowded among the remains of the
earlier leaves. Scapes erect or assurgent, 20 cm. tall and less, single or
several together, scantily glandular-pubescent, branching at the top into a
widely spreading corymbose many-flowered inflorescence with open cymules
and the branches arising from small linear-oblong or lanceolate bracts,
copiously pubescent on the back, the upper much reduced. Perennial from
a short stout fibrous-rooted caudex.

28 MINNESOTA STUDIES IN PLANT SCIENCE

Distribution: “Washington,” according to Small, Lc.
Specimens examined:

WASHINGTON: on wet rocks, mountains near Lower Cascades. W.N. Suksdorf,
No. 967, type, referred to Saxifraga nivalis L. Watson, in herb.; Skamania County,
May 29, 1886, W. N. Suksdorf (UMH).

Orecon: Oregon City, April 1885, Thomas Howell (USNH 45801, and UMH),
referred to Saxifraga refiexa Hook.

The filaments of the type collection of Micranthes aequidentata Small
(Suksdorf No. 967) do not agree with Sma.u’s description.

SAXIFRAGA MarRSHALLII Greene, Pitt. 1:159. 1887-89.
Micranthes Marshall (Greene) Small, N. Am. Fl. 22:2:145. 1905.

Very similar to Saxifraga aequidentata (Small) Engl. and Irmsch, but
differs in its smaller size and in having clavate filaments and reflexed sepals.
Greene’s specimen was from Hoopa Valley, California (Plate VII).

Distribution —Oregon to California (Small, l.c.).

Specimens examined: :

CALIFORNIA: ridges and meadows near Marble Mountains, Siskiyou County, alt.
7,000 ft., June, 1901, H. P. Chandler, No. 1613 (UMH, MBG 83916).

Saxifraga microcarpa sp. nov.

Receptaculum ovarii base adnatum, carpidia 2, nonnunquam 3, parva supra
basin glandula dilatata cincta conica et superne multo contracta; folliculi
parvi medio dilatati stylis brevibus stigmatibus parvis; filamenta latiss.me
clavata antheris luteis; sepala oblongo-ovata ciliolata base conspicue coalita
reflexa; petala elliptico-oblonga vel ovata base maxime contracta; folia
basalia ovata vel elliptica infra sparse pubescentia marginibus purpureis
grande dentatis vel serrato-dentatis; scapi erecti superne paniculato-ramosi
ramis patentibus adscendentibus rigidis floribus parvis paucis vel multis.

Receptacle adnate to the base of the ovary; carpels 2, sometimes 3,
small, surrounded above the base by a dilated gland, conical, and strongly
contracted above; follicles small, dilated at the middle; styles short; stigmas
small; filaments very broadly clavate, anthers yellow; sepals oblong-ovate,
ciliolate, conspicuously united at the base, reflexed; petals elliptic-oblong or
ovate, strongly contracted into a clawlike base; leaves basal, ovate or ellip-
tic, sparsely pubescent beneath, margins, purplish, coarsely dentate or serrate-
dentate ; scapes erect, paniculately branched above, the branches spreading,
ascending, rigid; flowers small, few to many; bracts linear-lanceolate, small,
cilioate ; leaves bluish-green and brownish pubescent beneath; perennial from
a rather slender, fibrous-rooted caudex (Plate IX).

Distribution—Known only from the type locality.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 29

Specimens examined:

Montana: Herbarium of the University of Montana, two sheets, for convenience
designated “a” and “b.” Sheet “a” type, determined by P. A. Rydberg as S. idahoensis;
Missoula, M. F. Elrod, No. 98, determined as S. reflexa montana by J. W. Blankinship
with the note: “Has leaves of S. reflexa Hook. and inflorescence of S. Nutkana Moc.
Hither an intergrade or a new species.” .

SAXIFRAGA OCCIDENTALIS Wats., Proc. Am. Acad. 23:264. 1888.

Micranthes occidentalis (Wats.). Small, N. Am. Fl. 22:2:144. 1905.

Distribution.—The exact range of this species is not known at present. According
to SMALL (l.c.) it is distributed from “Alberta to Montana,” a delimitation which is
quite at variance with that given by him in Bull. Torr. Bot. Club, 23:362, 1896, which
reads, “apparently confined to Vancouver island and the mainland in the immediate
vicinity.” No published explanation of the difference is known to the writer. Dr.
WartTSoN’s own account reads: “From the Rocky Mountains of British America
(Drummond) to British Columbia and Vancouver Island (Lyall, Macoun). Oregon
(Cusick, Henderson, Howell), and the northern Sierra Nevada (Chico, Mrs. J. Bid-
well; Gray).” The Cusick specimen is very likely the one listed above. The type
locality, according to SMALL (l.c.), is the “Rocky Mountains.” ,

It is to be noted in this connection that SMALL’s description of this
species in Bull. Torr. Bot. Club, 23:362 is conspicuously at variance with
the description he has given in North American Flora, 22:2:145, 1905, par-
ticularly in respect to the leaves. It seems that the descriptions must have
been based on two rather different plants.

S. occidentalis Wats. is a small species with finely crenate leaves, which
_are also commonly deep vinous red and more or less reddish tomentose
beneath. The scapes are sometimes stout, with a few-flowered, cymose in-
florescence, and stout carpels. It is easily mistaken for S. rufidula (Small),
except for its clavate filaments and obviously deeper receptacle. Elmer’s
specimen (2646) has been erroneously referred to S. rufidula by ENGLER
and IRMSCHER on their label on the herbarium sheet (Plate X).

Specimens examined:

OrEGoN: Powder River Mountains, as S. nivalis; W. C. Cusick (UMH).

WasHINGTON: Mt. Rainier, “On rock cliffs,” alt. 7,000 ft., August, 1895, as S.

nivalis, C. V. Piper, No. 2034 (UMH); Olympic Mountains, Clallam County, July,
1900, Elmer, No. 2646 (USNH 401969), as S. nivalis.

Saxifraga petiolata sp. nov.

Receptaculum ovarii base adnatum, carpidia 2, nonnunquam 3, supra
basin glandula dilatata cincta dorso conspicue uni-costato, stylis attenuatis
elongatis, stigmatibus parvis oblongis, filamenta clavata inferne angustata,
antheris purpureis, sepala oblongo-ovata reflexa base conspicue coalita 2-2.5
mm., petala ovata vel elliptica obscure retusa ad basin maxime contracta
3-3.5 mm., semina ellipitico-ovata superne latissima et ad apicem tuberculata
longitudinaliter sulcata, folia basalia longe petiolata ovata vel oblongo-ovata
subcordata marginibus sinuato- vel crenato-dentatis, scapus altus erectus

30 MINNESOTA STUDIES IN PLANT SCIENCE

graciliusculus sparse glanduloso-pubescentis ramis paniculatis diffusis paten-
tibus multifloribus bracteis linearibus vel lanceolatis; pilorum cellulae in
seriem unam dispositae glandulis multicellularibus globosis vel oblongis
saepe purpureis.

Receptacle adnate to the base of the ovary, carpels 2, sometimes 3, sur-
rounded above the base by a dilated gland, and with a conspicuous dorsal
rib ; stigmas small, oblong ; filaments clavate, narrow below; anthers purplish;
sepals oblong-ovate, reflexed, conspicuously united at the base, 2-2.5 mm.;
petals ovate or elliptic, obscurely retuse, greatly contracted at the base,
3-3.5 mm., seeds elliptic-ovate, broadest above and tuberculate towards the
apex, longitudinally grooved; leaves basal, long petroled, ovate or oblong-
ovate, subcordate, margins sinuate or crenate-dentate ; scape tall, erect, some-
what slender, sparsely glandular-pubescent, paniculately branched, the
branches diffuse and spreading and many-flowered; bracts linear or ianceo-
late; hairs uniseriate, glands globose or oblong, multicellular, often purple.

Scapes single or several; perennial from a short, stout, simple, or
branched caudex.

Prominent characteristics: strongly contracted receptacle, inflated folli-
cles, greatly elongated petioles, subcordate leaves, costate carpels, and large
globose glands of hairs. It is also a tall plant with an open diffuse inflo-
rescence of rather few branches (Plate VII).

Distribution—Known only from the type locality.

Specimens examined:

OREGON: on wet banks, Woodville, May, 1889, Thomas Howell (UMH, MBG
83917). Type!

SAXIFRAGA RUFIDULA (Small) n. comb.
Micranthes rufidula Small, N. Am. FI., 22:2:140. 1905.

A small species very similar to S. occidentalis (q.v.). The shallow
receptacle gives the appearance of a hypogynous condition. Follicles rela-
tively large (Plate VI).

Distribution—Appears to be limited to Vancouver Island, adjacent British Colum-
bia, and the Olympic mountain region of Washington.

Specimens examined:

BritisH CotumBta: Mt. Finlayson, Vancouver Island, May 17, 1887 (type col-
lection), J. Macoun; Mt. Arrowsmith, July 18, 1887, J. Macoun; Mt. Arrowsmith,
Vancouver Island, alt. 3,000-5,000 ft., June 29, 1906, C. O. Rosendahl, No. 1631; slopes
of Mt. Arrowsmith, Vancouver Island, alt. 3,000-5,000 ft., June 28, 1907, C. O. Rosen-
dahl, No. 2030.

WASHINGTON: on wet cliffs, Olympic Mountains, alt. 3,000-5,000 ft., August, 1895,
C. V. Piper, No. 2212; Mt. Angeles, on the highest elevations, 1909, Webster; Mt.
Baldy, alt. 4,000 ft., July 7, 1902, H. S. Conrad, No. 279. ;

JOHNSON: A REVISION OF THE SECTION BORAPHILA 31

SAXIFRAGA SAXIMONTANA E. Nelson, Erythea 7:168. 1899.

Micranthes saximontana (E. Nelson) Small, N. Am. Fl. 22:2:145.
1905.

(Plate X.)

Distribution.—Rocky Mountains of British Columbia and Alberta to the Yellow-
stone National Park, with outposts in the Wallowa Mountains of eastern Oregon, and
in the Black Hills of South Dakota.

Specimens examined:

Wyominc: under a cliff, Yancey’s Yellowstone National Park, July 17, 1899, A.
and E. Nelson, No. 5917 (type) (MBG 84383, USNH 365280, UMH) ; rich marly hills
in Jackson’s Hole, Snake Fork, expedition of Capt. W. F. Reynolds, U.S.A., 1859-60,
alt. 6,000 ft., June 12, 1860, Dr. F. V. Hayden (MBG 8419, MBG 84140).

IDAHO: granite ledges, Kootenai County, June, 1891, J. B. Letberg, No. &
(UMH) ; mountains, Kootenai County, July, 1889, J. H. Sandberg (UMH) ; moun-
tains, Kootenai County, July, 1889, J. H. Sandberg (MBG 84278) ; Little North Fork
Basin, alt, 1,000 m., September 4, 1895, J. B. Leiberg, No. 1640 (MBG 84600).

Orecon: Little Fork, Wallowa Mountains, July 12, 1897, E. P. Sheldon, No. a
(USNH 528321), referred to S. aequidentata by ENGLER and IRMSCHER,

Montana: Spanish Creek, Gallatin County, J. Vogel (USNH 416291), 1901,
referred to §. Allentt (Small) Engl. and Irmsch. by Engler and Irmscher.

SoutH Daxota: Black Hills, near Fort Meade, May 6, 1887, W. H. Forwood,
No. 121, referred to S. Allenit (Small) Engl. and Irmsch. by EncLer and IRMSCHER
(USNH 317662).

The following specimens from the United States National Herbarium
are considerably stouter, but in other respects are similar to the above. They
have a heavier and, in earlier stages, a more thyrsoid inflorescence, but
otherwise show the variations in the margins of the leaves as the above.
In floral characters they are identical.

ALBERTA: vicinity of Banff, alt. 5,300-7,000 ft., July 7, August 4, 1899, W. S.
McCalla, No. 2290 (USNH 368516), referred to S. Allenit (Small) Engl. and Irmsch.
by ENGLER and IRMSCHER.

Montana: foothills, Midvale, June 18, 1903, L. Jf. Umback, No. 100a, referred to
S. Allents (Small) Engl. and Irmsch. (USNH 541458); Duck Lake, June 23, 1901,
Stuart Weller (USNH 411603), referred to S. Alleniit (Small) Engl. and Irmsch.
This form possesses crenately toothed leaves and a stout scape with rather compact
cymules.

The following specimens from the Selkirks and the Canadian Rockies
are also here referred to this species, with reservation.

BritisH CotumsBiA: Carbonate Draw, July 13, 1904, C. H. Shaw, No. 302, T. R.
Snyder (MBG 84110, UMH). These two sheets show the obvious differences in habit
assumed by this species. The Minnesota Herbarium specimens would be easily mis-
taken for S. rhomboidea,

WASHINGTON: near Whistler Pass, East Fork of Pasayton River, Okanogan
Forest, Okanogan County, alt. 2,340 m., August 4, 1916, Eggleston, 13416 (USNH
886307). Cymules at first compact. Some of the leaves are brown-pubescent beneath.

32 MINNESOTA STUDIES IN PLANT SCIENCE

The following specimens from the Selkirks and the Canadian Rockies
are similar to the Montana plants.

AgerTA: Laggan, Rocky Mountains, alt. 7,000 ft, F. K. Butters (UMH);
Canadian Rockies, F. K. Butters, No. 262 (UMH); Canadian Rockies, F. K. Butters,
No. 281 (UMH).

British CoLumsBia: Cougar Valley, alt. 6,500 ft., on limestone ledges, F. K. But-
ters, No. 340 (UMH); Cougar Valley, 5,500 ft, F. K. Butters, No. 363 (UMH);
Avalanche Crest, Selkirks, alt. 7,200 ft., F. K. Butters, No. 401 (UMH) ; Mt. Cheops,
alt. 8,000 ft., July 18, 1908, Butters and Holway, No. 347 (UMH).

Nelson’s type specimens of this species are much more slender and
delicate plants than the others enumerated above, probably due to the
peculiar conditions under which they were found growing, but the coinci-
dence of geographical distribution and the close similarity in floral and
vegetative characteristics leave little doubt that they are all S. saximontana
E. Nelson. The isolated outposts in the Black Hills and in the Wallowa
Mountains of Oregon seem rather remotely connected, but it is exceedingly
difficult to detect any differences in the plants except in the minutest details.
The cotype material of S. saximontana shows more orbicular petals, broader
and ciliolate sepals, lanceolate rather than linear bracts, and more multi-
cellular glands on the hairs in some cases, but the same variations are to be
found in other specimens.

The specimens referred to Saxifraga Allenu (Small), Engl. and Irmsch.
by ENGLER and IRMSCHER are erroneously determined inasmuch as the
leaves of this particular species are distinctly crenately toothed and the
inflorescence thyrsoid. When placed side by side the plants are obviously
different.

SAXIFRAGA TENNESSEENSIS Small, Bull. Torr. Bot. Club 23 :364. 1896.
Micranthes tennesseensis Small, Fl. S.E.U.S., 501. 1903.
A plant of characteristically stout habit (Plate VII).

Distribution—Along the Tennessee River in the vicinity of Knoxville, Tennessee.

Specimens examined:

TENNESSEE: Knoxville, May, 1898, Albert Ruth (MBG 84397, 84120); April to
May, 1897, MBG 84398; April, 1896, MBG 84396; April, 1895, 34120; April, 1895, MBG
84128; April, 1894, MBG 84290, UMH; Knoxville, May 5, 1895, J. G. Smith (MBG
34348, 84576).

NortH CaRoLInA: moist rocks, slopes of Craggy Mountain, Buncombe County,
May 18, 1898, Biltmore Herbarium, No. 565la (MBG 84333).

SAXIFRAGA VIRGINIENSIS Michx., Fl. Bor. Am. 1:269. 1803.

Saxifraga vernalis Willd., Hort. Berol. 43. 1805. (Unverified.)

Saxifraga virginica Nutt., Gen. 1:285. 1818.

Saxifraga pilosa Banks; D. Don. Trans. Linn. Soc. 13 :386. 1822, as syn.
(Unverified. )

JOHNSON: A REVISION OF THE SECTION BORAPHILA 33

Dermasea pilosa Haw. Saxifr. Enum., 9. 1821. (Unverified. )

Dermasea elongata Haw., l.c. (Unverified. )

A polymorphic species. Several rather distinct forms have been found
which differ considerably in carpel and seed characters, but the exact status
of which can not be determined at this time. Examination of Michaux’s type
specimens is necessary. The range of this species is peculiar and there
appears to be some marked differences between the northern and the south-
ern forms. The Minnesota specimens are without flowers and fruit so that
their nature can not be learned and must await further collecting. The
specimens from Missouri show certain characters which may prove suffi-
ciently distinct to warrant setting them apart as distinct species (Plate IX).

Dr. FERNALD (8) has described several forms of this species from
Massachusetts and New York. He notes that several aberrant forms of
S. virginiensis occur. In attempting to distinguish between S. nivalis and
S. virginiensis, however, Dr. FERNALD dwells on unimportant vegetative
and floral characters, whereas the outstanding differences between these two
species which are to be found in the carpels, gland, and receptacle, receive
no notice.

Distribution—New Brunswick and Quebec to northern Minnesota; in the South,
westward to Missouri.

Specimens examined:

MINNESOTA: exposed rocks, Two Harbors, May, 1891, J. H. Sandberg (UMH);
rocky slopes of Pelican Lake, August 30, 1909, C. O. Rosendahl (UMH); Flag
Island, Lake of the Woods, August 10, 1894, MacMillan and Sheldon, No. 2836.

Ontario: Kingston, May 7, 1881, J. Fowler (MBG 84353) ; Kingston, Jones Falls,
May 21, 1881, J. Fowler (MBG 84354); Kingston, May 30, 1901, J. Fowler (MBG
84568) ; Sparta, May 14, 1899, G. E. Fisher, J. Macoum (MBG 84347).

Micuican: Isle Royal at Tobin’s Harbor, August, 1900, W. A. Wheeler (“716
and 722”).

Onto: rocks, Hinckley, June 1, 1894, J. R. Watson (UMH).

New Yorxk: Ithaca, April 30, 1891, Herman Schrenk (MBG 84371); Sullivan
County, 1873, Eggert (MBG 84263); East Greenwich, 1861, Dy. Asa Fitch, No.
2419 (UMH). j

PENNSYLVANIA: Rawlinsville, 1884, Eggert (MBG 84366); along Conestoga
Creek, Lancaster County, April 12, 1889, J. K. Small (UMH); Allegheny County,
April 28, 1892, Adolph Koenig (UMH); Rawlinsville, ex herbarium, James Galen
(UMH); Safe Harbor, Lancaster County, April, 1893, Heller and Halbach, No. 804
(UMH); Glen Rock, York County, May 30, 1895, N. N. Glotfelter, M.D. (MBG
84352) ; rocks, Robinson’s Knoll, near Philadelphia, April 25, 1872, Redfield, No. 2021
(MBG 84357); Conestoga, May, 1887 (no collector cited), (MBG 84596): Swarth-
more, 1891, J. M. Greenman, No. 62L (MBG 737966) ; Westmoreland County, April
16, 1878, B. E. Pierron, No. 73 (MBG 777399); Mt. Alto, 1900, Jos. Illick (MBG
84070) ; on the Conestoga, Lancaster County, April 24, 1893, Heller and Halbach, No.
804 (MBG 84378) ; rocky places, Milford, May 15, 1905, C. A. Mill (MBG 84374);
banks of Conestoga Creek, May, 1894, A. F. Eby (MBG 84066) ; near Philadelphia,
1889, J. M. Greenman, No. 628 (MBG 738696) ; ex Bernhardi Herbarium, Philadelphia

34 MINNESOTA STUDIES IN PLANT SCIENCE

(MBG 84360); banks of the Conestoga, May 1, 1895, Chas. W. Etsenhour (MBG
84362) ; Philadelphia, Beynick (MBG 84355) ; Quakertown, May 12, 1907, Ethel Knight
Ball (MBG 83940).

MaIneE: rocky ground, Seal Harbor, Mt. Desert, June 27, 1890, Redfield herbarium
16492 (MBG 84356) ; exposed rocks, Orono, May 23, 1888, M. L. Fernald (UMH);
Orono, May, 1893, F. P. Briggs (UMH).

New HampsHirE: Hanover, May 5, 1883, C. H. Hitchcock (UMH).

MASSACHUSETTS: Amherst, May, 1890 and May, 1889, FE. L. Morris (UMH);
South Warmingham, 1890 (MBG 84369); Stony Brook, May 15, 1897, J. M. Green-
man, No. 629 (MBG 738779); West Roxbury, May 13, 1898, J. M. Greenman, No.
2950 (MBG 742616); Middlesex, Fels, April 19, 1898, J. M. Greenman, No. 569
(MBG 738766) ; Winchester, May 9, 1897, J. M. Greenman, No. 636 (MBG 738957) ;
Nonquit, E. P. (MBG 84367); Nonquit, May 1, 1889, E. P. No. 849 (UMH); dry
rocky banks, Lexington, Day and Fernald (MBG 84372); Arlington, May 3, 1891,
E. W. Dewart (MBG 84370) ; Nonquit, May 7, 1889, Miss Croll (MBG 84368).

Connecticut: Storrs, March, 1907, W. W. Ohlweiler (MBG 792271); rocky
woods, Southington, May 23, 1897, C. H. Bissell, Nos. 272 and 813 (MBG 84364).

New Jersey: Woodbridge, May, 1893, Rev. L. H. Lighthibe (UMH); Peapack,
April, R. C. Perry No. 50 (MBG 84562); Peapack, R. C. Perry (MBG 84564) ;
rocky banks, Greenwood Lake, Passaic County, May 19, 1907, Kenneth MacKenzie, No.
2560 (MBG 84584).

Section Chionophila, sect. nov.

Micranthes, Haw., Syn. Pl. Succ. 1812; in part.
Arabidia, Ledebour, Fl. Ross., 2. 1844; in part.
Boraphila, Engler, Verh. Zool.-Bot. Ver., Wien. 1869; in part.

Receptacle deeply adnate to the ovary; follicles elliptic-ovate, more or
less inflated, conspicuously contracted above; styles more or less elongate,
often stout; stigmas globose or discoid; gland discoid; the lower margin
closely approximated to the base of the calyx; filaments subulate, incurved
toward the apex; anthers oblong; sepals spreading; seeds narrowly ovate,
slightly winged on the margin, and more or less striate.

The most conspicuous characters of this section are the deep catigamitne
receptacle and the discoid gland which lies close to the base of the calyx.
The receptacle attains the greatest depth in Saxifraga rhomboidea, enclos-
ing the carpels for half their length. It is shallowest in S. franciscana. As
development proceeds the gland assumes a verrucose appearance and sur-
rounds the carpels as a broad zone immediately above the base of the calyx.
As a rule the beaks are stout, except in S. franciscana where they are quite
slender and also strongly bent at right angles or even recurved (Plates
II, III, XI).

The section embraces only three species, which, in their distribution, are
plants of high altitudes and of far northern latitudes. S. hieracifolia is
circumpolar in range, having its type locality in the Carpathian Moun-
tains of Hungary. The specimens examined are from scattering stations

ee

JOHNSON: A REVISION OF THE SECTION BORAPHILA 35

on the west coast of Alaska, from Point Barrow southward, including
the islands of Bering Sea. S. rhomboidea inhabits the higher elevations of
the Rocky Mountains, from southeastern British Columbia to central New
Mexico. Specimens of it have also been examined from a few scattering
stations in the Cascade Mountains of Washington and Oregon and the
Sierras of California. S. franciscana is restricted to the San Francisco
Mountains of Arizona. It somewhat resembles reduced forms of S. rhom-
boidea of which it is probably an offshoot.

The few species represented in the section can be readily separated
into two categories on characters of the inflorescence, and also on the color
of the petals. Thus we obtain one group with a capitate inflorescence and
white petals, which includes S. rhomboidea and S. franciscana, two rather
closely related species. The second group is represented by the single
species S. hieracifolia which possesses a spicate inflorescence and purple
petals. The two first named species are separated on the characters of the
_carpels and the beaks. The differences in the inflorescences are quite obvi-
ous, although in reduced forms of S. hieracifolia where only a few flowers
are present the inflorescence is capitate.

KEY TO THE SPECIES

I. Inflorescence capitate; petals white

1. Follicles stout; beaks erect or spreading.................... S. rhomboidea
2. Follicles small; beaks slender, strongly divergent............ S. franciscana
II. Inflorescence spicate, occasionally capitate; petals purple............ S. hieractfolia

SAXIFRAGA RHOMBOIDEA Greene, Pitt]. 3:353. 1896-1898.
Saxtfraga interrupta Greene, Plantae Bakerianae. (Unverified.)
Saxifraga rhomboidea austrina A. Nelson, Bot. Gaz. 31:394. 1901.
' Micranthes rhomboidea (Greene) Small, N. Am. Fl. 22:2:136. 1905.
Saxifraga nivalis Hook., Fl. Bor. Am. 1:248. 1832; in part; not S.
nivalis L.

Specimens of this species have been frequently referred to Saxifraga
nivalis L., from which it differs, however, both in general appearance and
in floral characters. The inflorescence of S. nivalis is distinctly corymbose
rather than capitate, the leaves are broadly ovate rather than oblong-ovate,
and the gland is a thick, fleshy, lobed ring raised above the base of the
calyx. S. nivalis is also typically a much smaller plant than S. rhomboidea
(Plates X, XI). Differences in the glands of the hairs will also be noted
on comparing the plates referred to.

Distribution—Rocky Mountains from British Columbia and Alberta to New
Mexico, and in the Cascade Mountains, and the Sierra Nevada.

Specimens examined:

ALASKA: Shumagin Island, 1871-72, M. W. Harrington, U.S. Coast Survey.
W. H. Dall’s exploration (MBG 84020).

36 MINNESOTA STUDIES IN PLANT SCIENCE

WasHIncTon: Cascade Mountains, Yakima region, June, 1882, ex herbarium,
Redfield, No. 14158.

IpAHO: ridges south from Wiesner’s Peak, alt. 2,050 ft., July 27, 1895, J. B.
Leiberg, No. 1390 (MBG 84077).

CALIFORNIA: Sierra County, 1874, J. G. Lemmon, No. 74. Sheet bears a specimen
labeled S. nivalis L. from Disco, Greenland, lat. 70°, legit Dr. Parry, ex herbarium
Dr. A. Gray, Redfield herbarium, No. 13683.

MontTANA: wet meadows, Flathead Pass, alt. 7,400 ft., June 29, 1905, J. W. Blank-
inship, No. 183 (MBG 84114) ; May 30, 1892, Dr. J. D. Heald (MBG 84373).

Wyominc: Bald Mountain, August 15, 1892, B. C. Buffum (UMH); near Mam-
moth Hot Springs, alt. 7,000 ft., June, 1893, E. H. Burglehaus (UMH, two sheets) ;
on Madison River, alt. 5,500 ft., June 26, 1860, Dr. F. V. Hayden (MBG 84134) ; Pole
Creek Hills, June 2, 1894, Aven Nelson, No. 128, as “Senecio lugens foliosus Gray.”—
Evidently a mistake in labels (UMH); Hone Creek, June 6, 1893, Aven Nelson
(UMH); Saw Mill Creeks, May 25, 1895, Aven Nelson, No. 1239 (MBG 84429,
UMH); Table Mountain, June 2, 1894, Aven Nelson, No. 127 (MBG 84430), as Sazri-
fraga nivalis L.; on grassy hillsides, Glen Creek, June 29, 1899, Aven and Elias Nelson,
No. 5559 (MBG 84542, UMH); on the moist slopes, Telephone Mines, Albany
County, August 1, 1900, Aven Nelson, No. 7900 (MBG 84540, UMH) ; head of Muddy
River, May 4, 1860, Dr. F. V. Hayden (MBG 84135) ; Gros Ventre Fork, Wind River
Mountains, on gravelly hills, alt. 6,800 ft., June 5, 1860, Dr. F. V. Hayden (MBG
84136 and 84137).

UtTaH: moist rocky soil, La Sal Mountains, alt. 9,000-10,000 ft., June, 1899,
C. A. Purpus, No. 6625 (MBG 84424); Fish Lake Mountain, alt. 11,500 ft., July 8,
1875, L. F. Ward, No. 324 (MBG 776622).

Cororapo: foothills west of Fort Collins, alt. 5,500 ft., May 7, 1896, C. F. Baker
(MBG 84435); Grand Mesa, alt. 9,000 ft., June 23, 1901, region of the Gunnison
watershed, C. B. Baker, No. 232 (MBG 84539, UMH); near Pagosa Peak, alk.
11,500 ft., August, 1899, C. F. Baker, No. 375 (MBG 83888); Howe’s Gulch, five
miles west of Fort Collins, alt. 5,500 ft., May 20, 1893, C. F. Baker, No. 3171 (UMH) ;
Marshall Pass, region of the Gunnison watershed, alt. 10,000 ft., July 19, 1901, C. F.
Baker, No. 493 (MBG 84541). (Plant on this sheet is very similar to Baker’s No. 509,
Small’s type of S. arnoglossa); Bear Creek Divide, west La Plata. Mountains, alt.
11,500 ft., June 29, 1898, Baker, Earle, and Tracy, No. 227 (MBG 84101, MBG 84433,
UMH); Breckenridge, 1887, S. D. Bereman, No. 731 (MBG 84578) ; summit Pike’s
Peak, July 29, 1901, F. FE. and FE. S. Clements, No. 430 as S. interrupta nana. (MBG
83890, UMH) ; Seven Lakes, alt. 3,500 m., July 8, 1901, F. E. and E. S. Clements, No.
442 (UMH, MBG 83889) ; foothills, alt. 6,000 ft., May 5, 1896, C. S. Crandall (MBG
84428); in rather dry woods, Berthoud’s Pass, as “Sarifraga nivalis Lin. forma
munor compacta.” (MBG 84253); no locality, September 14, 1874, G. Engelmann
(MBG 84181) ; Gray’s Peak; July 16, 1891, no collector cited, ex herbarium Fritchey,
1891 (MBG 84074) ; semi-meadow. Rabbit Ear Range, Routt County, July 17, 1803,
L. N. Gooding, No. 1582 (MBG 84093) ; Pike’s Peak, alt. 12,000 ft., August 13, 1884,
G. W. Letterman, No. 154 (MBG 84423); as S. nivalis L.; Gray’s Peak, No.— date,
G. W. Letterman, “S. nivalis’ (MBG 777454); Manitou Trail to Pike’s Peak at
timber line, August 13, 1884, G. W. Letterman (MBG 777456); foothills, Larimer
County, May, 1893, G. E. Osterhout (UMH); Tolland, dry sandy lowlands, alt. 9,000
ft., June 27, 1913, L. O. Overholts, No. 10026 (MBG 743694) ; from the headwaters
of Clear Creek, and the alpine ridges lying east of Middle Park, 1861, C. C. Parry,
No. 169, “S. nivalis L.” (MBG 84240); Mt. Baldy, alt. 11,000 ft., July 11, 1891,

JOHNSON: A REVISION OF THE SECTION BORAPHILA 37

E. C. C. (MBG 84421); Snow Edge, July 6, 1886, and Leadville, August 8, 1886,
Wm. Trelease (MBG 84436) ; Kempton, Mineral Pt., alt. 11,600 ft., June 20, 1887, ex
herbarium, Trelease.

New Mexico: Pecos River, July 4, 1898, G. E. Coghill, No. 54 (MBG 83873) ;
Santa Fe Mountains, August, 1884, F. H. Snow (MBG 776638) ; Pecos River National
Forest, approximate alt. 12,000 ft., July 11, 1908, Paul C. Standley, No. 4315 (MBG
84082).

SAXIFRAGA HIERACIFOLIA Waldst. and Kit., Pl., Hungar. 1:17, Table
XVIII. (Unverified.)

Micranthes hieracifolia Haw., Saxifr. Enum. 45, 1821.

Saxtfraga nivalis var. racemosa, Towns, it. Hung. 486.

Saxifraga rigida, Chr. Sm., in Herbarium DC., ex Engler, Monogr.
Saxifr. 151. 1872, as synonym.

Saxifraga uliginosa, Fisch., in litt. 1829. Engler, I.c., as synonym.

Saxifraga plantaginifolia Hook., in Parry’s Voyage, appendix 394.

Evaiezoa hieracifolia Raf., Fl. Tell., 2:70. 1836.

(Plates XI, V, VII.)

Specimens examined:

AvasKxa: Hall Island, July 14, 1899, Wm. Trelease, No. 4035, as S. hieracifolia
W. and K. (MBG 83975) ; Port Clarence, July 12, 1899, Wm. Trelease, No. 4936, as
S. hieracifolia W. and K. (MBG 83976); St. Matthew Island, July 15, 1899, Wm.
Trelease, No. 4037, as S. hieracifolia W. and K. (MBG 83975) ; St. Matthew Island,
August 11, 1891, J. Macoun No. “55” (MBG 83978); Oglamie, Pt. Barrow, Arctic
Sea, lat. 71°, 1883, Dr. John Murdock, No. 15704, ex herbarium, Dr. A. Gray as
S. hieractfolia W. and K. (MBG 83977) ; “Alaska,” Mr. Reaman, as S. nivalis L.,
Herbarium of Dartmouth College. Caulescent, leaves more deeply dentate.

It is very doubtful whether these specimens are actually Saxifraga
hieracifolia W. and K. Owing to conditions incident to the World War it
has been impossible to obtain authentic material from Hungary (the type
locality) for comparison. The specimens at hand are very different, how-
ever, from a Norwegian specimen in the Herbarium of the University of
Minnesota. The latter has broad elliptic-ovate leaves on short petioles, and
possesses a distinctly raised gland which persist sas a flange around the
carpels, a condition not found in the Alaskan specimens. In the characters
of the petals, sepals, and hairs, however, the two plants are similar. The
gland of the Alaskan specimens is conspicuously depressed and smooth in
the early stages, and the flowers are characteristically stellate. It was the
form of the petals no doubt that led Hooker (16) to infer that the American
plant, to quote his words, “appears exactly intermediate between S. penn-
sylvanica and S. nivalis, more allied, however, to the former” but its deep
receptacle, flat, appressed, discoid gland, and stout erect follicles indicate
no very close relationship to either, petal-form in this case furnishing no
criterion of relationship. Furthermore, the American specimens do not

38 MINNESOTA STUDIES IN PLANT SCIENCE

possess the plantagineous leaves that might suggest the S. plantaginifolia
of Hooker (In Parry’s Second Voyage, appendix 394), although this form
of leaf is characteristic of the Norwegian specimen. Hooxer’s (16, p. 49),
description reads “‘foliis petiolatis obovato-spatulatis repandodentatis.” In
a specimen cultivated by himself, HAwortu (14) found the leaves to be
“lanceolata obtusa subavenia crassiuscula,’—not so different from Willde-
now’s description “foliis oblongo-lanceloatis glabris repando-dentatis.”

SAXIFRAGA FRANCISCANA (Small) Fedde, Just, Bot. Jarhesb. 33:1:613.
1906.

Micranthes franciscana Small, N. Am. Fl. 22 :2:144. 1905.

(Plates XI, VIII-X.)

Specimens examined:

Arizona: San Francisco Mountains, alt. 11,500 ft., August 27, 1889. F. H.
Knowlton, No. 127, as S. nivalis L., referred to this species by ENGLER and IRMSCHER
on sheet No. 258261 (USNH).

The following specimen is doubtfully referred here:

CauirorniA: Mt. Goddard, Sierra Nevada Mountains, alt. 11,500 ft. July 24-26,
1900; Hall and Chandler, No. 70y, as S. nivalis L. (UMH). This plant differs from
the above in having a wholly glabrous, purple, and taller (11 cm.) scape, and a fewer-
flowered, less compact inflorescence. The differences in the floral characters are very
slight and difficult to detect. The beaks are shorter, and the gland seems to persist
as an inconspicuous flange above the base of the calyx. No seeds.

Section MicRANTHES (Haw.) Tausch, Hort. Canal., fase. 1. 1823.

Micranthes Haw., Syn. Pl. Succ., 320. 1812; as genus.
Boraphila Engler, Verh. Zool.-Bot. Ver., Wien. 19. 1869; in part.
Micranthes Small, N. Am. FI., 22:2:132. 1905; in part.

Ovary subinferior; receptacle bowl-shaped, shallow to deep. Gland
fleshy, broad, discoid, and almost completely enveloping the ovary, and
usually considerably raised above the base of the calyx, its surface at
length conspicuously convoluted and verrucose, and its lower margin persist-
ing as a distinct flange around each carpel. Carpels distinct, but connected
by the adnate receptacle and at least the basal portion of the gland. Fila-
ments subulate, flattened, uncinate. Sepals spreading or reflexed. Petals
lanceolate to orbicular. Seeds various, tuberculate to smooth, in some forms
winged. Inflorescence paniculate to corymbose, the cymules often glom-
erate. Hairs uniseriate, the glands various (Plates II, III, XII, XVIII).

HawortH (13) founded his genus on Saxrifraga pennsylvanica, distin-
guishing it by its “lorate” petals (“petala lorata distantia”). Tauscu (30)
included S. nivalis L., apparently on the similarities in the calyx, but he
erroneously describes the seeds as smooth. SERINGE (26) included S. vir-
giniensis Michx., S. davurica Pall., and S. hieracifolia W. and K., on the

JOHNSON: A REVISION OF THE SECTION BORAPHILA 39

same grounds, apparently, that TauscH included S. mivalis, but in the case of
S. davurica he was probably misled by inadequate material, as the species
properly belongs in the section Hydatica. LEDEBouR (18) combined the sec-
tion with Arabisa of TauscH on the non-stoloniferous habit. Torrey and
Gray (31) included S. erosa Pursh (S. micranthidifolia Haw.) with some
hesitation.

Hooker (16) made no mention of this species but included a new one
which he had named Saxifraga reflexa. Finally Smatt (27) adopted
‘Micranthes (Haw.) as a generic name for a combination of HAwortTH’s
genera Robertsonia, Dermasea, and Aulaxis, and RAFINESQUE’S (24) genus
Steiranisia. In that sense SMALL’S genus was synonymous with the sections
Micranthes and Hydatica of Tauscu and his followers. The result was
an aggregation of species of widely different floral characteristics.

The section Micranthes contains the majority of those species with a deep
receptacle in which the tendency toward epigyny has advanced the farthest
and which are further characterized by the conspicuous, fleshy, discoid gland
which is elevated a considerable distance above the base of the calyx, and
which persists as a distinct flange around the carpels at maturity. The
gland is the most distinctive characteristic of the section.

In Saxfraga pennsylvanica and S. californica the gland is more elevated
than in the other species. Other minor distinctions appear here and there.
A reference to the plates will show the striking similarities in the glands
and carpels, particularly in S. integrifolia, S. nidifica, and S. montanensis.

The carpels are distinct to the base but are connected laterally by the
adnate receptacle and at least by the lower portion of the gland. In some
cases as in S. pennsylvanica, S. Sierrae, and S. oregana, the receptacle
does not keep pace with the growth of the carpels so that as the follicles
are pushed apart at maturity the receptacle is compressed and gives the
appearance of a completely hypogynous condition.

As a whole the section presents the most difficult problem in the entire
series. Polymorphism appears to be a general characteristic and compli-
cates the situation. In no other group, it appears from the herbarium speci-
mens, have botanists been so at variance on the question of species.

Two main types of petals occur, the narrowly lanceolate and the orbicu-
late. Between these there are many variations. The strictly lanceolate
type is restricted exclusively to the eastern species, namely to S. pennsyl-
vanica and its allies. The extreme orbiculate type is represented by S.
californica and S. napensis. Curiously enough the orbiculate type is as
constant at this extreme as the lanceolate type at the other. In the group
represented by S. integrifolia the variations of petal-form are perplexing
in the extreme.

40 MINNESOTA STUDIES IN PLANT SCIENCE

Clavate filaments do not occur in the section. The subulate type pres-
ent is usually flattened with age and incurved more or less sharply at the
top.

The receptacle is typically bowl- or urn-shaped, but varies considerably
within certain groups, such as that represented by S. integrifolia. The
relative depth of the receptacle furnishes a basis for dividing the species
into two principal divisions.

Variations in the inflorescence are also marked within certain limits,
not only as between immature and mature stages but as between individuals
of the same species, as in S. nidifica and S. pennsylvanica. The writer has
not been able to determine whether the latter type is a specific or merely
an individual variation. A well-defined corymbose inflorescence occurs in
S. fragosa and S. napensis. Glomerate cymules are characteristic of S.
nidifica and its allies.

Seed characters appear to be quite constant within certain groups. Thus,
winged seeds are characteristic of S. Sierrae and related forms, and
tuberculate seeds of S. pennsylvanica and its allies. Intermediate forms
seem to show varying types, but in most cases the material has been defi-
cient in mature specimens. Several types of seeds have been found in
specimens referred to S. pennsylvanica.

The principal categories have been founded on the relative depth of the
receptacle—one shallow, the other deep. The species of the first division
are separated into two categories on the basis of petal-form, the first com-
prising all the species which have typically lanceolate petals, namely S.
pennsylvanica and allies, the second comprising a half dozen species which
have orbiculate or ovate petals, namely S. Sierrae and S. califormica and
their allies. Characters of the follicles, petals, and leaves, since they are
constant within the limits indicated, serve to separate the minor categories
of each group, respectively. Finally the specific distinctions are based on
certain vegetative and floral characters that are well marked. In the second
division some differences in the gland occur, showing two lines of develop-
ment, one in which the type has remained constant, the other in which it has
become considerably narrowed, at least as far as could be determined from
the material available. The latter group is limited to a single species. The
remaining species, only six in number, are grouped in two categories on the
basis of the colors of the flowers. Characters of the petals serve to separate
the species of the first category. The species of the second category fall
into two equal groups on characters of the inflorescence, in which the
species are designated by characters of the inflorescence and leaves. An
attempt has been made to arrange the species according to their affinities,
and if the specimens be laid out in series according to the accompanying
analytical key the significance of the arrangement may be evident.

JOHNSON: A REVISION OF THE SECTION BORAPHILA Aj

Geographically the species are almost wholly confined to the United
States. There are two well-marked distributional belts practically within
the same latitudinal range, one eastern, the other western. The eastern belt
runs westward from New Hampshire and Massachusetts into Minnesota,
Iowa, and Missouri, occupying practically all the area between the Great
Lakes and the Ohio River, with a narrow strip in Canada along Lake Erie
and Lake Ontario. The belt is dominated by S. pennsylvanica. At the
extremities of the range certain peculiar types have been differentiated, to
which may be added certain apparently distinct types from the center of
the range. The exact status of these types can not be established at present.
An interesting feature in this connection is the remarkable similarity be-
tween certain specimens from the Yellowstone National Park region and a
specimen from the Winnebago Valley in Minnesota.

The western distributional belt lies in the form of a deep inverted
crescent, the points of which rest on the same parallel of latitude, on the
southern boundary of Colorado, in the Rocky Mountains, and a short dis-
tance north of Tulare Lake, California, respectively. The arms of the
crescent follow the courses of the mountains northward, with the main
body occupying the watershed of the Clark’s Fork, the Clearwater, and the
Snake rivers, and the lower course of the Columbia. A small, apparently
detached belt, occupied by a single species, surrounds the shores of Puget
Sound. ‘This large crescent-shaped belt almost completely surrounds the
Great Basin. Practically nothing is known concerning the exact limits of
the ranges of the individual species, particularly the western ones, as the
collections are in most cases very meager. The species on the. whole are
plants of the lower altitudes, not a few being marsh plants.

KEY To THE SPECIES

I. Receptacle ccmparatively shallow; follicles spreading and often apparently superior
at maturity
A. Petals narrowly lanceolate
a. Follicles and beaks elongate
1. Petals white; follicles green

(a) Branches of inflorescence diffuse and delicate............ S. Forbestt

(b) Branches of inflorescence stout, ascending.......... S. pennsylvanica

2A, JEREMY XE cra Shgtaiethy SAU Lc LN ee eee ALC S. purpuripetala
b. Follicles globose, inflated; brownish

ieshollicles\large!stoutspbeaks thick 3% 505. 0s Sees. S. crassicarpa

Zs Hollictes!smalll;| beaks\very, short.: . 022. se een le S. winnebagoensis

B. Petals orbiculate to oblong
a. Leaves elongate, of a lanceolate type; tall species
1. Leaves narrowly lanceolate or oblanceolate; petals oblong..... S. oregana
2. Leaves broader and mostly shorter; petals ovate or orbiculate..S. Sierrae

42 MINNESOTA STUDIES IN PLANT SCIENCE

b. Leaves of an ovate type
1. Leaves small, sharply dentate
(a) Leaves shallowly dentate to subentire; petals oblong..... S. parvifolia
(b) Leaves coarsely and evenly serrate-dentate; petals elliptic...........
wheal et NONE eg Su URN NLS Poca sili d/o wa ch aici SUARLY ant a rr S. gracillima
2. Leaves larger, repand to sinuate dentate; petioles elongate
(a) Leaves glabrous, thin; petals ovate-lanceolate.............. S. fallax
(b) Leaves pubescent; petals orbiculate to oval............ S. californica
Ii. Receptacle deeper; follicles not strongly spreading
A. Gland broad, fleshy, verrucose or convoluted, conspicuous
a. Flowers commonly greenish; cymules glomerate
1. Petals mostly shorter than the sepals, sometimes longer; inflorescence

COMETACEE 8 SMe aN AN icizisies. bclenta atte vine eee S. nidifica
2. Petals mostly exceeding the sepals, or wanting; inflorescence commonly
Spicatel 2) soar Aleman cee eae eC IUAV AO LCD) NO oa ar S. montanensis

b. Flowers white, generally conspicuous
1. Inflorescence more or less contracted; cymules glomerate

(a) Inflorescence stout, thyrsoid; stout plants.............. S. integrifola
(b) Inflorescence delicate, subcapitate; delicate plants.......... S. aprica
2. Inflorescence broad, corymbose; petals orbicular
(a) Leaves broadly ovate; flowers large:...........225 meee S. fragosa
(b) Leaves narrowly ovate; flowers small.................... S. napensts
B. Gland narrow, smooth; follicles short, purple, very smooth....... S. laevicarpa

SAXIFRAGA APRICA Greene, Bull. Torr. Bot. Club 23:25. 1896.
Saxifraga umbellulata Greene, Erythea 1:222. 1893.
Micranthes aprica (Greene) Small, N. Am. Fl. 22:2:135. 1905.

Specimens examined:

CALIFORNIA: wet shady places above Donner Lake, Nevada County, July 4, 1892,
C. F. Sonne, No. 90 (MBG 84536) (doubtful) ; Mt. Eddy, Siskiyou County, August
18, 1903, Copeland, No. 3845, as Saxifraga nidifica, det. Dr. Greene; distributed by
C. F. Baker (MBG 84117) ; Siskiyou County, Mt. Eddy, im open gravelly places in the
forest, alt. 6,800 ft., 4. A. Heller, No. 12081.

SAXIFRAGA CALIFORNICA Greene, Pitt. 1:286. 1889.
Saxifraga virginiensis Benth., Pl. Hartw. 311. 1848.

Micranthes californica (Greene) Small, N. Am. Fl. 22:2:140. 1905.
Saxifraga virginiensis Michx. var. californica (Greene) Jepson, Fl. Mid.
Western Calif. 268. 1901.

Distribution—Central California, north and south of a line between Lake Tahoe
and San Francisco Bay.

Specimens examined:

CaLirorniIA: Mt. Tamalpais, March 30,1889, E. L. Greene (UMH) ; Sonoma County,
March 13, 1892, Michener and Bioletti (UMH); near Clear Creek, Butte County, alt.
175 ft., April 1-15, 1897, H. E. Brown, No. 150 (UMH); Loma Prieta, Santa Clara
County, eastern slope, April 1-7, 1894, J. Burt Davy, No. 476 (UMH); San Juan,
San Benito, June, 1903, A. D. E. Elmer, No. 4911 (UMH, MBG 84118); Clinton,
Amador County, Sequoia gigantea region, March 29, 1892, Geo. Hansen, No. 92 (UMH
and MBG 84282); Mokelumne Hill, Calaveras County, alt. 2,000 ft., April 1, 1896,

JOHNSON: A REVISION OF THE SECTION BORAPHILA 43

Geo. Hansen, No. 1533 (MBG 84283) ; woods near Cazadero, Sonoma County, April
24, A. A. Heller (MBG 84279) ; 1896, Mrs. R. M. Austin, No. 859 (MBG 84281) ;
Alma, Santa Clara County, common about moist banks and thickets, March 23, 1902,
C. F. Baker, No. 370 (MBG 84275); Sonoma Creek, eastern base of Mt. Hood,
Sonoma County, March 12, 1892, Heller and Brown, No. 5040 (MBG 84276) ; foot-
hills west of Los Gatos, Santa Clara County, April 6, 1904, A. A. Heller (MBG
84280) ; Santa Rosa, Sonoma County, March, 1880; near Mariposa, Mariposa County,
March and April, 1882; Glacier Point, Mariposa County, July 1, 1885; Mariposa,
Mariposa County, May, 1886; Elk Grove, Sacramento County, March 31, 1894; south
of Farrington, Stanislaus County, March 30, 1894; north of Folsom, Placer County,
April 5, 1894, Stockton Creek, Mariposa County, April 20, 1889, Santa Rosa, Sonoma
County, March 20, 1880; ex herbarium J. W. Congdon.

Saxifraga crassicarpa sp. nov.

Saxifraga pennsylvanica L., of authors in general.

Receptaculum ovarii parti inferiori adnatum; folliculi crassi subsphaerici

stylis crassis valde divergentibus; stigmatibus crassis discoideis; filamenta
subulata, angustata, 2.5 mm. longa; sepala cuneata, acuta, base coalitiuscula

reflexa ; petala 3 mm. longitudine angustatissime lanceolata, acuminata I-ner-
via ; semina ovalia conspicue. tuberculata.

Receptacle shallow; follicles thick, stout, subspherical, beaks stout,
strongly divergent; stigmas broad, thick, flattened; filaments 2.5 mm. long,
subulate, narrow ; sepals cuneate, acute, somewhat united at the base, reflexed
about 2 mm. long lateral nerves, 2 short; petals very narrowly lanceolate,
acuminate, l-nerved, about 3 mm. long; leaves 6-19 cm., uniformly elliptic-
lanceolate, purplish towards the base, thick, leathery, sparsely pubescent
on both sides, minutely and rather distantly and evenly denticulate, obtuse,
ciliolate, contracted into a short membranous margined petiole-like base;
scapes stout, about 90 cm. high, glandular-pubescent, more denselv so above,
branching from above the middle; inflorescence paniculate-cymose, branches
elongate, slender, ascending, bearing the few-flowered cymules towards the
ends; perennial from a stout, woody caudex.

The follicles of this plant are very different from the common Saxifraga
pennsylvanica; they are very stout, globose, and thick-beaked. The seeds
are relatively large and conspicuously tuberculate. The few-flowered coarse
inflorescence is also characteristic (Plate XII).

Distribution—As far as known occasional from northeastern Minnesota to north
central Illinois.

Specimens examined:

Minnesota: St. Louis River, July 23, 1886, Arthur, Bailey, and Holway, No.
B329 (UMH).

WIScoNSIN: swamps in Mirror Lake, Sauk County, July 14, 1903, H. Eggert
(MBG 83863); Walworth County, common in low grounds among thickets, near
Lake Beulah, July 26, 1905, James I. Shannon, No. 125a (MBG 84478, 84479).

44 MINNESOTA STUDIES IN PLANT SCIENCE

ILLINOIS: wet prairie near Wady Petra, Stark County, May 12, 1897, Virginius H.
Chase (MBG 84469).

SAXIFRAGA FALLAX Greene, Bull. Torr. Bot. Club 23:25. 1896.
Micranthes fallax (Greene) Small, N. Am. Fl. 22:2:141. 1905.

A delicate species of the aspect of Saxifraga Careyana Gray (Plate
TV):

Distribution—Not definitely known. “Oregon and California” according to
Small (l.c.).

Specimens examined:

CaLiForNIA: Emigrant Gap, July 1, 1882, Marcus E. Jones, No. 3587, as S. fallax
Greene by ENcLER and IrMscHER (USNH 221361).

SAXIFRAGA ForBEsII Vasey, Am. Entom. and Bot. 2:288. 1870.
Saxifraga pennsylvanica L., Sp. Pl. 399. 1753.

Distribution—Not definitely known; apparently limited mostly to eastern Mis-
souri, through St. Louis and Jefferson counties, and adjacent counties in Illinois.

Specimens examined:

Missourrt: wet shaded sandstone cliffs, Silica, Jefferson County, May 5, 1898,
Colton Russell (MBG 83960) ; Silica, May, 1890, ex herbarium Frederick Wislizenus,
No. 939 (MBG 84580); rocks, St. Louis, May, 1887, H. Eggert (UMH); Jefferson
County “Sandfelsen,” May 11, 1887, H. Eggert (MBG 83856, 739323); St. Louis,
shady rocks, May and July, 1880, H. Eggert (MBG 84582) ; Pacific, Franklin County,
May 8, 1896, H. Eggert (MBG 738820, 83861) ; St. Louis, shady sandstone, July, 1880,
H, Eggert (MBG 83857) ; Jefferson County, May 11, 1887, H. Eggert (MBG 83862) ;
St. Louis County, on the Meramec, under shady rocks, June 15, 1875 (MBG 83820) ;
April 15, 1880 (MBG 83858); June 20, 1884 (MBG 83859), with large foliaceous
bracts; Allenton, June 20, 1880 (MBG 772053, 780787) ; April 16, 1880 (MBG 775263,
83818), wet sand rocks, G. W. Letterman (MBG 777205).

Itt1nors: June, 1875, D. G. Vasey (MBG 84223); Makanda, “type locality,” on
moist cliffs, May 5, 1900, Baker, No. 6168 (MBG 83868).

SAXIFRAGA FRAGOSA Suksdorf ex Small, Bull. Torr. Bot. Club 23:363.
1896.

Saxifraga claytoniaefolia Canby, Small, Bull. Torr. Bot. Club 23:365.
1896.

Saxifraga integrifolia var. fragosa (Suks.) Rosendahl, Beibl. Bot. Jahrb.,
Nr. 83. 1905.

Saxifraga integrifolia Hook. var. claytoniaefolia (Canby) Rosendahl, J.c.

Micranthes claytoniaefolia (Canby) Small, N. Am. FI. 22:2:141. 1905.

Saxifraga fragosa var. leucandra Suks., in herb.

Distribution —‘Washington to Oregon” (Small), lc.
Specimens examined:

WASHINGTON: wet rocks near the Columbia River, Klickitat County, May, 1892,
W.N. Suksdorf, No. 1727 (MBG 84331, UMH); wet mossy cliffs, Almota, Whitman
County, May, 1897, A. D. E. Elmer, No. 139 (UMH).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 45

OrEGonN: the Dalles, Wasco County, April 19, 1902, E. P. Sheldon, No. 10109
(UMH and MBG 83872).

Saxifraga gracillima sp. nov.

Receptaculum ovario infra medium adnatum, ovariis depressis in
glandula latissima profunde insertis, folliculi parvi inflati, superne valde con-
tracti, stylis primum gracillimis demum plus minusve abbreviatis divergenti-
bus, stigmatibus globosis, filamenta subulata apice valde incurvata carpidiis
supercurrentia, sepala parva ovatave cuneata base minime coalita 1-nervia,
petala elliptica vel elliptico-lanceolata sepalis multi longiora base valde con-
tracta.

Receptacle adnate to the ovary below the middle; ovary depressed,
inserted deeply in a very broad gland; follicles small, inflated, strongly con-
tracted above styles, at first very slender, at length shortened and recurving,
globose; filaments subulate, strongly incurved at apex and exceeding the
(young) carpels; sepals small, ovate or cuneate, slightly united at the base,
1-nerved; petals elliptic or elliptic-lanceolate, much longer than the sepals;
strongly contracted at the base.

Leaves basal, 1.5-5 cm., ascending-spreading, ovate, smaller, sometimes
elliptic, coarsely serrate-dentate, thin, acute, sometimes purplish, margins
ciliolate or not at all so, abruptly contracted into a very slender petiole about
equalling or sometimes exceeding the blades. Scapes erect or assurgent,
very slender to delicate, commonly solitary, 15-22 cm. high, thinly glandu-
lar-pubescent, cormybosely branched above; branches of the inflorescence
few and filiform; flowers white, conspicuous, not numerous, in loose, open
cymules. Bracts small to minute, especially the upper, sparsely-ciliolate on
the margins. Perennial from a short, slender, fibrous-rooted caudex. A
graceful, delicate species. The relatively large petals, very small carpels
and calyx-lobes, and the strongly serrate-dentate leaves with their slender
petioles are the most obvious characteristics. The shallow receptacle gives
the appearance of a hypogynous condition with age. There is great varia-
tion in the size of the petals in different flowers. Mature fruit not seen
(Plate XTV).

Distribution.—Known only from type locality. —
Specimens examined:

CALIFORNIA: Cafion of Big Chico Creek, Butte County, March 19, 1914, 4. A.
Heller, No. 11214, as Micranthes californica (Greene) Small, Type (UMH, MBG
745924). Plants on MBG sheet smaller than the specimens in the Herbarium of the
University of Minnesota.

SAXIFRAGA INTEGRIFOLIA Hook., Fl. Bor. Amer. 1:249. 1840.

This species appears to be a perplexingly polymorphic one. Such has
been the confusion in regard to it that identical forms have been variously

46 MINNESOTA STUDIES IN PLANT SCIENCE

referred to Saxifraga fragosa, S. Sierrae, S. oregana, S. bracteosa, S.
plantaginea, and S. nivalis. A single specimen from Forest Grove, Oregon,
(Prof. Joseph Marsh, No. 85) which appears the closest to S. imtegrifolia
Hook. as to leaf characters, differs radically from that species in carpels and
petals. In spite of the excellent delineations in Hooxer’s (16) plates the fact
remains that no such fruit or carpels occur in the entire section Micranthes,
as are portrayed in Figures 2, 4, and 5 of Hooxer’s Plate LXXXVI.
Hooker’s figures fail to show the characteristic broad gland surrounding
the carpels and so generally well-marked in the section Micranthes; more-

over Figure 4 (and to less extent Figure 5) portrays the carpels as united’

above the middle, which is not the case except as they are adnate laterally
to the gland. This and the further fact that Figure 4 (and to less degree
Figure 5). portrays the carpels as seated in a deep receptacle are both con-
tradictory of the description “ovario omnino libero” (Vol. 1:249). Figure 5
might well belong to such species as S. saximontana (Sect. Dermasea) or
S. rhomboidea (Sec. Chionophila). No plants have been found that con-
form to Hooker’s characterizations of the leaf-margins as “imtegerrimis”’
(Plate XVI).

The region about the lower Columbia River whence Dr. Scouler’s plants
came needs to be thoroughly explored, and the original collection studied,
in order to establish the identity of this species.

For the present Savifraga bracteosa Suksdorf and its varieties are

included in this species. Better material is necessary to establish the exact |

status of S. bracteosa but as far as can be determined there is no essential
difference between it and specimens from the same locality that have been
referred to S. iniegrifolia Hook.

Saxifraga laevicarpa sp. nov.

Folliculis brevibus inflatis laevibus purpureis receptaculo infra medium
insertis, stylis brevibus valde divergentibus vel recurvatis, glandula laeve
crassa angustata elevata, filamentis subulatis superne attenuatis incurvatis
antheris parvis marginibus purpureis; sepalis virideis vel purpureis 2 mm.
longis angustatis cuneatis vel deltoideis base coalitiusculis acutis reflexis,
petalis 3-4 mm. longis purpureis perangustatis lanceolatis acutis 1-nerviis
base contractis; seminibus crassis ovatis saepe angulatis carinatis sulcatis
evidente tuberculatis.

Follicles short, inflated, smooth, purple, inserted below the middle in the
receptacle; styles short, strongly divergent or recurved; gland smooth,
thick, narrow, elevated; filaments subulate, attenuated above, incurved,
anthers small, the margins purple; sepals green or purple, 2 mm. long, nar-
row, cuneate or deltoid, somewhat united at the base, acute, reflexed; petals
3-4 mm. long, purple, very narrow, lanceolate, acute, 1-nerved, contracted
at the base; seeds thick, ovoid, often angled, carinate, sulcate, tuberculate.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 47

Leaves 2-4 cm., narrowly ovate, dull green to purplish, glabrous or
nearly so, shallowly denticulate, margins sometimes irregular, contracted
into a slender petiole about equalling the blade. Scapes 25 cm. or shorter,
solitary, erect, purplish, rather copiously glandular-pubescent. Inflorescence
racemose Or somewhat spicate, elongate to shorter; cymules contracted or
glomerate, apparently sessile, the lower much shorter than the elongate inter-
nodes; flowers white, rather few. Bracts lanceolate, small. the lowest about
$ mm., purplish, equalling or exceeding the small glomerate cymules. Peren-
nial by a short, fibrous-rooted caudex.

Distinctive features: purple, smooth carpels with widely divergent to

recurving beaks and the peculiar thick, smooth, flange-like gland (Plate
XVII).

Distribution—Puget Sound region.
Specimens examined:

Wasuincton: Friday Harbor, April 3, 1910, Wm. Moodie. Type! Tacoma,
May 11, ——, T. C. Frye. ‘

SAXIFRAGA MONTANENSIS Small, Bull. Torr. Bot. Club 23 :56 :367. 1896.

Saxifraga subapetala E. Nelson, Erythea 7:169. 1899.

Micranthes montanensis Small, N. Am. FI. 22:2:139. 1905.

Micranthes subapetala (E. Nels.) Small, l.c.

Micranthes brachypus Small, l.c. (Unverified.)

This seems to be a good species, but it intergrades strongly with the
more western S. columbiana Piper, and S. plantaginea Small (here included
under S. nidifica Greene, g.v.), while some forms that have been referred to
S. montanensis Small, by some authorities, closely approximate Micranthes
arnoglossa Small in floral characters. The species can not be distinguished
from E. Netson’s S. subapetala, but the leaves of the latter have longer
petioles. Other specimens are difficult to distinguish from Suksdorf’s S.
bracteosa (No. 1728) from the Columbia River, West Klickitat County,
Washington.

SMALL’S species is based on Mr. Frank Tweedy’s No. 58, collected in
southwestern Montana in July, 1888. The original description is greatly at
variance, however, with the later one in North America Flora 22:2 :139.
1905. It is difficult to connect the two with the same plant, and leads to the
suspicion that Tweedy’s No. 58 is not the plant that the author in his later
work considers to be Micranthes inontanensis.

Distribution—Region of Yellowstone National Park; limits of range not defi-
nitely known.

Specimens examined:

Montana: Bozeman, meadows June 14, 1905, Blankinship, No. 521 (doubtful) ;
Spanish Basin, Gallatin County, alt. 6,500 ft., June 30, 1897, Rydberg and Bessey, No.
4263, referred to S. montanensis by ENGLER and IrMSCHER (USNH 360877) (doubtful).

48 MINNESOTA STUDIES IN PLANT SCIENCE

Wyominc: near Mammoth Hot Springs, alt. 6,000 ft. July, 1893, and 7,000 ft.,
July, 1893, both along meadow borders, F. H. Burglehause (UMH); Mammoth Hot
Springs, July, 1893, F. H. Burglehause, referred to S. montanensis by ENGLER and
IrMScHER (USNH 45808).

SAXIFRAGA NAPENSIS Small, Bull. Torr. Bot. Club 25:316. 1898.

Micranthes napensis Small, N. Am. Fl. 22:2:140. 1905.

A tall graceful species bearing a strong resemblance to Saxifraga fragosa
Suks. It is based on a coilection of J. W. Bigelow (Whipple’s expedition)
and George Thurber No. 496 (Small, /.c.), Napa Valley, California. There
is a discrepancy between the original description (/.c.) and that in the North
American Flora (l.c.). In the latter the petals are described as “‘wanting.”
The descriptions of the calyx and the carpels also differ. The leaves are
described as pubescent. In the specimens cited below they are glabrous.
No mature fruit available (Plate XVII).

Distribution—Not definitely known; apparently restricted to a small region north
of Tulare Lake, California. “California,” according to Small (l.c.).

Specimens examined:

CaLIFoRNIA: region of Dinkey Creek, Fresno County, alt. 7,500 ft., June 25, July
15, 1900, Hall and Chandler, No. 386 (UMH, two sheets; MBG 84426).

SAXIFRAGA NIDIFICA Greene, Erythea 1:222. 1893.

Saxifraga plantaginea Small, Bull. Torr. Bot. Club 23:366. 1896.

Saxifraga columbiana Piper, Bull. Torr. Bot. Club 27 :393. 1900.

Micranthes columbiana (Piper) Small, N. Am. Fl. 22:2:135. 1905.

Micranthes plantaginea Small, l.c.

Receptacle deep ; ovary deeply immersed in the gland; follicles stout, erect
or not strongly spreading; beaks stout, spreading; stigmas globose, at length
flattened, thick. Gland conspicuous, fleshy, at length rugose, verrucose or
convoluted, the margin lobed, at first crowding the base of the calyx, at
length elevated and persisting as a distinct flange around the follicles. Fila-
ments subulate, flat, uncinate, mostly shorter than the sepals, often very
short ; anthers commonly large, the margins purple with age. Sepals 1.5-3.0
mm., oblong to triangular-ovate or deltoid, at length reflexed, acute to obtuse
or rounded at apex, margins entire or denticulate; 1-2 nerved, occasionally
more, the nerves simple or branched, reaching to or beyond middle, free or
uniting with the mid-nerve below the apex. Petals 1-3 mm., sometimes very
small to minute, oblong, elliptic, obovate to spatulate-orbicular, varying
greatly in the same individual and even in the same flower; acute, obtuse, to
rounded and retuse at apex, orbicular forms with a more or less clawlike
base; mid-nerve simple or with one or more elongated branches, these some-
times coalescing with mid-nerve below apex. Seeds thick, fusiform, some-
times pointed or slightly winged at the ends and marked with more or less

JOHNSON: A REVISION OF THE SECTION BORAPHILA 49

prominent longitudinal ridges of inconspicuous tuberculate teeth, or nearly
smooth or striate. Leaves 2-10 cm., lanceolate to oblong or ovate, varying
considerably, mostly glabrous, margins ciliolate and obsoletely denticulate to
sinuate or rarely dentate. Scapes about 45 cm. high and less, stout, erect,
finely glandular-pubescent, more copiously so above, branched near the top
or occasionally with one or two long slender branches ‘arising from below
the middle. Inflorescence paniculate-cymose, compact and spicate in some
forms to open and branched below; upper cymules mostly sessile or nearly
so, glomerate. Perennial from a short, stout caudex; sometimes a cluster
of reddish bulblets at the base of the scape.

GREENE’S (11) original specimens of S. nidifica have not been available,
but his description of it (J.c.) indicates that it is identical with S. planta-
ginea Small and S. columbiana Piper. The mass of bulblets noted by Greene
at the base of the scape in his plants is also to be found in the specimens of
S. plantaginea from Spokane, and in plants from Idaho and Montana.
SMALL (27) separates S. nidifica from S. plantaginea by its “arachnoid” as
well as glandular pubescence, while the latter is regarded as being merely
glandular-pubescent, a distinction which does not hold since both kinds ot
hairs occur in specimens of S. plantaginea from Montana and Idaho.

More or less confusion has also prevailed as to the distinctions between
S. plantaginea and S. columbiana. S. plantaginea is based on Sandberg and
Leiberg’s collection of May, 1893, from Spokane, Washington. The species
is distinguished, according to SMALL, by its “broad greenish petals which
are exceeded by the calyx.” Later Piper (22, p. 393) based a new species,
S. columbiana, on specimens from the same region, which he differentiated
from S. plantaginea Small on size, and on the characters of the petals, the
latter, he writes, being “a larger plant in every way,” with “white, elliptic
or oblong, 1-nerved” petals. Comparing the cotypes and other authentic
specimens it is found that Piper’s own collection of S. columbiana, namely
No. 1808, from Pullman, Washington, is a larger plant than any of the
specimens of S. plantaginea, Small; at the same time other collections, iden-
tical with Piper’s No. 1808, from adjacent localities in Idaho and western
Montana, show the same characteristics as to size (Heller No. 3037, Lewis-
ton, Idaho; Sandberg, Kootenai County, Idaho, July, 1887; Williams No.
401, Columbia Falls, Montana). Another specimen collected by Piper at
Pullman, Washington, May 24, 1894 (without number) and identical with
his No. 1808, can not be distinguished from the cotype of S. plantaginea
(namely that of Sandberg and Leiberg, Spokane, Washington, May, 1893).
A detailed examination of a great deal of material of the two species shows
the petals to be extremely variable and therefore wholly unreliable as specific
characters (Plate XV).

50 MINNESOTA STUDIES IN PLANT SCIENCE

Disiributton—Montana, northern Idaho, and eastern Washington to Nevada.
Limits of the range not definitely known.

Specimens examined:

WasHInctTon: Spokane, alt. 1,200 ft., May, 1893, Sandberg and Leiberg (UMH);
Spokane, alt. 1,000-2,000 ft., May, 1893, Sandberg and Leiberg (UMH); Spokane, alt.
2,000-3,000 ft., May, 1893, Sandberg and Leiberg (UMH, MBG 84123); Hangman
Creek, Spokane County, alt. 1,510 ft., May 16, 1893, Sandberg and Leiberg, No. 16
(MBG &560) ; in pebbly moist hillsides, Pullman, May,.1897, A. D. E. Elmer, No. 126
(UMH, MBG 83874); on springy hillsides, Pullman, May 24, 1894, C. V. Piper, No.
1808 (UMH).

IpaHO: moist hillsides, Lewiston, April, 1892, J. H. Sandberg (UMH); Nez
Perces County, May, 1892, J. H. Sandberg (MBG 84422, UMH); meadows Kootenai
County, May, 1891, J. B. Leiberg, No. 81 (UMH); grassy slopes, upper ferry, Clear-
water River above Lewiston, April 30, 1892, Sandberg, MacDougal, and Heller, No.
73 (MBG 775340); about Lewiston, Nez Perces County, alt. 1,500-2,000 ft., May 7,
1896, A. A. and E. G. Heller, No. 3037 (MBG 83886, UMH).

SAXIFRAGA OREGANA Howell, Erythea 3:34. 1895.
Micranthes oregana (Howell) Small, N. Am. Fl. 22:2. 1905.

This tall species differs from its allies by its very shallow receptacle,
elongated seeds, and long, narrow, oblanceolate leaves which are wholly
entire. The flattening of the receptacle by the subsequent growth and wide
divergence of the follicles produces a nearly hypogynous condition which
was clearly pointed out by HoweEtt (17) as follows: “calyx attached to the
ovary at the very base.” To this species Piper (21) referred Suksdorf’s
No. 1729 and Henderson’s No. 2488 both of which, in leaf characters differ
radically from Howell’s specimens, although very similar as to inflorescence.
The receptacle also is deeper and broader than in Howell’s plant, the petals
consistently narrower, and the seeds narrower and more winged at the ends.
SMALL (27) attributed the elongated narrow leaves of Howell’s plant to its
growing in water, but this hardly accounts for the differences in the recep-
tacle. The description of Micranthes oregana Howell in North American
Flora 22 :2 :138, 1905 is decidedly at variance with Howell’s original descrip-
tion, from which it might be inferred that it was based not on Howell’s
specimens, but on Suksdorf’s No. 1729, a plant referred to S. oregana
Howell by Prrer (Plate XIII).

Distribution—Known only from the type locality.
Specimens examined:

Orecon: Lake Labish near Salem, June 21, 1893, Thos. Howell, No. 1498, ex
herbarium, J. W. Congdon (UMH, MBG 84460).

SAXIFRAGA PARVIFOLIA Greene, Pitt. 3:116. 1896-1898.
Micranthes parvifolia (Greene) Small, N. Am. Fl. 22:2:141. 1905.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 51

Distribution—Not definitely known. “Oregon to California” according to Small

(he.).
Specimens examined :

OrEecon: Grant’s Pass, April 13, 1887, Thos. Howell, No. 1134, as Savifraga
wirginiensis Michx., var. (MBG 84381).

Dr. Greene’s type specimen was collected by Howell at Grant’s Pass,
Oregon, 1892. No specimens of this collection have been available, but the
above-named specimen agrees with Greene’s description (Plate XIV).

SAXIFRAGA PENNSYLVANICA L. Sp. Pl. 399. 1752.

A detailed study of a great deal of material that has passed among
herbaria as S. pennsylvanica has disclosed a number of very different forms
which the writer suspects may prove to be sufficiently distinct to merit
specific rank. The differences are most marked in the carpels, gland, and
seeds, but are also plainly evident in the leaves and the inflorescence. As
far as can be determined these forms differ also in their habitat, which is
significant and calls for investigation. Before their status can be deter-
mined it will be necessary also to see the Linnean type. Certain obvious
variations were noted by the early botanists; thus MicuHaux (19) in a note
writes as follows: “Variat floribus in ramis paniculae conglobatis et
corymbosis.’ These forms were given varietal rank by SERINGE (26): (a),
conglomerata and (b), corymbifera, respectively, the former having a dense
panicle and conglomerate flowers (“panicula densa floribus conglomeratis’),
the latter a lax panicle and open corymbs (“panicula laxa corymbosis dis-
tantibus”). Both varieties were recognized by Torrey and Gray (31).
Whether these varieties are merely individual variations, denoting early and
late stages of growth, or specific distinctions can not be determined; that
remains to be learned in the field. An eastern form described by Dr.
FERNALD (8) as S. pennsylvanica L. forma fultior, with large “dilated oval’
bracts is of the type possessing a stout contracted or conglomerate inflores-
cence, a form which reappears in western parts of the range. What aspect
this plant assumes at maturity is not known, but the writer is of the opinion,
based on a careful study of the carpels and the gland, that it is probably a
distinct species. The problem of species in this connection must therefore
be left to future investigations (Plate XII).

Saxifraga purpuripetala sp. nov.

Receptaculum ovario infra medium adnatum; carpidia inferne distenta
superne valde contracta stylis conicis conniventibus stigmatibus parvis;
folliculi infra medium inflata stylis divergentibus vel recurvatis; glandula
lata elevata convulata vel verrucosa; filamenta subulata angustata viridia
vel purpurescentia ; sepala 2-4 mm. longa cuneata vel deltoidea reflexa base

52 MINNESOTA STUDIES IN PLANT SCIENCE

coalita viridia vel purpurea; petala 3-4 mm. longa purpurea lanceolata saepe
perangustata; semina crassa ovata 1-1.3 mm. longa angulata carinata con-
spicue longitudinaliter tuberculata.

Receptacle united to the ovary below the middle; carpels distended
below, strongly contracted above; styles conical, connivent; stigmas small;
follicles inflated below the middle, styles divergent or recurved; gland broad,
elevated, convoluted or verrucose; filaments subulate, narrow, green, or
purple; sepals 2-4 mm. long, cuneate or deltoid, reflexed, united at the base,
green or purple; petals 3-4 mm. long, purple, lanceolate, often very narrow;
seeds thick, ovoid, 1-1.3 mm., angled, carinate, prominently, longitudinally
tuberculate. i

Leaves oblong, elliptic or elliptic-lanceolate, 5-20 cm., dull green to
brownish green above, paler yellowish-tinged beneath, glabrous, thin, laevi-
gate, cbscurely or shallowly denticulate, tapering to short, broad, mem-
branous-margined, sheathing petiole-like bases. Scapes 45 cm. tall or less,
at first stout, later more slender, sometimes branching from below, densely
glandular-pubescent, the glands purple, thinner below but hairs longer,
whitish and mostly not gland-bearing. Inflorescence paniculate-cymose,
elongate, rather compact at first, opening with age, the branches ascending,
the upper much shortened. Bracts narrow lanceolate, the lower sometimes
foliaceous. Haris uniseriate, of various lengths; glands oblong. Perennial
from a stout, woody caudex.

This new species has been long regarded as a purple-flowered form of
S. pennsylvanica L. but the color of the petals, the peculiarities of the car-
pels, gland, seeds, and habit are sufficiently distinct, it seems, to warrant
specific rank (Plate XII).

Distribution—Known only from two localities.

Specimens examined:

VERMONT: Peacham. June 13, 1894, Ferdinand Blanchard (MBG 751771, June 3,
1883, MBG 751772, August, 1892, MBG 751775).

New Jersey: near Peapack, ex herbarium, R. C. Perry (MBG 84563).

SAXIFRAGA SIERRAE (Coville) Smali, Bull. Torr. Bot. Club 23 :362.
1896.

Saxifraga integrifolia Sierrae Coville, Proc. Biol. Soc., Wash. 7:78.
1892.

Micranthes Sierrae (Coville) Heller, Muhlenbergia 2:52. 1905.

Micranthes arnoglossa Small, N. Am. Fl. 22 :2:138. 1905.

“Blades of the larger leaves 8-12 cm. long, oblong-lanceolate to elliptical-lanceolate,
conspicuously serrate-denticulate, from glabrous to sparingly clammy-hairy above and
beneath, thinner and more distinctly veined than in the type; petiole and margin of
the leaf toward the base ciliate with clammy hairs; otherwise as the type form.”

JOHNSON: A REVISION OF THE SECTION BORAPHILA 53

The above description is borrowed from the original of CoviILLE (2)
who described the plant as a variety of Saxifraga integrifolia Hook. The
type specimen, which has not been available for study, is in the United
States National Herbarium. “No. 1705, Death Valley Expedition, Col-
lected August 25, 1891, about eight miles northwest of Whitney Meadows
on the Headwaters of Kern River, Sierra Nevada, Tulare County, Cali-
fornia, by Frederick V. Coville.” CoviLLe states that the plant is found in
the Rocky Mountains also. Collections from various localities show several
distinct forms referred to S. Sierrae of which only those are here given
recognition which conform most closely to the descriptions of CovILLe.
The others constitute such a confusing array of forms that the problem of
their status must await further study. In addition all Rocky Mountain
specimens referred to Micranthes arnoglossa Small are included here.

Specimens examined:

CauirorniA: Tulare County, summer, 1904, C. F. Baker, No. 4428, collector,
J. D. Culbertson, det. by Dr. Greene as Saxifraga oregana Howell (MBG 84087);
Coyote Creek, Tulare County, July 20, 1904, C. F. Baker, No. 3265, collector, J. D.
Culbertson, det. by Dr. Greene as Saxifraga oregana Howell. (MBG 84085) ; Sisson,
Siskiyou County, June 15, 1905, A. A. Heller, No. 8031 (MBG 84091) as Micranthes
Sierrae (Coville) Heller.

Cotorapo: Marshall Pass, alt. 10,000 ft., July 19, 1901, C. F. Baker, No. 509
(UMH). This specimen cited as the “type collection” of S. arnoglossa Greene n. sp.

Saxifraga winnebagoenis sp. nov.

Receptaculum ovario non alte adnatum; folliculi praeparvi globosi
superne multo contracti stylis brevissimis stigmatibus parvis; filamenta
subulata gracilia brevia; sepala parva oblonga vel ovata vel deltoidea acuta
base evidente coalita reflexa; petala non adsunt; semina ovata longitu-
dinaliter tuberculata.

Receptacle shallow; follicles very small, globose, slightly spreading,
abruptly contracted above into very short beaks; stigmas small. Filaments
subulate, slender, 1.7 mm., exceeding the sepals. Sepals 1.3-1.5 mm., oblong
or ovate to triangular, acute, plainly united at base, green, reflexed; lateral
nerves 2, reaching to the middle. Petals wanting in specimens. Seeds about
1 mm., broadly ovoid-fusiform, carinate, with longitudinal ridges of short
tuberculate teeth. Hairs of various lengths, uniseriate; glands oblong, some-
times broadest at top, consisting of a row of superposed cells. Leaves 12
cm. long and less, few, elliptic-lanceolate or obovate, minutely denticulate to
subentire, shallowly or inconspicuously repand, obtuse to acute, strigillose
pubescent on both sides, somewhat leathery, gradually contracted into short,
sheathing bases. Scape erect, solitary, about 35 cm., yellowish-green, loosely
glandular-pubescent, branching above. Inflorescence racemose-cymose, nar-
row; cymules contracted at the ends of the branches of the inflorescence.
Perennial from a short, stout, coarsely rooted caudex (Plate XIII).

54 MINNESOTA STUDIES IN PLANT SCIENCE

This plant is very different from the eastern S. pennsylvanica. The
unusually small follicles and the rather leathery, yellowish-tinged leaves
which are velvety in appearance are the most distinctive characters. In
many respects it is very similar to some Rocky Mountain specimens from
the United States National Herbarium which ENGLER and IRMSCHER (7)
have referred to S. montanensis Small. Spanish Basin, Gallatin County,
Montana. Rydberg and Bessey No. 4263 (USNH 360877); Bozeman,
Montana, Plankinship, No. 521 (MBG 84116).

Distribution—Known only from the type locality.
Specimens examined:

MINNESOTA: Winnebago Valley, Houston County, June 12, 1899, W. A. Wheeler,
No. 180 (UMH) Type!

Section Tricarpum, sect. nov.

Micranthes, Small, Fl. S.E.U.S. 501. 1903, as genus; in part.
Boraphila, Rosendahl, Beibl. Bot. Jahrb., Nr. 83:70. 1905; in part.

Receptacle deeply adnate to the ovary; carpels three, distinct, connected
by the receptacle and above by the gland; follicles stout, inflated; styles
stout, divergent, rigid; stigmas discoid; gland thick, slightly elevated above
the base of the calyx.

Monotypic, including the species described below.

The carpels of this section are generally three together, although four
are sometimes present. The receptacle attains a considerable depth, enclos-
ing the carpels for half their length, in this respect resembling the section
Chionophila, and Micranthes in part. The beaks are peculiarly rigid and
conical, or somewhat pyramidal, and frequently widely spreading. The
stigmas are considerably flattened and sometimes incompletely two-lobed.
The gland is not elevated to such an extent as in Micranthes, but as in that
section the lower margin persists as a distinct flange around the follicles.
The follicles remain connected by the receptacle and the gland, even almost
to the base of the styles, but are otherwise distinct to the base, although
they appear to be united entirely (Plates II, III, XVIII).

The single species of which the section is composed is distributed over a
rather limited region from southwestern Missouri to the vicinity of Dallas,
Texas. Collectors have described it as a plant of sandy or cherty soils.
Its range follows the southwestern extension of the Ozark vegetation belt
and flanks the Atlantic forest zone, according to HARSHBERGER (15).

SAXIFRAGA TEXANA Buckl., Proc. Phil. Acad. 1861:455. 1862.
Micranthes texana (Buckl.) Small. Fl. S.E.U.S. 501. 1903.

Carpels 3, occasionally 4; at maturity stout, pyramidal, contracted above
into stout conical, spreading beaks. Stigmas small, flattened or discoid.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 55

Gland a fleshy, more or less verrucose or convoluted, flattened disc, its lower
margin persisting as a thin flange around each follicle, and slightly elevated
above the base of the calyx. Filaments subulate, flattened, uncinate; anthers
oblong, somewhat contracted above, the margins at length purple. Sepals
mostly broad, oblong to triangular-ovate, 1.5-2.5 mm., obtuse to rounded
and sometimes retuse at the apex, slightly united at the base; lateral nerves
usually 2, variously branched, sometimes coalescing with the mid-nerve
below the apex; entire, glabrous, sometimes irregularly denticulate, spread-
ing. Petals 2.4 mm., white, broadly orbicular or ovate, to spatulate or ellip-
tic-lanceolate; base broad, clawlike in orbicular forms; mid-nerve with 2-4
branches which are often variously forked and coalescent. Leaves basal,
small, glabrous ovate to elliptic-lanceolate, 6 cm. long or less, abruptly to
gradually contracted into a narrow, flattened, petiole-like base mostly
shorter than the blade; margins of blades sinuate to shallowly crenate-
dentate, ciliolate or eciliolate. Scapes 7-20 cm. high, slender to stout, soli-
tary or several together, rather sparsely glandular-pubescent, the hairs
commonly elongated, uniseriate. Inflorescence contracted, paniculate-cymose
or merely cymose, sometimes capitate, the cymules glomerate; lowest
branches usually shorter than the internodes in branched inflorescences.
Perennial from a short fibrous-rooted caudex which often produces several
short shoots.

The species has sometimes been referred to S. wirginiensis from which,
however, it can easily be distinguished by the carpels and especially by the
mature follicles, the number of which is generally three, as noted above.
The two species differ also strongly in seed characters (Plate XVIII).

Distribution—From southwestern Missouri to the region of Dallas in northeastern
Texas.

Specimens examined:
Missouri: Reding’s Mill, April 8, 1909, B. F. Bush, No. 5475 (MBG 84597);

Reding’s Mill, Newton County, local in chert barrens, April 9, 1909, Ernest J. Palmer,
No. 1647 (MBG 760519).

Texas: Terrell-Elmo, April 6, 1903, J. Reverchon, No. 3729 (MBG 84557) ; black
sandy land, between French Town and Dallas, J. Reverchon, No. 281 (MBG 84556) ;
Brazos County, sandy land, March 10, 1900, N. Ness, No. 1994 (MBG 84554) ; damp
sands, Elmo, April 6, 1903, J. Reverchon, No. 3729 (MBG 84555).

No state cited, February 27, 1878, J. F. Joor (MBG 84552); ex herbarium,
J. Reverchon (MBG 84552).

Section ARABISA Tausch, Hort, Canal, Fasc. 1. 1823.

Spatularia, Haworth, Saxifr. Enum. 47. 1821.

Gymnopera, D. Don, Trans. Linn. Soc. 13 :343. 1822; in part.
Arabidia Seringe, DC. Prodr. 4:40. 1839; in part.

Hexaphoma, Raf., Fl. Tell. 2:66. 1836.

Hydatica, Neck. Elem. 2:387. 1790. Small, N. Am. Fl. 22 :2:148. 1905.
Boraphila, Engler, Verh. Zool.-Bot. Ver., Wein. 19:521. 1869; in part.

56 MINNESOTA STUDIES IN PLANT SCIENCE

Ovary superior; carpels elliptic-oblong or ovoid, united to below the
middle; gland a somewhat constricted, yellowish to purplish girdle above
the base of the carpels; follicles ovoid, constricted more or less at the base,
the sides marked with more or less prominent longitudinal lines or ribs;
strongly contracted above into slender divergent beaks. Stigmas small,
globose. Filaments subulate. Sepals oblong, lanceolate-oblong, to ovate-
lanceolate, acute to obtuse, becoming strongly reflexed. Corolla irregular,
white, or yellowish-tinged; upper three petals sagittate, the two lower
oblanceolate or merely lanceolate. Seeds elliptic-fusiform, carinate, marked
with deeply pectinate, longitudinal ridges. Hairs multiseriate, with globose,
multicellular glands. Leaves basal, rosulate, alternate, with elliptic to
oblanceolate, deeply and coarsely to shallowly dentate blades, contracted
below into broad and flat to slender, petiole-like bases. Scapes solitary or
tufted, stout to slender, erect or assurgent, more or less branched above or
from near the base, in some forms prolifically, mostly coarsely glandular
pubescent. Inflorescence paniculate or paniculate-cymose, often diffuse,
with few to numerous flowers, these in the western species replaced to a

greater or less degree with bulbils. Perennial herbs from short, slender, ©

fibrous-rooted caudices.

The species comprising this section form a clearly defined natural group,
exhibiting striking similarities in the characters of the floral organs. The
similarities in the carpels and capsules are so close that it is difficult to dis-
tinguish one species from another on these characters.

One of the marked characteristics of this section is the complete
hypogyny. In addition there is a prolific production of bulbils in the
western species. Only two of our American species, namely S. leucanthe-
mifolia and S. ferruginea, attain any appreciable size, and these two are also
very similar in vegetative characters, herbarium collections of the latter
having been often referred to the former. Their capsules and seeds can be
distinguished with difficulty.

All the species appear to be calciphobe. With the exception of S.
foliolosa and S. ferruginea they do not present contiguous or overlapping
ranges but occupy separate belts along the periphery of the continent. S.
foliolosa is circumpolar in its range, with an isolated island of it on Mt.
Katahdin, Maine, the only locality from which it is known south of extreme
Arctic America (FERNALD, in Rhodora 2:167). It is probable that it tran-
sects the limit of the range of S. ferruginea on the Bering Sea coast of
Alaska. The most extended range is that of S. ferruginea, namely from the
coast of Alaska, along Bering Sea, to Montana and Oregon. S. bryophora
occupies a limited alpine region of the Sierra Nevada of California, radiating
from Lake Tahoe. According to HARSHBERGER (15) it is unknown from the
Cascades. It is no doubt an offshoot of S. ferruginea, with which it once

JOHNSON: A REVISION OF THE SECTION BORAPHILA SY,

formed a continuous specific line. S. leucanthemifolia occupies a narrow
belt of the higher Appalachians, from Maryland to northeastern Georgia.
Only a single American specimen of what is probably Saxifraga stellaris
L. has been available for study—a specimen from Labrador. None of the
western plants examined can be referred to this species, although a specimen
from Mt. Hood, Oregon, mentioned later, bears a strong resemblance to it
in some vegetative characters (Plates II, XIX).

When HawortH (14) in 1821 divided the old Linnean genus Saxifraga
he brought together the species S. leucanthemifolia Michx., S. Clusit Gouan,
and S. stellaris L. under the new genus Spatularia. Haworrs clearly diag-
nosed the characters which define the group. He further divided the genus
into two divisions, viz., Verae Rectirostraiae, including S. leuwcanthemi-
folia and S. Clusu; and Spuriae: Curvirostratae, including S. stellaris.
The material available is inadequate to verify this distinction. TAauscu
and his followers include these species in the section Arabidia, but SERINGE
(26) erroneously included S. arguta D. Don. Leprzour (18) failed to.
recognize the distinction made by these writers when he combined the sec-
tions Arabidia and Micranthes, as did also ENGLER (4) in creating the sec-
tion Boraphila. SmMatt (27) was the first to recognize the interpretation put
upon this group by HawortH, when he adopted that author’s genus Spatu-
laria.

SECTION ARABISA

Key TO THE SPECIES

I. Leaves dentate

A. Leaves deeply and coarsely dentate; inflorescence loose....S. leucanthemifoha

B. Leaves more shallowly dentate, mostly towards apex
a. Inflorescence commonly diffuse; some or most of flowers often replaced by
[PONTE EDS) Gia) GMOS Gl IIE IEPD PARRY SN AS Ie and UR RE GO SUNT DU S. ferruginea

b. Inflorescence reduced

1. Flowers, except the terminal, replaced by bulbils; scapes erect, rigid,
glabrous; branches of inflorescence short and mostly at right angles with

SEEM ene eves ane n ema ar A cL cuaiulle 2 Mac, SugiaVc lial eia) ge! tual aia aline tole aia S. foliolosa

2. Not bulbil-bearing, flowers conspicuous; scapes pubescent or

IAD EOL SMC Nee re Ue ROR DAR REDS LARUE) YUN CE UN ENGR S. stellaris®

ime Weavesventize. small lanceolate... use Pi iisles Cee cvs eels c's saekiee ieee S. bryophora

SAXIFRAGA BRYOPHORA A. Gray, Proc. Am. Acad. 6:533. 1865.
_ Saxifraga leucanthemifolia Michx. var. integrifolia Engler, Monogr.
Saxiir, 135, 1872.

Spatularia bryphora (A. Gray) Small, N. Am. Fl. 22:2:148. 1905.
(Plate XIX).

10 The detailed treatment of this species is deferred for a future paper.

58 MINNESOTA STUDIES IN PLANT SCIENCE

Distribution—Eastern California in the Sierras, from Fresno County northward
to Sierra County, region of Lake Tahoe.

Specimens examined:

CaLIFoRNIA: road to Meadow Lake from Webber, Nevada County, grassy slope,
August, 1892, Sonne, No. 89 (MBG 83870); Placer County, August-October, 1892,
A. M. Carpenter (UMH); meadows near Black Mountain, Fresno County, alt. 9,500
ft. July, 1900, Hall and Chandler, No. 595 (UMH, USNH 390665); state survey
(5943), Bolander (UMH, USNH 320768) ; 1875, J. G. Lemmon, No. 97 (MBG $4199
and 84200); Sierra Nevada, peak northeast of Camp 117, alt. 9,000 ft., July 6, 1863,
W. H. Brewer, No. 1807 (USNH 320777) ; Kings’s Valley, Lassenfs Peak, July, i882,
Mrs, Austin (USNH 139349); on same sheet, Sierra County, J. G. Lemmon, No. 97
(USNH 139348); Donner Lake, July, 1889, T. S. Brandegee (USNH 735784).; ridge
south of Donner Pass, alt. 7,500 ft., Nevada County, August 10, 1903, 4A. A. Heller,
No. 7143 (USNH 467528) ; on dry mountain slope near Webber Lake, alt. 7,000 ft.,
August 4, 1900, J. B. Leiberg, No. 5260 (USNH 610507) ; Half Moon Lake, Lake
Tahoe region, alt. 7,760 ft., August 9, 1909, E. A. McGregor, No. 73 (USNH 612809) ;
Heather Lake, Lake Tahoe region, August 18, 1909, E. A. McGregor, No. 148 (USNH
612876).

SAXIFRAGA FERRUGINEA Graham, Edinb. New Phil. Jour., p. 348. July,
1828.

Saxifraga notkana Moc., ic. ined., Seringe, DC. Prodr. 4:40. 1830, as
synonym.

Hexaphoma ferruginea Raf., Fl. Tell. 2:66. 1836.

Saxifraga Bongardi Presl., ex Engler, Monogr. Saxifr. 135, 1872, as
synonym of Saxifraga leucanthemifolia var. Brunoniana (Bong.) Engler.

Saxtfraga leucanthemifolia var. Brunoniana (Bong.) Engler, Monogr.
Saxdirwloos Lose:

Saxifraga stellaris Brunoniana Bong., Mem. Acad. St. Petersb. 6:2:140.
1832. (Unverified.)

Saxifraga nootkana Moc., ex Rosendahl, Beibl. Bot. Jahrb., Nr. 84, p. 65.
1905.

Spatularia ferruginea (Graham) Small, N. Am. Fl., 22:2:150. 1905.

Spatularia Vreelandu Small, N. Am. Fl., 22:2:149. 1905.

Spatularia Brunoniana Small, N. Am. Fl. 22:2:149. 1905.

Spatularia Newcombei Small, N. Am. Fl. 22:2:149. 1905.

Follicles elliptic-ovoid, united below the middle, contracted above into
short slender beaks; stigmas small, globose; sides of follicles marked with
more or less conspicuous lines or ribs; carpels ovoid. Gland a fleshy narrow
girdle above the base of the carpels. Filaments subulate, flattened, rigid,
dilated to adnate at the base; anthers oblong, yellowish to purple. Sepals
oblong, oblong-ovate, elliptic-oblong, 2-4 mm., reflexed, acute to obtuse,
sometimes ciliolate ; lateral nerves commonly 2, free, mostly simple, reaching
beyond the middle and sometimes uniting with the mid-nerve at the apex.
Petals 3-7 mm., the upper three more or less sharply sagittate to unequally

JOHNSON: A REVISION OF THE SECTION BORAPHILA 59

or scarcely at all so, commonly with two yellowish spots below the middle;
the two lower oblanceolate to elliptic-lanceolate or ovate; all contracted at
the base into conspicuous, mostly narrow claws. Hairs generally elongate;
glands globose, whitish to purple. Leaves basal, crowded, 1.5-10 cm., long,
oblanceolate to spatulate or cuneate, more or less pubescent, sometimes
copiously so—to glabrous or nearly so; more or less coarsely serrate-
dentate above the middle, or incised-toothed, especially towards the apex and
above the middle, or remotely serrate-dentate, the teeth elongate, narrow;
margins ciliate. Scapes 3-4.5 cm. high, solitary or clustered, erect or assur-
gent, more or less glandular pubescent, branching at the top or from near
the base, often profusely; branches ascending or spreading. Inflorescence
paniculate-cymose, the flowers few to numerous, sometimes all but the
terminal replaced by bulbils. Bracts lanceolate to oblanceolate, the upper
small, the lower foliaceous with a few to several coarse apical teeth; leaf-
like in some forms.
A polymorphic species of wide range displaying considerable variation
in vegetative characters. The different forms, however, do not appear to be
of sufficient stability to warrant according them specific rank (Plate XIX).

Distribution.—Islands of Bering Sea, and Alaska to Montana, Idaho, and Oregon.
Specimens examined :

ALASKA: summit of high mountains near Yes Bay, August 21, 1890, Thomas
Howell, No. 1621 (UMH); Windham Bay, June 11, 1905, J. D. Culbertson; Kodiak,
July 3, 1898, Wm. Trelease, Harriman Alaska expedition.

BritisH CotumBiA: Nanaimo, Vancouver Island, June 4, 1889, John Macoun
(USNH 219860) ; Moresby Island, July 9, 1900, Wilfred H. Osgood (USNH 620999) ;
Glacier, alt. 5,800 ft., July 20, 1901, C. O. Rosendahl, No. 139 (UMH); Avalanche
Mountain, Glacier, August 21, 1902, C. O. Rosendahl, No. 968 (UMH) ; Asulkan Pass,
alt. 8,000 ft., August 17, 1904, F. H. Walker, No. 493 (UMH); woods above Revel-
stoke, alt. 4,000 ft., July 10, 1905, Chas. H. Shaw, No. 897 (UMH) ; Cowichan-Alberni
Trail, summit of pass, Vancouver Island, alt. 3,000 ft., June 27, 1906, C. O. Rosendahl,
No. 1735 (UMH); along the trail above Alberni, Vancouver Island, June 30, 1906,
C. O. Rosendahl et al., No. 1623 (UMH); summit of Mt. Edinburgh, Vancouver
Island, July 30, 1906, C. O. Rosendahl, No. 1735 (UMH); Fish Creek Valley, alt.
5,500 ft., August, 1906, F. K. Butters and E. W. D. Holway, No. 240; Mt. Edinburgh,
Vancouver Island, alt. 2,000 ft, June 7, 1907, C. O. Rosendahl and F. K. Butters
(UMB) ; rocky ledge, Cowichan Lake, Vancouver Island, June 15, 1907, C. O. Rosen-
dahl, No. 1809 (UMH); Glacier, alt. 4,100 ft., July 28, 1907, F. K. Butters and
E. W. D. Holway (UMH); snow sheds, Glacier, alt. 4,200 ft., August 24, 1907, F. K.
Butters and E. W. D. Holway (UMH); Mt. Abbott, Glacier, alt. 7,000 ft. July 22,
1908, F. K. Butters and E. W. D. Holway (UMH); Glacier, H. C. Cowles, No. 167
(MBG 744924) ; Queen Charlotte Islands (MBG 71701), as S. Newcombei; Selkirks,
C. H. Shaw, No. 451.

WasuincTon: Skagit Pass, Cascade Mountains, August 24, 1892, Lake and Hull,
No. 468 (UMH) ; Stevens Pass, alt. 4,000-6,000 ft., August, 1893, J. H. Sandberg and
J. B. Leiberg, No. 1836 (UMH); dry rocky places, Goat Mountains, July 12, 1895,

60 MINNESOTA STUDIES IN PLANT SCIENCE

O. D. Allen, No. 199 (UMH); along rivulets, Mt. Ranier, alt. 6,500 ft., August, 1895,
C. V. Piper, No. 2026 (UMH); north fork of Bridge Creek, Okanogan County, alt.
5,000 ft., September, 1897, A. D. E. Elmer, No. 691 (UMH); Olympic Mountains,
Clallam County, August, 1900, A. D. E. Eliner, No. 2644 (UMH).

MonTANA: Sperry Glacier, Marcus E. Jones (UMH); mountain sides, Sperry
Glacier, September 1, 1903, L. M. Umbach, No. 812 (USNH 542027) ; Sperry Glacier,
upper temperate life zone, alt. 7,500 ft., Marcus E. Jones, Herbarium of the University
of Montana. .

Collections of this plant were by some earlier writers referred to S. leu-
canthemijolia of the Appalachians, which is, however, distinct. Herbarium
specimens referred to Saxifraga Bongardi Presl. “in herb. Engler,” are all
S. ferruginea. ENGLER (5) gives S. Bongardi Presl., S. notkana Moc. ic.
ined., and S. stellaris var. Brunoniana Bong. as synonyms of his S. leucan-
themtfolia var. Brunoniana (Bong.) ic. ined. t. 422? (“in America bor.
occid. prope Notka.”) as a synonym of his S. stellaris var. vulgaris (Ser.
MSS.). RosenpDauL (25), referring to SERINGE, accepts Mocin’s Saxifraga
notkana as a distinct species apart from S. ferruginea, with a slight modifi-
cation of the orthography. As synonyms ROSENDAHL adjoins S. stellaris
var. Brunoniana Bong., S. Bongardi Presl., and S. leucanthemifolia. var.
Brunoniana (Bong.) Engler. Contemporaneously with RosENDAHL, SMALL,
in North American Flora 22:2:149, published two new species, under the
genus Spatularia Haw., namely Spatularia Newcombei, the former based on
a collection by F. K. Vreeland 1048, from Glacier Basin, Montana, and the
latter on a collection from Queen Charlotte Islands (Torreya 2:55. 1902).
The range of the former, to quote SMALL, extends from “British Columbia
to Montana and Colorado.” ‘That of the latter is limited to “Queen Char-
lotte Islands, British America.” At the same time SMALL recognizes Saxi-
fraga stellaris Brunoniana Bong. as a distinct species under the name Spatu-
laria Brunomiana (Bong.) Small, citing in addition as synonyms Saxifraga
leucanthemifolia Brunoniana Engler (!), Saxifraga notkana Moc., and
Saxifraga Bongardi Presl. Sma. also recognizes as distinct Saxifraga
ferruginea of Graham, under the name Spatularia ferruginea (Graham)
Small, giving its distribution as “Alaska to British Columbia,” and adjoining
as a synonym Hexaphoma ferruginea Raf. Fl. Tell. 2:66. 1836. Smatt’s
descriptions of these four species fail to show any sound and stable char-
acters by which they may be recognized, and the single characters, made use
of in his key to separate the species, are fluctuating and unreliable. Spatu-
laria Vreelandu and S. Brunoniana, for example, (along with S. bryophora
and S. foliolosa) are distinguished from S. Newcombei, and S. ferruginea
(along with S. stellaris and S. petiolaris (S. leucanthemifolia)) by the fact
that their inflorescences are bulblet-bearing, the inflorescences of the latter
being without bulblets. Further fine distinctions are made in regard to the
relative number of flowers and bulblets in distinguishing S. Brunoniana

JOHNSON: A REVISION OF THE SECTION BORAPHILA 61

from S. Vreelandw and in distinguishing S. Newcombei from S. fer-
rugimea by the relative length of their petals in millimetric denomina-
‘tions. While, as pointed out above, several forms are indicated as
occurring in, and limited to, certain regions of the entire range of our
species, they can not be separated on the characters mentioned in the preced-
ing lines. With large collections from several herbaria before me, covering
the entire range of the above-named species, it appears, if the collections are
grouped according to localities, that the bulblet-bearing character is common
_ to all, and that such great fluctuations in the form and size of the petals
occur, even in specimens from the same collection and on the same sheet,
as to make valueless any specific distinctions on these grounds. SMALL’s
species Spatularia Newcombe, of which a photograph only is available
(MBG 717014), and which, according to SMALL, is known only from the
type locality (Queen Charlotte Islands) has its homotype in Cowles’ 167,
from Glacier, British Columbia (MBG 744924), and in MacDougal’s 384,
from Kootenai County, Idaho (UMH); except for the few bulblets on the
last named, it is, in leaf character and pubescence, indistinguishable from
Spatularia Newcombei. The same applies to No. 451, Shaw’s Selkirk flora.
SMALL’s Spatularia Vreelandu is merely a small statured form of the plant,
homotypes of which occur in collections from all parts of the entire range.
On sheet No. 422334 USNH, collected by Culbertson at Windham Bay,
Alaska (Baker 4921), for instance, are two plants, one large and leafy-
bracted, with numerous large flowers and no bulblets and strikingly similar
in floral characteristics to SMALL’s S. Newcombei; the other small, (12 cm.
high) with all but the terminal flowers replaced by bracts, and corresponding
in every way with Smatvt’s description of S. Vreelandu. In fact the several
forms which have been described as species occur together constantly
throughout the entire range, often appearing together in collections from
various herbaria, on the same sheet. A good comparison of coast and
inland forms is found in Rosendahl’s No. 1809, from Cowichan Lake, Van-
couver Island, which is exactly similar to a specimen collected by Marcus
E. Jones at Sperry Glacier, Montana, at least in foliage characters. Accord-
ing to Dr. F. K. Butters, who has made a thorough and comprehensive
study of the Selkirk flora, both bulblet-bearing and non-bulblet-bearing
_ forms may be found repeatedly growing together, and he has observed that
the bulblet-bearing habit is commonly associated with altitude. Seeds of the
Selkirk plants planted by Dr. Butters in the greenhouse of the University
of Minnesota developed into plants which ran to bulblets entirely. Plants
collected by Butters and Rosendahl on the snowshed at Glacier, British
Columbia, differ markedly from others of the same locality in being tall,
excessively tufted, and profusely branched, with an abundance of flowers,
fruit, and glabrous leaves.

62 MINNESOTA STUDIES IN PLANT SCIENCE

A study of a great deal of material at hand has failed to disclose any
appreciable differences in the characters of those organs which are usually
regarded as showing the greatest constancy, namely fruit and seed. For
this reason all the various segregates which have been made are herein
regarded as one species. And the specific name ferruginea of Graham is
adopted, since it antedates others. The specific name Brunoniana, as used
by some writers, appears invalid, having a previous application to a species
of the section Trachyphyllum.

Those forms which exhibit constant peculiarities of form and habit,
correlated with a continuous range over a particular region, may be regarded
as varieties, and, as regards our species, at least two such varieties occur
continuously at the northern and the southern extremes of the range, with
a probable third in Washington and Oregon in the southern extreme. More
extensive collections and a more careful survey of the particular regions
are to be desired. A description of the varieties together with their ranges
follows:

Var. foliacea var. nov. Scapus aut solitarius aut caespitosus tenuis
ample ramosus grosse atque laxe glanduloso-pubescens ad medium bracteas
foliaceas gerens, foliis magnis ad basim cuneatis grosse serrato-dentatis.

Scapes solitary or tufted, erect or assurgent, commonly weak, coarsely
and loosely glandular-pubescent, often prolifically branched, with large leafy
bracts to above the middle, and numerous bulblets and flowers, the flowers
usually terminal. Leaves large, thin, broadly oblanceolate-elliptic, rounded
at the apex, coarsely serrate-dentate, the teeth few, cuneately narrowed
below into a broad or narrow base; glabrous or sparingly pubescent, con-
spicuously ciliate on the margins. The large leafy bracts and weak habit of
the plant are characteristic.

Distribution—High mountains of Kootenai County, and probably also Soshone
County, Idaho.

Specimens examined:

IpaHO: Palouse County and Lake Coeur d’Alene, August, 1892, J. H. Sandberg
(UMH) ; Packsaddle Mountain, Kootenai County, August 10, 1892, D. T. MacDougal,
No. 384, (UMH); crevices of ledges of porphyritic granite of Packsaddle Mountain,
alt. 8,000 ft., Lake Pend d’Oreille, July 12, 1889, J. B. Leiberg, No. 468 (UMH, MBG
84379) ; Lake Pend d’Oreille, alt. 3,000 ft., August, 1892, D. T. MacDougal (UMH) ;
Kootenai County, High Mountains, July, 1892, J. H. Sandberg (UMH); crevices of
rocks, summit of Packsaddle Peak, Kootenai County, August 5, 1892, Sandberg, Mac-
Dougal, and Heller, No. 834.

Var. grandiflora var. nov. Foliis ad 6 cm. longis ad basim cuneato-
contractis, floribus majoribus nonnumquam numerosis, bulbillis paucis,
inflorescentiae ramis diffusis.

Flowers unusually large, sometimes numerous; bulblets few. Scapes
rather stout, solitary or several together, coarsely and loosely pubescent,

ae JOHNSON: A REVISION OF THE SECTION BORAPHILA 63

branching above the middle, the branches spreading; leaves thin, oblanceo-
late-elliptic, glabrous or scantily pubescent, coarsely serrate-dentate above
the middle, cuneately contracted below into a broad or narrow base, 6 cm.
and less.

Distribution.—Alaska westward to Kodiak Island.

Specimens examined:

AtasKa: Windham Bay, June 11, 1905, C. F. Baker, No. 4921, J. D. Culbertson
(USNH 422334) Type; on rocky bluffs, shores of Behm Canal, 1894, M. W. Gorman
(UMH) ; Kodiak, July 3, 1899, Wm. Trelease, No. 4046 (MBG 84045).

Also the following peculiar form based on a single specimen:

Var. cuneata var. nov. Scapus 17 cm. altus, foliis ad basim latam
cuneato-contractis grosse serrato-dentatis, inflorescentiae ramis flores mag-
nos paucos atque bulbillos paucos gerentibus, fructibus crassis.

Leaves with elongated broad, flat, cuneate bases, slightly expanded above
into a short coarsely serrate-dentate, few-toothed blade; pale green, thinly
pubescent, margins ciliate, apex acute. Scape about 17 cm. high, stout,
branching into a few-flowered stout cymose inflorescence at the top. Bulb-
lets very few, flowers large, fruit stout, brownish. Filaments stout, rigid,
light-colored against the brown follicles.

Specimens examined:

Moresby Island, Queen Charlotte Islands, July 2, 1900, W. H. Osgood (USNH
620099) .

The following specimens are characterized by possessing elongated nar-
row leaves and a loose spreading inflorescence of slender branches.

Var. diffusa var. nov. Scapus 15-22 cm. altus, laxe pubescens supra
medium paniculato-cymose diffuse ramosus, foliis ad 9 cm. longis glabris
oblanceolatis grosse serrato-dentatis, floribus magnis, bulbillis paucis vel
nullis.

Leaves 9 cm. and less, narrow, glabrous, oblanceolate, acute, coarsely
serrate-dentate, the teeth few and remote, scapes 15-22 cm., solitary or
several together, assurgent, sparsely pubescent, diffusely branched above
the middle or from below the middle, the branches slender, spreading.
Inflorescence paniculate-cymose, open. Flowers rather few, large, with or
without bulblets.

Specimens examined:

. ALASKA: summit of high mountains near Yes Bay, August 21, 1895, Thos. Howe?t,
No. 1621 (UMH, MBG 84021).

The Minnesota sheet contains one small specimen, among two others,
which has small leaves of the usual ferruginea type. One of the speci-
mens, from the Missouri Botanical Gardens, possesses what appears to be
bulblets.

64 MINNESOTA STUDIES IN PLANT SCIENCE

The following collections from Mt. Hood and Mt. Adams approach
Saxifraga stellaris in their leaf characters: ;

Var. stellariformis var. nov.. Caulescens, scapo tenui, inflorescentiae
ramis filiformibus flores paucos atque bulbillos gerentibus, foliis grosse
paucidentatis sparsim pubescentibus ad basim attenuate contractis.

Thomas Howell, Mt. Hood, Oregon, August, 1881 (UMH) ; caulescent.
scapes slender, branches of inflorescence filiform, few-flowered, producing
flowers and bulblets. Leaves broad in proportion to length, thin, coarsely
few-toothed, or with a few apical teeth, sparsely pubescent, cuneately con-
tracted below into slender petiole-like bases, mostly equalling or exceeding
the blades.

Howell’s specimen is the nearest approach to S. stellaris of any plant
found in the western material examined, except for the form of the scape,
which appears to be typical ferruginea. A more extensive collection is
needed here.

Var. nivea var. nov. Scapus 5-10 cm. longus crassus, inflorescentia
flores magnos atque bulbillos gerens, foliis ad 1.5 longis ad scaporum basis
dense aggregatis apice grosse 2-4-dentatis glabris vel sparsim pubescentibus.

Snow-line, Mt. Adams, warm, moist, sandy slopes, August 10, 1882,
L. F. Henderson, No. 288, as S. stellaris (MBG 84387); scapes short,
stoutish, 5-10 cm., inflorescence bulblet and flower-bearing, the flowers
large in proportion to the size of the plants. Leaves short, 1.5 cm. and less,
crowded at the base of the scape, thick with 2-4 coarse apical teeth, glabrous
in one or two specimens, scantily pubescent in the others. The smaller
plants show a strong similarity to other small forms of S. ferruginea, how-
ever.

SAXIFRAGA FOLIOLOSA R. Br., Chlor. Melv. 17. 1823.

Saxifraga stellaris comosa Poir. in Lam. Encyc. 6:680. 1804. (Unveri-
fied.)

Saxifraga stellaris prolifera Cham., in Lit., ex Engler, Monogr. Saxifr.
133, 1872, as syn.

Saxifraga stellaris foliolosa Turez., Lac. Baic. 496.

Saxifraga comosa Britton, Mem. Torr. Bot. Club 5:178. 1894.

Spatularia foliolosa (R. Br.) Small, N. Am. FI. 22:2:158. 1905.

The leaves of the specimens examined are smaller than those of the
European specimens of Saxifraga stellaris L. The rigid erect scapes and
stiff branches of the inflorescence are quite characteristic of this species.

Only a single specimen of those examined possessed a terminal flower
(Plate XIX).

Distribution—According to Small: “Circumboreal, southward to the mountains
of Maine; also in Europe and Asia.” (/.c.). According to Engler fc.: “In der polar
region ziemlich verbreitet.”

JOHNSON: A REVISION OF THE SECTION BORAPHILA 65

Specimens examined:

Nugsuak Peninsula: “Collected by the Cornell Party on the Peary Voyage of
1896,” August 12 (UMH and MBG 84303).

St. George Island, July 31, 1891, Jas. AV. Mlacoun (MBG 84046).

SAXIFRAGA LEUCANTHEMIFOLIA Michx. Fl. Bor. Am. 1:268. 1803.
Spatularia leucanthemifolia Haw., Enum. Saxifr. 48. 1821.
Hexaphoma petiolaris Rat., Fl. Tell. 67. 1836.

Saxifraga Michauxu Britton, Mem. Torr. Bot. Club 4:118. 1894.
Spatularia Michaux (Britton) Small, Fl. S.E. U.S. 502. 1903.
Spatularia peticlaris (Raf.) Small, N. Am. FI. 22:2:150. 1905.

Distribution—Mountains, Maryland to Georgia.
Specimens examined:

TENNESSEE: Cades Coves, eastern Tennessee, August, 1899, Ff. H. Ferriss (MBG
83933).

MaryYLanpD: June 18, 1899, J. R. Churchill? (MBG 738695).

VIRGINIA: plants of Stony Man Mountain and vicinity, Blue Ridge, near Luray,
alt. 3.700 ft., September 4, 1901, F. S. and Mrs. Steele, No. 76 (UMH and MBG
83932) ; Bedford County, A. H. Curtiss (MBG 777207); peaks of Otter, Bedford
County, July 27, 1871, A. H. Curtiss (MBG 84235); Salt Pond Mountains, Giles
County, alt. 4,800 ft., August 19, 1875, John H. Redfield (MBG 83908); 4. F. E.
Creeson (MBG 84062) ; summit of White Top Mountain, Washington County, August
11-12, 1892, J. K. Small (MBG 83934).

NortH Caroitina: Statesville, M. E. Hyams (UMH); Roan Mountain, A. Ruth,
(UMH); Roan Mountain, alt. 6,000 ft., July 15, 1880, /. IW. Chickering; Grand-
father Mountain, Mitchell County, alt. 5,500-6,000 ft., August 25, 1893 (UMH and
MBG 83935); “In montibus Carolinae et Georgiae,’ Buckley (MBG 83911); on
rocks, Roan Mountain. alt. 6,100 ft., July, 4. H. Curtiss, No. 854 (UMH); Enos Plott,
Balsam Mountain, alt. 6,100 ft., August 28, and Roan Mountain, 6,400 ft., September
8, 1876, Redfield, Nos. 5500 and 5561 (MBG 83907); mountains, Leroy, Ruger, and
Redfield, No. 2005 (MBG 83906) ; Swain County, “Pinnacle Rock, Clingman’s Dome,”
alt. 6,660 ft., 1891; Beardslee and Kofoid (MBG 83910, 83896) ; Grandfather Mountain,
July 22, 1890, A. A. Heller, No. 51 (MBG 83904, UMH) ; Blowing Rock, July 19, 1890
(UMH) ; top of Roan Mountain, September 8, 1876, G. Engelmann (MBG 84236) ;
Cranberry Mountain, journey of September 22, 1898, Sargent, Muir, and Canby, No.
45 (MBG 83901) ; Grandfather Mountain, Mitchell County, July, 1895, herbarium of
Fred. Wislizenus (MBG 84579) ; Buncombe County, wet rocks near the summit of
Craggy Mountain, alt. 6,000 ft., July 8, 1897, Biltmore Herbarium, No. 1231d, (MBG
83898, UMH).

Section Heterisia (Raf.) sect. nov.

Heterisia Raf., Fl. Tell. 2:60. 1836; as genus.

Steiranisia, Raf., Fl. Tell. 2:69. 1836; as syn.

Hydatica, Torrey and Gray, Fl. N. Am. 1:567. 1838-1840; in part.
Hooker, Fl. Bor. Am. 1:252. 1840; in part. Ledebour. Fl. Ross. 2:216.
1844; in part.

Boraphila, Engler, Verh. Zool.-Bot. Ver., Wien. 19:522. 1869; in part.

66 MINNESOTA STUDIES IN PLANT SCIENCE

Ovary free; carpels united to the middle and surrounded by a deep,
thick, convoluted gland. Follicles pyriform, inflated below, strongly con-
tracted above into slender, divergent beaks. Filaments clavate. Sepals at
length reflexed. Seeds fusiform, faintly striate to smooth, slightly winged
by the loose seed-coat. Hairs, multiseriate (Plates II, XII).

Monotypic, comprising the following species:

SAXIFRAGA MERTENSIANA Bong., Mem. Acad. St. Petersb. VI:2:141.
1832.

Saxifraga heterantha Hook., Fl. Bor. Am. 1:252. 1832.

Steiranisia heterantha Raft., Fl. Tell. 2:62. 1836.

Heterisia Mertensiana (Bong.) Small, lc.

Heterisia Eastwoodiae Small, I.c.

Distribution—From Sonoma County, California, the Blue Mountains of south-
eastern Oregon, and the Bitter Root Mountains of Idaho and Montana to the region
of Yakutat Bay, Alaska.

Specimens examined:

AasKA: Sitka, June 17, 1899, Wm. Trelease, No. 4063 (MBG 83987); Sitka,
June 17, 1899, Wm. Trelease, No. 4063 (MBG 83988); Port Wells, Prince William
Sound, June 26, 1899, Wm. Trelease, No. 4064 (MBG 83989); at the base of Mount
Tebenkoff, Yakutat Bay, June 22, 1892, Frederick Funston, No. 41 (MBG 83990).

BritisH CoLuMBIA: stream sides, Prairie Hills, alt. 5,400 ft., July 26, 1907, But-
ters and Holway 235 (UMH); Avalanche Crest, Glacier, alt. 6,200 ft., July 28, 1907,
Butters and Holway, No. 235 (UMH); wet ledges above Cougar Valley, alt. 5,600 ft.,
July 18, 1908, Butters and Holway (UMH); along banks of streams, Cowichan
Alberni Trail, Vancouver Island, alt. 2,600 ft., June 26, 1906, C. O. Rosendahl (UMH) ;
Cameron River Valley, Vancouver Island, June 28, 1907, C. O. Rosendahl, No. 1991
(UMH and MBG 84054) ; alpine meadow in the Big Bend district, about 118° 20’ W.
long. 51° 45’ N. lat., alt. 7,500 ft., July 26, 1905, H. Shaw, No. 1016 (MBG 83919).

WaASHINGTON: north fork of Bridge Creek, Okanogan County, on moist mossy
flanks of cliffs, September, 1897, A. D. E. Elmer, No. 716 (UMH) ; Snoqualmie, June
4, 1889, E. C. Smith (MBG 83918); clefts of rocks, Goat Mountains, alt. 5,500 ft.,
August 12, 1896, O. D. Allen, No. 241 (MBG 83926); Olympic Mountains, Clallam
County, July, 1900, 4. D. E. Elmer, No. 2643 (UMH, MBG 83929) ; mossy rocks at
Quiniault Falls, June 21, 1902, H. S. Conard, No. 390 (UMH); Baldy Peak, Chehalis
County, alt. 3,500 ft., July 21, 1897, H. Lamb, No. 1348 (MBG 83927).

IpaHo: ledges, Kootenai County, June, 1892, J. B. Letberg, No. 467 (UMH) ;
divide between St. Joseph and Clearwater River, alt. 1,600 m. July 9, 1895, J. B. Le#-
berg, No. 1206 (MBG 84081) ; Palouse County and about Lake Coeur d’Alene during
June and July, 1892, and May, 1892, J. H. Sandberg (UMH); Nez Perces County,
moist hillsides, May, 1892, NSS, No. 3170 (UMH, MBG 83930, 83924); mountain
slopes, Kootenai County, July, 1887, J. H. Sandberg (UMH).

Orecon: 1871, Hall, No. 152 (MBG 84225); 1871, Hall, No. 152 (MBG 83921) ;
wet rocks on the Wallowa Mountains, September 1, 1899, W. C. Cusick, No. 2327
(UMH, MBG 83925); Bingham Springs, western: slope of the Blue Mountains, alt.
766 m., June 12, 1908, and 3321, wet granite cliffs, Wallowa Mountains, alt. 3,000 m.
August 17, 1908, W. C. Cusick, No. 3330, (MBG 84049) (both on same sheet) ;

JOHNSON: A REVISION OF THE SECTION BORAPHILA 67

Oneonta, June, 1886, Thos. Howell (UMH); left fork of Wallowa River, Wallowa
Mountains, alt. 6,000 ft., July 12, 1897, E. P. Sheldon, No. 8548 (MBG 84072).

Catirornia: Gasquet, April 20, 1892, Thos. Howell, No. 1428 (MBG 83931, UMH) ;
Ukiah, 1866, H. N. Bolander, No. 4652 (MBG 84224) ; Mill Creek, Mendocino County,
1866, H. N. Bolander, No. 4652 (MBG 83928) ; Emigrant Gap, July 1, 1882, M. E.
Jones, No. 3588 (MBG 810802) ; along Little River, Humboldt County five miles from
the coast, alt. about 400 ft., June 4, 1911, P. Tracy, No. 3217 (MBG 84598) ; woods
near Cazadero, Sonoma County, April 24, 1903, 4. A. Heller, No. 6616 (MBG 83923),

Montana: Darby, upper temperate life zone, alpine, alt. 7,500 ft., July 30, 1909,
M. E. Jones (MBG 84959).

SPECIES INCERTAE SEDIS

MICRANTHES APETALA (Piper) Small
MICRANTHES BIDENS Small a
MICRANTHES BRACHYPUS Small
MICRANTHES CRENATIFOLIA Small
MICRANTHES ERIOPHORA (S. Wats.) Small
MICRANTHES GALACIFOLIA Small
MICRANTHES GREENET Blankinship
MIcCRANTHES KUMLIENII Small
MICRANTHES LATA Small

MICRANTHES MONTANA Small
MICRANTHES REFLEXA (Hook.) Small
MICRANTHES RYDBERGII Small
MICRANTHES TENUIS (Wahl.) Small
MICRANTHES YUKONENSIS Small
SAXIFRAGA GASPENSIS Fernald

No specimens of the above species have been available for study, and as
the descriptions of the authors deal principally with vegetative characters
their status can not be definitely determined at this time.

Micranthes eriophora (S. Wats.) Small appears to be very different
from any species described in this paper, but the specimen at hand is in
such poor condition that no definite conclusions can be drawn from it.

Saxifraga gaspensis Fernald was separated by its author from Saxi-
fraga nivalis. The distinguishing characters as pointed out by FERNALD
appear, however, to be wholly vegetative.

Micranthes apetala (Piper) Small is probably a form of Saxifraga
nidifica Greene.

This monograph was undertaken at the suggestion of Dr. C. O. Rosendahl, and
has been carried out under his direction. The writer was privileged to use the large
collection of Saxifragas in the Herbarium of the University of Minnesota, and other
large and valuable collections kindly loaned by other herbaria.

The writer desires to express his thanks to Dr. Rosendahl for much needed advice,
for his efforts in securing valuable herbarium materials, and providing ample facilities
for carrying on the work; to Dr. Frederic K. Butters for much practical advice, help-
ful criticism, and information about the Saxifraga flora of the Selkirks and the

68 MINNESOTA STUDIES IN PLANT SCIENCE

Canadian Rocky Mountains; to Dr. William S. Cooper for helpful suggestions touch-
ing ecological factors; to Dr. Oscar W. Oestlund for many inspiring ideas and guiding
principles drawn from his fund of knowledge of the science and philosophy of tax-
onomy. Especial thanks are due to Dr. W. R. Maxon, of the United States National
Herbarium, for the loan of indispensable herbarium material; likewise to Dr. George
T. Moore and Dr. J. M. Greenman for the loan of a large collection from the Her-
barium of the Missouri Botanical Garden. The writer wishes also to express his
thanks to Dr. H. M. Hall for specimens from the Herbarium of the University of
California; to Mr. F. S. Hall for specimens from the Herbarium of the State Museum,
University of Washington, Seattle; to Dr. F. L. Pickett, Washington State College,
Pullman, Washington, for specimens from that region; to Dr. J. E. Kirkwood, Uni-
versity of Montana, Missoula, Montana, for specimens from the Rocky Mountain
region; and to Mr. Alonzo P. Troth, of the Lewis and Clark High School, Spokane,
Washington, for fresh material from the Spokane Valley.

Owing to conditions resulting from the World War Engler’s monograph on
Saxifraga in Das Pflanzenreich (1V :117:1. 1916, 1V:117:II. 1919) was not obtain-
able when this paper was in process of preparation—AvTHor.

University of Minnesota, 1919.

LITERATURE CITED

. Britton, N. L. and Brown, A. Illustrated flora of the northern United States,

Canada, and the British Possessions. Vol. 2, page 174. 1897.

. Covirte, F. V. Proc. Biol. Soc. Washington 7:78. 1893.
. Don, D. Monograph of the genus Saxifraga. Trans. Linn. Soc. 13. 1822.
. ENcLER, A. Index criticus specierum atque synonymorum generis Saxifraga.

Verh. Zool.-Bot. Ver., Wien. Vol. 19. 1869.
—. Monographie der Gattung Saxifraga. Breslau. 1872.
Saxifragaceae, in Die Natiirlichen Pflanzen Familien 3:2a:41-93. 1891.
, and IrmMscHER. Manuscript notes on sheets of the United States National
Herbarium.

. FERNALD, M. L. New species, varieties, and forms of Saxifraga. Rhodora 19:141.

1917.

. Gray, Asa. Notes on a botanical excursion to the mountains of South Carolina.

London Journ. of Bot. 2:113. 1843.
Chloris Boreali Americana. Mem. Am. Acad. 3:39. 1846.

. GREENE, E. L. Novitates Occidentalis. Erythea 1:222. 1893.
. Gopron, D. A. Flore de France. 1848-56.
. HaworrH, A. H. Synopsis plantarum succulentarum. 1812.

Saxifragarum enumeratio. 1821.

. HarsHBERGER, J. W. Phytogeographic survey of North America. 1911.

. Hooxer, W. J. Flora Bocreali-Americana. Vol. 1, pages 242-55. 1840.

. Howett, THomas. New species of Pacific Coast plants. Erythea 8. 1895.

. LepEBour, C. F. Flora Rossica. Vol. 2, pages 204-25. Stuttgart. 1844.

. Micuaux, A. Flora Boreali Americana. Vol. 1, page 269, 1803.

. Patras, P. S. Reise durch verschiedene Provinzen Russischen Reich, 3. 1878.

Pirrer, C. V. Flora of Washington, Contrib. U. S. Natl. Herb. 11. 1906.
—. New and noteworthy northwest plants IV. Bull. Torr. Bot. Club
27 :393. 1900.

. PursH, Frepertck. Flora Americae Septentrionalis. Vol. 1, page 311. 1814.
. Rarinesoug, C. S. Flora Telluriana 2:68. 1836.
. RosENDAHL, C. O. Die nordamerikanischen Saxifraginae u. ihre Verwantschafts

Verhaltnisse in bez. ihrer geo. Verbreitung. Beibl. Engl. Bot. Jahrb. 83. 1905.

. SERINGE, N. C. Saxifraga in DC. Prod. 4:17-47, 1839.
. SMALL, J. K. Saxifragaceae in North American Flora 22 :2 :126-56.

-——-——. Saxifragaceae in Britton and Brown, Illustrated flora. Second edition.
Vol. 2, pages 214-30. 1913.

, and Varit, ANNA Murray. Report of a botanical exploration of south-
western Virginia. Mem. Torr. Bot. Club 5:118. 1893.

. Tauscu, I. F. Hortus Canalius. fase. 1. 1823.
. Torrey and Gray. Flora of North America. Vol. 1, pages 563-76. 1838-1840.
32. WILLDENow, C. L. Linnaeus Sp. Pl. Fourth edition. Vol. 2, pages 639-58. 1799.

_ EXPLANATION OF PLATES

72 MINNESOTA STUDIES IN PLANT SCIENCE

Piate I

A, B, section Hydatica. A, capsule in the subsection Rotundifoliatae; B, follicles
in the subsection Cuneifoliatae; a, b, young stages of A; c, young stages of B; gl,
gland; A, d, e, types of hairs in the section Hydatica; C, follicles of the section Cal-
thophyllum; f, young stage of same; gl, gland; g, type of hair in Calthophyllum; D,
capsule of the section Aulaxis; h, types of hairs in Aularis; k, young stage of pistil
of Aulaxis; gl, gland; E, type of carpels in the section Dermasea; i, 7, young stages of
the same; gl, gland; J, type of hair in the section Dermasea.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 73

74 MINNESOTA STUDIES IN PLANT SCIENCE

PrateE II

F, follicles of the section Chionophila; m, young stage of same, dorsal aspect; gl,
gland; G, follicles of the section Micranthes; n, young stage of same; gl, gland; fi,
flange of the gland; H, follicles of the section Tricarpum; o, young stage of same;
gl, gland; ff, flange of gland; p, p, p, types of hairs in sections Chionophila, Micran-
thes, and Tricarpum; I, capsule of the section Arabisa; s, young stage of same; gl,
gland; J, capsule of the section Heterisia; r, young stage of same; gl, gland; q, type
of hair in sections Arabisa and Heterisia.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 75

Piate II

76 MINNESOTA STUDIES IN PLANT SCIENCE

PLaTE III

Ventral aspects of the carpels of the sections in which the receptacle is more or
less adnate to the ovary (in all cases the gland is shaded black) : section Micranthes:
1, S. californica; 2, S. fragosa; 3, S. pennsylvanica; 4, S. montanensis; 1'-4', carpels
of the same seen from above, showing nature of the gland in each; section Chiono-
phila; 5, S. rhomboidea; 5’, young stage of same seen from above, showing gland;
¥, point of connection of carpels; section Tricarpum: 6, S. texana; 6’, young stage of
same seen from above; +, line of connection of carpels (shaded in line); section
Dermasea: 7, young stage of carpel of S. caroliniana; 8, 8’, old and young stages of
carpels of S. rufidula; 9, 9’, the same of S. occidentalis; 10, carpel of S. idahoensis;
11, 11’, early stages of carpels of S. virginiensis. Figures 7 and 11 also show the
variations in the depth of the receptacle in species of the section Dermasea.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 77

Q

PiateE III

78 MINNESOTA STUDIES IN PLANT SCIENCE

Piate LV*

I. Section Hydatica. capsule of S. arguta; 1, petal; 2, sepal; 3, filament; 4, seed;
5, hair and its gland; 6, leaf; drawn from a specimen collected by S. G. Stokes in the
vicinity of Clayton Peak, Wasatch Mountains, Utah (MBG 84500).

II. Immature capsule of S. aestivalis; 7, petal; 8, sepal; 9, filament; 10, non-gland-
bearing hair; 11, seed; gl, gland; Ila, pistil, showing bottle-shaped form of young
carpels, typical of the section; drawn from Piper’s No. 2622, from Mount Rainier,
Washington (UMH).

III. Mature capsule of 5. Nelsontana; 12, sepal; 13, glands of hairs; 14, seed;
drawn from Jas. M. Macoun’s No. 23i, from Copper Islands, Bering Sea (USNH
249366).

IV. Mature capsule of S. unalaschensis; 15, filament; 16, petal; 17, sepal; 18, seed:
- drawn from a specimen collected by Trevor Kincaid on Popoff Island, Alaska (USNH
375422). ;

V. Mature capsule S. davurica; 19, sepal; 20, filament; 21, leaf; 22, seed; drawn
from a specimen collected by Chester Washburn, near Cape Sabine, Alaska (USNH
421209). ;

VI. Mature capsule of S. Lyallii; 23, petal; 24, sepal; 25, filament; 26, hair and
gland; 27, seed; drawn from Butters and Holway’s No. 818, from the Cougar Valley,
Selkirk Mountains, British Columbia (UMH).

*The small numbers associated with figures of organs refer to the length in millimeters,
natural size.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 79

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PiaTE ITV

80 MINNESOTA STUDIES IN PLANT SCIENCE

PLate V

Section Calthophyllum. I, mature follicles of S. spicata; 1, petal; 2, sepals; 3,
stamen and filaments; 4, leaf; 5, hair; 6, immature seeds; II, pistil, showing shape of
the young carpel, with broad base; III, ventral side of young carpel, with adherent
receptacle; I, drawn from a specimen collected by Dr. J. M. Coulter at Cape Van-
couver (USNH 249391), the others drawn from a specimen collected by W. C. Men-
denhall in the Valley of Alatna, Alaska (USNH 377352).

Section Aulaxis. IV. Flower of S. micranthidifolia; V, mature capsule; VI,
immature carpels; VII, pistil with swollen gland at base; 7, petals; 8, sepals; 9,
stamen and filament; 10, leaf; 11, seeds; 12, a seed much enlarged; 13, hairs and their
glands; drawn from Small and Heller’s No. 246, from Grandfather Mountain (UMH).

81

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JOHNSON: A REVISION OF THE SECTION BORAPHILA

PLaTeE V

82 MINNESOTA STUDIES IN PLANT SCIENCE

PLaTE VI

Section Dermasea. I, mature follicles of S. rufidula; II, lateral; III, ventral; IV,
dorsal, views of a single follicle, the latter showing the shallow adherent receptacle;
1, petals; 2, sepals; 3, filaments; 4, seeds; 5, a bract of the inflorescence; 6, leaf; 7,
receptacle and calyx; drawn from Dr. C. O. Rosendahl’s No. 1631, from Mount Arrow-
smith, Vancouver Island, British Columbia (UMH).

V. Dorsal and ventral views of immature carpels of S. laevicaulis; 8, petals; 9,
sepals; 10, stamen and filament; 11, pistil of three carpels; drawn from C. V. Piper’s
No. 5061, from Mount Grayback, Oregon (UMH).

VI. Flower of S. aequidentata; VII, follicles; VIII, more mature follicles; 12,
petals; 13, sepals; 14, filament; 15, receptacle and calyx; 16, hairs; drawn from Suks-
dorf’s No. 967, from Skamania County, Washington (UMH); IX and X, immature
carpels, showing gland at base; XI, pistil of three carpels; XII, ventral side of young
carpel showing depth of adherent receptacle; 17, petals; 18, sepals; 19, filaments; 20,
seed; 21, a bract of the inflorescence; 22, leaves; 23, gland of hair; drawn from a
specimen collected by Thomas Howell at Oregon City (UMH).

JOHNSON: A REVISION OF THE SECTION BORAPHILA

Piate VI

84 MINNESOTA STUDIES IN PLANT SCIENCE

PLATE VII

I. Immature follicles of S. Allenii; 11, dorsal and ventral views of a young carpel,
showing gland and shallow receptacle; 1, petals; 2, sepals; 3, stamen; 4, bracts of the
inflorescence; 5, gland of hair; drawn from H. D. Langille’s No. 37, from Mount
Hood, Oregon (USNH 525065).

III. Immature follicles of S, Marshall; IV, young carpels showing gland at base,
and dorsal rib; 6, petals; 7, sepals; 8, stamens; 9, hairs and their glands; drawn from
Harley P. Chandler’s No. 1613, from Marble Mountain, Siskiyou County, California
(MBG 83016, UMH).

V. Follicles of S. Careyana; VI, ventral view of same; VII, dorsal and ventral
view of young carpel showing gland and shallow receptacle; 10, petals; 11, sepals;
12, filaments; 13, glands of hairs; drawn from John H. Redfield’s No. 11721, from
Roanoke Mountain, North Carolina (MBG 84285).

VIII. Follicles of S. tennesseensis; TX, dorsal side of an immature ‘follicle show-
ing region of the gland; 14, petals: 15, sepals; 16, filament; 17, seed; 18, glands of

hairs; drawn from a specimen collected by A. Ruth at Knoxville, Tennessee (MBG
84398).

A REVISION OF THE SECTION BORAFPHILA 85

JOHNSON:

Priate VII

86 MINNESOTA STUDIES IN PLANT SCIENCE

Prate VIII

I. A flower of S. caroliniana; II, mature follicles of same; III, immature follicles ;
IV, ventral side of a young carpel showing shallow adherent receptacle; V, early
stage of the carpels; VI, ventral view of a mature follicle; 1, petals; 2, sepals; 3,
stamen; 4, leaf; 5, seeds at left immature; 6, glands of hairs; 7, a bract of the
inflorescence; I, drawn from a specimen in the Herbarium of the University of Minne-
sota; II and VI, drawn from a specimen collected by Dr. Asa Gray, C S. Sargent,
J. H. Redfield, and William M. Canby on Negro Mountain, North Carolina (MBG
84291), III, IV, and V from a specimen, collected by J. D. Small at Chatham Hill Gap,
Virginia (MBG 84288).

VII. Mature follicles of S. tdahoensis; VIII, dorsal; IX, ventral sides of young
carpels; X, a very young carpel showing gland; 8, petals; 9, sepals; 10, stamen; 11,
leaves; 12, bract of inflorescence; 13, glands of hairs; VII and 11 drawn from J. H.
Sandberg’s collection from the Palouse County, Washington, the others from Aiton’s
collection from Kootenai County, Idaho (UMH).

XII. Mature follicles of S. petiolata; XIII, ventral view of a mature follicle,
showing the slightly adherent receptacle; 14, petals; 15, sepals; 16, filament; 17, seeds;
18, a seed much enlarged; 19, bract of inflorescence; 20, glands of hairs; 21, side view
of mature fruit.

87

JOHNSON: A REVISION OF THE SECTION BORAPHILA

Piate VIII

88 MINNESOTA STUDIES IN PLANT SCIENCE

PLate [X

I. Follicles of §. Hallii; II, dorsal; III, ventral views of immature follicles; IV,
ventral side of a young carpel; 1, petals; 2, sepals; 3, stamen and a filament showing
variation; 4, bracts of the inflorescence; 5, glands of the hairs; drawn from Elihu
Hall’s No. 151, Oregon (MBG 84380).

V. Follicles of S. microcarpa; VI, dorsal; VII, ventral views of a single follicle;
6, petals; 7, sepals; 8, stamen and a developed filament; 9, bract of the inflorescence;
10, glands of the hairs; drawn from M. J. Elrod’s No. 98, from Missoula, Montana,
Herbarium of the University of Montana.

VIII. Immature follicles of S. klickttatensis; IX, ventral; X, dorsal sides of a
single follicle; 11, petals; 12, sepals; 13, stamen; 14, bract of inflorescence; 15, hairs
and their glands; drawn from a specimen collected by W. N. Suksdorf in West
Klickitat County, Washington (USNH 45806).

XI. Flower of S. virginiensis, with parts cut away to show young carpels and
gland; 16, stamens at different stages of development; drawn from J. H. Sandberg’s
collection from Two Harbors, Minnesota; 17, 18, and 19, various forms of petals
found in different collections of S. virginiensts; 17, from Milford, Pennsylvania, C. A.
Mill, collector (MBG 84374); XIV, XV, XVI, young carpels of same; 18, from
Sandberg’s collection from Two Harbors, Minnesota; 19, from Southington, Con-
necticut, C. H. Bissell, collector, No. 272 (and 813) (MBG 84364); 20, bracts of
inflorescence; 21, glands of hairs from different specimens; 22, seeds; XII, two
carpels with the parts of flower cut away; XIII, ventral side of single carpel showing
depth of the adherent receptacle; XII-XVI, and 22, drawn from Dr. J. M. Green-
man’s No. 629, from Stony Brook, Massachusetts (MBG 738779).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 89

90 MINNESOTA STUDIES IN PLANT SCIENCE

PLATE X

I. Follicles of S. nivalis, showing gland and deep adherent receptacle; II, young
carpels; III, dorsal; IV, ventral sides of single carpel; 1, petals; 2, sepals; 3, fila-
ments; 4, seeds; 5, glands of the hairs; drawn from Karl Hedbom’s collection from
the Dovre Fjeld, Norway (UMH).

V. Immature follicles of 5. saximontana; VI, dorsal and ventral sides of a single
carpel; VII, dorsal side of a young carpel; 7, petals; 8, sepals; 9, stamens; 10, seeds
(the lower much enlarged) ; 11, glands of the hairs; drawn from Sheldon’s No. 8536,
from the Wallowa Mountains, Oregon (USNH 528321) ; 12, petals; 13, sepal; from a
specimen collected by J. H. Sandberg in Kootenai County, Idaho (UMH) ; 14, stamen;
15, gland of hair; 16, bract of inflorescence; drawn from A. and E. Nelson’s No. 5917,
Yellowstone Park, Wyoming (UMH).

VIII. Follicles of S. occidentalis; drawn from a specimen collected by W. N.
Cusick in the Powder River Mountains, Oregon (UMH); IX, ventral view of a
mature carpel; X and XI, young carpels; 17, petals; 18, sepals; 19, seeds; 20, stamens:
21, bracts of the inflorescence; 22, glands of the hairs; drawn from Elmer’s No. 2646,
from the Olympic Mountains, Clallam County, Washington (USNH 401969).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 91

1
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PEATE OX

92 MINNESOTA STUDIES IN PLANT SCIENCE

PLaTE XI

Section Chionophila. I, follicles of S. rhomboidea; II, ventral view of a follicle
of same; 1, petals; 2, sepals; 3, stamen; 4, hairs and their glands; 5, bract of the
inflorescence; drawn from J, W. Blankinship’s No. 183, from Flathead Pass, Montana
(MBG 84114); III, dorsal; IV, ventral views of carpels of another specimen of
Blankinship’s No. 183; 6, petals; 7, sepals; 8, stamen; 9, hairs and their glands; from
Blankinship’s No. 183 (MBG 84114); 10, a type of gland commonly found on the
hairs of S. rhomboidea.

V. A flower of S. hieracifolia; drawn from Dr. Wm. Trelease’s No. 4035, from
Hall Island, Alaska (MBG 83975) ; VI, follicles; VII, ventral view of a follicle; 11,
petals; 12, sepals; 13, seeds; 14, glands of hairs; drawn from Jas. M. Macoun’s No.
55, from St. Matthew Island, Bering Sea (MBG 83978).

VIII. Follicles of S. franciscana; IX, ventral; X, lateral views of a single
follicle; 15 and 16, types of petals occurring in S. franciscana; 17, receptacle and
calyx; 18, seed; 19, hairs and their glands; drawn from F. H. Knowlton’s No. 127,
from the San Francisco Mountains, Arizona (USNH 258261).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 93

PLaTE XI

Od. MINNESOTA STUDIES IN PLANT SCIENCE

Piate XII

Section Micranthes. I, follicles of S. Forbesti; 1, seeds; from MBG S582; II and
III, ventral views of mature follicles; II, from MBG 83820; III, from MBG 775263;
IV and V, dorsal and lateral views of carpels; IV, from MBG 83859; V, from MBG
83862; VI, dorsal; VII, ventral views of young carpels; 2, petals; 3, sepals; 4, stamen
and filaments drawn from Vasey’s specimen collected in Illinois, June, 1875 (MBG
84223) ; 5, seed; drawn from a specimen collected by Eggert near St. Louis, Missouri
(MBG 84480).

VIIL. Follicles of S. purpuripetala; 6, petal; 7, seeds of same; drawn from a
specimen collected by Ferdinand Blanchard at Peacham, Vermont (MBG 751775) ;
IX, dorsal view of a mature carpel of S. purpuripetala; 8, two petals; 9, speals; 10,
stamens; drawn from MBG 751771, Ferdinand Blanchard, Peacham, Vermont.

X. Follicles of S. pennsylvanica; XI, ventral view of a mature follicle; XII,
dorsal view of a young carpel; 12, petals; 13, sepals; 14, a seed; drawn from a speci-
men collected by G. P. Ashcroft, Munson, Indiana (UMH) ; XIII, mature follicles; 15,
a seed; drawn from a specimen collected by C. H. Bissell, No. 69, near Southington,
Connecticut (MBG 84487); XIV, ventral view of an immature follicle; 16, a petal;
drawn from a specimen collected by Thomas Morong near Cambridge, Massachusetts
(MBG 84488) ; XV, ventral view of young carpel; 17, three petals; 18, a sepal; 19 and
20, seeds from different specimens of S. pennsylvanica; 19 (MBG 84473), Redfield No.
2010 (MBG 84481) ; 21, common types of glands of hairs occurring in S. pennsyl-
vanica.

XVI. Follicles of S. crassicarpa; XVII, dorsal view of a mature follicle (drawn
on reduced scale) ; 22, pétal; 23, a sepal; 24, a filament; drawn from a specimen col-
lected by H. Eggert in Wisconsin (MBG 83863); XVIII, XIX, dorsal views of
two young carpels; XX, a more advanced carpel of same; 25, a petal; 26, a sepal;
27, a stamen; 28, a bract of inflorescence; 29, gland of hair; drawn from a specimen
collected by Virginius H. Chase near Wady Petra, Illinois (MBG 84469).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 95

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Pirate XII

96 MINNESOTA STUDIES IN PLANT. SCIENCE

PLate XIII

I. Follicles of S. winnebagoensis; II, ventral; III, dorsal views of a mature
follicle; 1, sepals; 2, a filament; 3, seeds; 4, glands of hairs; drawn from W. A.
Wheeler’s No. 180, from the Winnebago Valley, Minnesota (UMH).

IV. A flower of S. oregana; V, follicles; VI, ventral view of a mature follicle;
VII and VIII, very young carpels; 5, petals; 6, sepals; 7, a stamen; 8, seeds (drawn
to different scales) ; 9, bract of inflorescence; 10, glands of hairs; drawn from Thomas
Howell’s No, 1498, from Lake Labish, Oregon (UMH).

IX. Follicles of S. Sierrae; X, dorsal; XI, ventral views of mature follicles : XII,
young carpels showing gland; 11, petals; 12, sepals; 13, filament; 14, seeds; 15, glands
of hairs; drawn from Mrs. R. M. Austin’s No. 83, from Colby, Butte County, Cali-
fornia (UMH).

XIII. Advanced stage of pistil; XIV, ventral view of carpel; 16, petals; 17, a
sepal; 18, glands of hairs; drawn from a specimen of Baker’s type collection of S.
arnoglossa, from Marshall Pass, Colorado (UMH); 19, a petal; 20, a sepal; from
Baker’s No. 4635; 21, two seeds; drawn from a specimen of S. arnoglossa; from
Kenneth D. Mackenzie’s No. 222, Colorado (MBG 83876).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 97

v.09 ci ih

yes os)

Puate XIII

98 MINNESOTA STUDIES IN PLANT SCIENCE

PLateE XIV

I. Dorsal; II, ventral views of mature follicles of S. parvifolia; III, two young
carpels of same, from above and side; 1, petals; 2, sepals; 3, stamen and filaments ;
4, glands of hairs; drawn from Thomas Howell’s No. 1134, from Grant’s Pass, Ore-
gon (MBG 84381).

IV. Follicles of S. gracillima; V, ventral view of a mature follicle; VI, same of
young carpel; VII, two young carpels, seen from above, showing gland; 6, petals; 7,
sepals; 8, bracts of inflorescence; 9, glands of hairs; drawn from A. A. Heller’s No.
11214, from Butte County, California (MBG 745924).

VIII. Immature follicles of S. fallax; IX, ventral view of a follicle; X, lateral
view of same; 10, petals; 11, sepals; 12, filament; 13, seeds; 14, seed enlarged; 15,
hairs and their glands; drawn from Marcus E. Jones’ No. 3587, from Emigrant Gap,
California (USNH 221361).

XI. A flower of S. californica; drawn from Pitt’s No. 286, from Mount Tamal-
pais, California (UMH) ; XII, mature follicles; XIII, dorsal; XIV, ventral views of
a carpel; 16, petal; 17, sepal; drawn from Baker’s No. 370, from Santa Clara County,
California (MBG 84275); 18, petals; 19, glands of hairs; drawn from Mrs. R. M.
Austin’s No. 859, California (MBG 84281) ; 20, stamens; 21, bracts of the inflorescence:
22, diagram of a branchlet of the inflorescence; from H. E. Brown’s No. 150, from
Butte County, California; 23, a seed enlarged; 24, glands of hairs.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 99

PLaTE XIV

100 MINNESOTA STUDIES IN PLANT SCIENCE

Pee XV

S. nidifica: I and II, two flowers of the same collection of “S. plantagineo”
Small; 1 and 3, petals and sepals of I; 2 and 4, the same of II; drawn from the type
collection of Sandberg and Leiberg, from Spokane, Washington, May, 1893; 5, varia-
tions in petals of different flowers of the same plant; drawn from a specimen of a
collection furnished by Mr. Alonzo P. Troth, of the Lewis and Clark High School,
Spokane, Washington; III and IV, two different types of mature follicles in Piper’s
S. columbiana; V, dorsal view of a single follicle of III; 6, two petals of same; 7
and 8, mature and immature seeds; 9, glands of hairs; VI, a flower showing both
narrow and broad petals in the same flower (compare with I and II); drawn from
Piper’s No. 1808, from Pullman, Washington; VII, a flower of Suksdorf’s S$. brac-
teosa; VIII, dorsal; IX, ventral views of mature follicles of same (compare with III,
IV, V); 10, seeds; 11, glands of hairs of same; drawn from Suksdorf’s No. 1728,
from the Columbia River, southwestern Washington (UMH); X, a flower of Suks-
dorf’s No. 4014, S. bracteosa var. leptopetala, from West Klickitat County, Washington
(UMH) ; XI, a young carpel from a flower of I, showing prominent verrucose gland.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 101

PLATE XV

102 MINNESOTA STUDIES IN PLANT SCIENCE

Pirate XVI

I. Pistil of S. montanensis; II, dorsal view of a young carpel; 1, a petal; 2,
sepals; 3, a filament; 4, glands of hairs; g, gland of carpels; drawn from “E. C. S,’s”
No. 155, from Electric Peak, Yellowstone National Park (USNH 488735); III,
dorsal; IV, ventral views of a carpel; 5, a petal; 6, sepals; 7, filaments; 8, glands of
hairs; g, gland of carpel; I, a pistil; drawn from another specimen of the same col-
lection, (E. C. S. No. 155) ; V, dorsal; VI, ventral views of mature follicles; 9, petals;
10, sepals; 11, a filament; 12, seeds; 13, glands of hairs; drawn from Rydberg and
Bessey’s No. 4264, from Spanish Basin, Gallatin County, Montana (USNH 360878).

Vil. Dorsal view of a mature follicle of a specimen referred to S. integrsfolha
by Professor Joseph Marsh; VIII, a young carpel; IX, a more advanced stage of
young carpel of same; 14, petals; 15, sepals; 16, glands of hairs; drawn from Marsh’s
No. 85, University of Washington Herbarium; X, dorso-lateral view of an immature
follicle; XI, carpels at opening of flower; 17, petals; 18, sepals; 19, stamen; 20,
glands of hairs; drawn from a specimen from Whidby Island, Washington, referred
to S. integrifolia by N. L. Gardner, University of Washington Herbarium.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 103

a

8

PuiaTE XVI

104 MINNESOTA STUDIES IN PLANT SCIENCE

Piate XVII

I. A flower of S. fragosa; II, follicles; III, ventral; IV, dorsal views of imma-
ture follicles; V, ventral views of a very young carpel, showing gland g; 1, petals;
2, sepals; 3, filament; 4, seeds, the lowest much enlarged; 5, glands of hairs; drawn
from Suksdorf’s No. 1427, from Klickitat County, Washington (UMH); VI, dorsal;
VII, ventral views of mature follicles; 6, petals; 7, sepals; 8, stamen; 9, glands of
hairs; drawn from Sheldon’s No. 10109, from the Dalles, Oregon (UMH); VIII,
lateral view of a mature; IX, dorsal view of an immature follicle; 10, a bract of the
inflorescence; drawn from Elmer’s No. 139, Almota, Whitman County, Washington
(UMH).

X. A flower of S. napensis; XI, young carpels; XII, dorsal view of an immature
follicle; XIII, ventral view of same; 11, a filament; 12, glands of the hairs; drawn
from Hall and Chandler’s No. 386, from the region of Dinky Creek, Fresno County,
California (UMH).

XIV. Immature follicles of S. laevicarpa; XV, ventral; XVI, dorsal views of a
follicle; 13, petals; 14, sepals; 15, glands of hairs; drawn from a specimen collected
by William Moody at Friday Harbor, Washington, University of Washington Her-
barium.

JOHNSON: A REVISION OF THE SECTION BORAPHILA 105

E/

Pirate XVII

106 MINNESOTA STUDIES IN PLANT SCIENCE

Prate XVIII

Section Tricarpum. I, capsule of S. texana; II, dorsal view of an immature
carpel of same; 1, sepals; 2, petals; 3, stamen and filaments; 4, seeds, hairs, and their
glands; drawn from Ernest J. Palmer’s No. 1647, from Reding’s Mill, Newton
County, Missouri (MBG 760519) ; III, capsule; 6, sepals; 7, immature seed; 8, bracts
of inflorescence; 9, stigma enlarged; drawn from B. F. Bush’s No. 5475, from Red-
ing’s Mill, Missouri (MBG 84597); IV, ventral view of an immature carpel; 10,
sepal; 11, petals; 12, seed; drawn from J. Reverchon’s No. 3729, from Terrell-Elmo,
Texas (MBG 84557); V and VI, ventral views of mature carpels; 13, sepals; drawn
from H. Ness’ No. 1994, from Brazos County, Texas (MBG).

JOHNSON: A REVISION OF THE SECTION BORAPHILA 107

Pirate XVIII

108 MINNESOTA STUDIES IN PLANT SCIENCE

PLATE XIX

Section Arabisa. I, capsule of S. leucanthemifolia; 1, petals; 2, sepals; 3, seed;
4, hair of same; drawn from specimen 123lb of the Biltmore Herbarium collection
(UMB).

II. Capsule of S. ferruginea; 5, petals; 6, sepals of same; drawn from Dr.
William Trelease’s No. 4046, from Kodiak, Alaska (MBG 84045); 7, a stamen; 8, a
seed; from Baker’s No. 4921, collected by J. D. Culbertson at Windham Bay, Alaska
(USNH 422334).

III. Pistil of S. bryophora; 9, filament; 10, petals; 11, sepals; drawn from
Bolander’s No. 5043, California; IV, capsule; drawn from Lemmon’s No. 97 (MBG
84200) ; 12, bulbils of Saxifraga foltolosa.

Section Heterisia. V, a flower of S. Meretensiana; VI, advanced pistil; 13, a
sepal; drawn from Thomas Howell’s No. 1428, California (UMH); 14, a filament;
15, petals; 16, a sepal; 17, a cluster of bulbils; 18, a pair of bulbils of S. Mertensiana;
drawn from Elmer’s No. 2645, from the Olympic Mountains, Washington (UMH) ;
VII, capsule of S. Mertenstana; 19, type of hairs in the section Heterésia,

JOHNSON: A REVISION OF THE SECTION BORAPHILA 109

Pirate XIX

VITA

Arthur Monrad Johnson was born at Frederickstad, Norway, January
19, 1878, and emigrated to the United States with his mother and younger
brother in 1882, having been preceded by his father in 1880. The family
settled near Warren, Minnesota. He attended the common schools at War-
ren and later graduated from the state high school at Warren in June, 1897.

He taught in the elementary schools of Marshall County, Minnesota,
for three years and then entered the University of Minnesota in 1900. He
graduated in June, 1904, with a Bachelor of Arts degree; pursued graduate
work in botany at the Minnesota Seaside Station, Vancouver Island, in the
summer of 1906; and in 1919 received the degree of Doctor of Philosophy
from the University of Minnesota. He was an assistant in botany at the
University of Minnesota during the summer session of 1902; assistant prin-
cipal and instructor in science in the high school at St. Cloud, Minnesota,
from 1904 to 1905, and instructor in science in the high school at Colfax,
Washington, from 1905 to 1909. During the summer sessions of 1908 and
1909 he was instructor in botany and geology in the Washington State
College at Pullman, Washington; from 1909 to 1910 he was deputy county
treasurer of Whitman County, Washington; from 1910 to 1916, instructor
in biological science in the: North Central High School, Spokane, Washing-
ton; instructor in botany, University of Washington during the summer
session of 1913; instructor in botany during the summer quarters, De Pauw
University, Greencastle, Indiana, in the summers of 1917 and 1919; from
1916 to 1919, teaching fellow in the University of Minnesota ; from 1919 to
the present, instructor in botany in that institution, and in the summer of
1921, he was appraiser for the Northern Pacific Railroad, Pacific Coast
Division.

He has published the following: The Honey Bee and the Pollination of
Ambrosia, American Botanist, 1917; The Use of the Text-Book in Begin-
ning Classes in Botany, School Science and Mathematics, 1921; The Use of
the Weed-Patch in Teaching High School Botany, School Science and
Mathematics, 1923; Observations on the Carnivorous Propensities of the
Gray Gopher, Journal of Mammology, 1922; Cryptomorpha, a New Section
of the Genus Saxifraga, American Journal of Botany, May, 1923.

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