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MADRONO

A WEST AMERICAN JOURNAL OF BOTANY
VOLUME XX
1969-1970

BOARD OF EDITORS

Class of:

1970—LyMAN BENSON, Pomona College, Claremont, California
MivprepD E. Matutas, University of California, Los Angeles

1971—MarIon OwnsBeEy, Washington State University, Pullman
JoHN F. Davinson, University of Nebraska, Lincoln

1972—IrRA L. Wicctns, Stanford University, Stanford, California
REED C. Roiiins, Harvard University, Cambridge, Massachusetts

1973—WaALLACE R. ERNsT, Smithsonian Institution, Washington, D.C.
ROBERT ORNDUFF, University of California, Berkeley
Roy L. TaAytor, University of British Columbia, Vancouver

1974—-KENTON L. CHAMBERS, Oregon State University, Corvallis
EMLEN T. LITTEL, Simon Frazer University, Burnaby, British Columbia

1975—ArTURO GOMEz-PompPa, Universidad Nacional Autonoma de México
DuNcAN M. Porter, Missouri Botanical Garden, St. Louis

Editor—JoHN H. THOMAS
Department of Biological Sciences
Stanford University, Stanford, California 94305

BUSINESS MANAGER AND TREASURER
June McCAsKILL
P.O. Box 23, Davis, California 95616

Published quarterly by the
California Botanical Society, Inc.
Life Sciences Building, University of California, Berkeley 94720

Printed by Gillick Printing, Inc., Berkeley, California 94710

Es
RN a i
oat
ee
VOLUME 20, NUMBER 1 JANUARY, 1969
Contents

AN ANALYSIS OF GEOGRAPHICAL VARIATION IN
WESTERN NorTH AMERICAN MENZIESIA (ERICACEAE),
James C, Hickman and Michael P. Johnson 1

THE DISTRIBUTION OF PINACEAE IN AND NEAR
NorTHERN NeEvapA, William B. Critchfield and
Gordon L, Allenbaugh 12

LaurA M. LorratneE, 1904-1968, Roxana S. Ferris 26

REvIEws: Paul R. Ehrlich, The Population Bomb
(Kenneth E. F. Watt) ; William A. Weber,
Rocky Mountain Flora (Wallace R. Ernst) ;
C. L. Porter, Taxonomy of Flowering Planis
(Kingsley R. Stern) ; Helen M. Gilkey and
La Rea M. Dennis, Handbook of Northwestern
Plants (Robert Ornduff) 28

Notes AND News: Zoe; NEw DISTRIBUTION RECORD
FOR CLAYTONIA NEVADENSIS FROM NORTHWESTERN
CALIFORNIA, William J. Ferlatte; NEw PuBLicATIons 31

A WEST AMERICAN JOURNAL OF BOTANY

‘UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

POT HCO,
Ke wal! he PS

\
MAR 10 1969
PRGA 7) che iM at 2

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established
1916. Subscription price $6.00 per year ($4.00 for students). Published quarterly in
January, April, July, and October by the California Botanical Society, Inc., and
issued from the office of Madrono, Herbarium, Life Sciences Building, University of
California, Berkeley, California. Orders for subscriptions, changes in address, and un-
delivered copies should be sent to the Corresponding Secretary, California Botanical
Society, Department of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS

LyMAN BENSON, Pomona College, Claremont, California
KENTON L. CHAMBERS, Oregon State University, Corvallis
Joun F. Davinson, University of Nebraska, Lincoln
WALLACE R. Ernst, Smithsonian Institution, Washington, D. C.
ARTURO GOMEZ PompPaA, Universidad Nacional Autonoma de México
EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia
Mitprep E. Maruias, University of California, Los Angeles
ROBERT ORNDUFF, University of California, Berkeley
Marion OwneBey, Washington State University, Pullman
Duncan M. Porter, Missouri Botanical Garden, St. Louis
REED C. CoLiins, Harvard University, Cambridge, Massachusetts
Ira L. Wiccrns, Stanford University, Stanford, California

Editor — JoHn H. THOMAS
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCCASKILL
P. O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Harry Thiers, Department of Ecology and Systematic Biology, San
Francisco State College. First Vice-President: Grady Webster, Department of Bot-
any, University of California, Davis. Second Vce-President: Ernest C. Twisselmann,
Cholame, California. Recording Secretary: John West, Department of Botany, Uni-
versity of California, Berkeley. Treasurer: June McCaskill, Department of Botany,
University of California, Davis.

The Council of the Calfornia Botanical Society consists of the officers listed
above plus the immediate past President, Elizabeth McClintock, California Academy
of Sciences, San Francisco; the Editor of Madrofio; and three elected Council Mem-
bers: Annetta Carter, Department of Botany, University of California, Berkeley;
Robert Ornduff, Department of Botany, University of California, Berkeley; and
Malcolm Nobs, Carnegie Institution of Washington, Stanford.

AN ANALYSIS OF GEOGRAPHICAL VARIATION IN WESTERN
NORTH AMERICAN MENZIESIA (ERICACEAE)

James C. Hickman and MicHAeEt P. JoHNSON

The boreal shrub genus Menziesia J. E. Smith consists of four species
in Japan and two rather closely related species in North America, one
occupying the Appalachian region and the other, M. ferruginea Smith
(1791), occupying coastal and mountainous areas throughout the moist
regions of western North America. All members of the genus are found
in mesic habitats. Occasionally they constitute the dominant understory
vegetation, especially in coastal bogs and forests and at lake margins in
areas of high rainfall or persistent fog or mist.

The western populations were originally described as two separate
species. M. ferruginea Smith included the coastal populations, and M.
glabella Gray (1878), a less pubescent and glandular form, occurred in
the Rocky Mountains. Intermediate specimens, subsequently collected
in the Cascade Range, led Peck (1941) to consider the Cascade and
Rocky Mountain plants together as M. ferruginea var. glabella (Gray)
Peck. Calder and Taylor (1965) have recently made the combination
M. ferruginea ssp. glabella.

The pubescence characters used to differentiate the two entities lack
the geographical coherence suggested by the proposed systems of classi-
fication. The purpose of this study is to examine more carefully and
discuss the geographical differentiation within M. ferruginea.

METHODS

A total of 143 herbarium specimens was studied from throughout the
range of the species. In so far as can be determined, each represents a
distinct population. The geographical range of the species has been di-
vided into seven areas on the basis of physiography, climate, geology and
political boundaries (Detling, 1948). The areas are: 1, the Alaskan
coast; 2, the Canadian coast; 3, the United States coast from Washing-
ton to northern California; 4, the northern Cascade Mountains from
British Columbia to the Columbia River; 5, the southern Cascade
Mountains of Oregon; 6, the Canadian Rocky Mountains; and 7, the
Rocky Mountains of the United States as far south as Wyoming. The
localities and areas are shown in Fig. 1.

The only well-defined geographical discontinuity between areas occurs
in central British Columbia. It isolates the Rocky Mountain populations
from all others. The coastal areas are essentially continuous from north-
ern California to the Kenai Peninsula and merge with the Cascade areas
through the Manning Park-Mt. Seymour area in southwestern British
Columbia. In this instance the line between areas was drawn on geo-
logical grounds, keeping the volcanic Cascades as an entity.

MaproNo, Vol. 20, No. 1, pp. 1-32. February 20, 1969.

2 MADRONO [Vol. 19

GEOGRAPHIC *
DISTRIBUTION OF
M. FERRUGINEA

SE
140 MILES

¢=1 OTU
@= 3-6 0TU's ,

Fic. 1. The localities of analyzed specimens and the boundaries of the geographi-
cal sample areas: 1, Alaskan coast; 2, Canadian coast; 3, United States coast; 4,
northern Cascade iyo ne De southern Cascade Mountains; 6, Canadian Rocky
Mountains; 7, United States Rec: Mountains.

1969 | HICKMAN & JOHNSON: MENZIESIA S

Twenty-nine characters were measured. They included leaf tip shape
and density and length of glandular and puberulent hairs on the young
stem, pedicel, calyx and carpels. For both the upper and lower leaf sur-
faces, density and length of subulate, glandular and puberulent hairs
were measured. Two other characters were derived by summing the
densities of subulate and glandular hairs for both surfaces, giving a
measure of total leaf pubescence. All leaf density measurements for sub-
ulate and glandular hairs were made by superimposing a grid over the
leaf surface and counting all those hairs that fell completely within the
grid, as well as those along two adjacent sides which had their points of
attachment within the grid. Densities of hairs on stems and petioles were
measured by counting the number of hair bases visible on a 1 mm seg-
ment taken immediately proximal to the oldest leaf, and 1 mm below
the calyx, respectively. The puberulent hairs were too small to be
counted by these methods, and their densities were approximated by
measuring the distance of several hairs to their nearest neighbors. The
average distance was then converted to a density measurement. All
measurements of length were made with a micrometer. The leaf tip
shape index resulted from numerically grading a minimum of ten leaves
per specimen from round (1) to acuminate (8).

Sokal and Sneath (1963) suggest using no fewer than sixty characters
in numerical taxonomic studies. However, this study concerns subspe-
cific variation, necessitating analysis of a large number of individuals, and
the number of characters consequently had to be sacrificed. Ehrlich and
Ehrlich (1967) have shown that classifications of butterflies based on
small subsets of characters are highly and significantly correlated with
the classification based on the total characters sampled. This has also
been demonstrated in plants by Johnson and Holm (1968). Although
sufficient data are not available to generalize this phenomenon, we might
accordingly expect the narrow character set we have chosen to signifi-
cantly approximate the total variation pattern in M. ferruginea.

The variation in each character was analyzed by comparing the means
for that character in all paired combinations among the seven areas
using analysis of variance (F tests). Multivariate analysis was made
following the methodology of Sokal and Sneath (1963). Each herbarium
specimen was taken as an operational taxonomic unit and compared
with every other OTU to construct two matrices of similarity measures
between the OTU’s. Taxonomic distance and product moment correla-
tion coefficients were the similarity measures employed. All characters
were standardized such that the mean for each is zero with a variance
of one. Each matrix was clustered using the unweighted pair-group
method with arithmetic means. A phenogram was constructed from each
cluster analysis.

RESULTS

For all characters measured, the variation within any one geographi-

4 MADRONO [Vol. 19

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Fic. 2. Character state distributions for four characters for the seven areas (fig.
1). To the right of each histogram are given all areas which differ significantly for
that character at the five and one percent levels based on the F test.

HICKMAN & JOHNSON: MENZIESIA

1969]

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Fig. 2 continued.

6 MADRONO [Vol. 19

TABLE I. A List oF THOSE CHARACTERS FOR WHICH THERE ARE NUMEROUS SIG-
NIFICANT DIFFERENCES BETWEEN GEOGRAPHICAL AREAS ~-IN PAIRED COMBINATIONS.
Atso GIVEN ARE THE GENERAL GEOGRAPHICAL PATTERNS OF VARIATION OF THESE

CHARACTERS.

Character

Density of subulate hairs on
upper leaf surface

Total density of hairs on
upper leaf surface

Length of subulate hairs on
upper leaf surface

Length of glandular hairs on
upper leaf surface

Density of glandular hairs on
lower leaf surface

Length of glandular hairs on
lower leaf surface

Density of glandular hairs
on young stem

Length of glandular hairs on
young stem

Density of puberulent hairs
on pedicel

Length of glandular hairs on
pedicel

Density of puberulent hairs
on calyx

Length of longest cilium
on calyx

Density of puberulent hairs
on carpel

Density of glandular hairs
on carpel

Length of glandular hairs
on carpel

Leaf tip index

Description of Variation Pattern

Decreasing clinally inland; Cascade and Rocky
Mountains similar (fig. 2)

Decreasing clinally south and inland, but reversed in
the Rocky Mountains

Decreasing clinally south and inland

Decreasing clinally south and inland, but reversed
in the Rocky Mountains

Unclear, generally decreasing south and inland

Decreasing clinally south and inland, but reversed
in the Rocky Mountains

Increasing clinally south on coast, decreasing inland

Increasing clinally south and inland; southern
Cascade and U.S. Rocky Mountains similar

Decreasing clinally inland; three groups: Coast,
Cascades and Rocky Mountains (fig. 2)

Two groups, no clinal change: southern Cascades
and Rocky Mountains similar (fig. 2)

Increasing clinally south and inland, but reversed
in the Rocky Mountains

Coast constant; others increasing south and inland

Coast constant; others increasing south and inland
Not clinal: Coast lowest; Cascades highest; Rocky
Mountains intermediate

Decreasing clinally south and inland; jump down-
scale (to roundness) at Rocky Mountains (fig. 2)

cal area overlaps the variation in all other areas. A series of histograms
of four characters for each of the seven areas is given in Fig. 2. Also
included are code numbers for those areas which differ significantly
from the area-character under consideration. Significance at p < .05 and
p < .01 is based on the F test between the means. Those characters
which exhibit significant differences between samples in several paired
combinations are given in Table I, with descriptions of the patterns of
character variation.

1969] HICKMAN & JOHNSON: MENZIESIA fl

For character sets involving size, such as those used in this study,
Sokal and Sneath (1963) suggest that a correlation coefficient gives a
better measure of similarity than taxonomic distance. As predicted, in
the phenogram based on taxonomic distance, several geographically un-
related and morphologically diverse specimens clustered with one an-
other or with the major clusters only at unusually high levels. Since
these individuals are extreme for one or more measurements, it seems
that the clustering pattern here is adversely affected by characters in-
volving size. For this reason, we consider the phenogram based on cor-
relation coefficients to be the more appropriate measure of similarity,
and further discussion will pertain to it.

The OTU’s fall into three large clusters which we refer to as Rocky
Mountain, Cascade and Coastal (fig. 3). The Rocky Mountain cluster is
the most homogeneous: it contains Rocky Mountain OTU’s almost ex-
clusively, and almost every Rocky Mountain OTU is included in it. The
Cascade and Coastal clusters are heterogeneous. They contain a pre-
dominance of OTU’s from the areas for which they are named, but also
include a significant number of OTU’s from other areas. The Cascade
and Rocky Mountain clusters show greater affinity to one another than
either does to the Coastal cluster.

DISCUSSION

The patterns of variation of characters in Table I show an independ-
ence, or “discordance.”’ Regarding the problem of concordance and dis-
cordance at subspecific levels, Ehrlich and Holm (1964, p. 166) state
that “with discordance predominaitng, subspecies recognized on the
basis of one or a few convenient characters would not be evolutionary
units. They would be simply units of convenience for filing specimens.”

Figure 2 presents a fraction of the original data and demonstrates
some of the more important variation patterns in detail. The density of
subulate hairs on the upper leaf surface is the most readily observed
character for differentiating the described M. ferruginea var. glabella
(Gray) Peck from the more pubescent typical M. ferruginea of coastal
regions. This character shows a clinal decrease inland, with the Rocky
Mountain and Cascade populations varying as a unit. The coastal popu-
lations form another quite distinct statistical unit. This character, taken
alone, would indicate that Peck was correct in stating that the var.
glabella should include both Rocky Mountain and Cascade plants
(1941). If we consider leaf tip shape, however, another pattern becomes
apparent. The coastal and Cascade specimens show a clinal increase in
roundness of the leaf tip as one progresses south and inland. The Rocky
Mountain plants show a sharp break from all the other populations,
having for the most part quite round leaf tips. This pattern, if taken
alone, would lead one to consider the Rocky Mountain form to consti-
tute the most nearly distinct subspecific unit. The length of glandular
hairs on the pedicel exhibits yet another pattern of variation. Here all

8 MADRONO [Vol. 19

6 55:4
a 4 3 5 4
a 4 3 5

CASCADE CLUSTER

5 Rocky Mountain OTU's
25 Cascade OTU's
8 Coastal OTU's

COASTAL CLUSTER

15 Cascade OTU's
57 Coastal OTU's

ROCKY MOUNTAIN CLUSTER

32 Rocky Mountain OTU's
| Cascade OTU

Fic. 3. Phenocycle resulting from the cluster analysis of the matrix of correla-
tion coefficients. The OTU’s are labeled as to the geographical area from which
they came (fig. 1). A summary of the constituents of each of the major clusters is
given. A dotted line marks the boundary between the Cascade and Rocky Moun-
tain clusters.

of the three major groups of populations are significantly different from
one another, with the cline again decreasing inland. Although the clines
for these three characters are generally similar, their relative expres-
sions follow different patterns, and they must thus be considered dis-
cordant characters.

Hitchcock, e¢ al. (1959) state the range of M. ferruginea var. glabella
(Gray) Peck as follows: “Rocky Mts., Alta. and B.C. to Wyo., west-
ward to e. Wash. and Oreg. and down the Columbia to Mt. Hood and
Mt. Adams, where the two varieties freely interbreed.” The only speci-
men from the Columbia Plateau Province of which we are aware was
collected at Twin Lakes, in northeastern Washington (Ferry Co.). This
collection came to our attention too late to be included in the analyses.
In fact, Menziesia seems to be poorly collected throughout the critical

1969 | HICKMAN & JOHNSON: MENZIESIA 9

western part of the Rocky Mountain areas. However, some indirect evi-
dence supporting Hitchcock’s statement is found by referring to the
density of puberulent hairs on the calyx in Fig. 2. The Oregon Cascade
and United States Rocky Mountain areas form a single statistical popu-
lation, separate from all others, being the only areas in which the mode
for this character is not zero. The pattern of variation for this character
would certainly indicate some degree of gene flow across the Columbia
Plateau or the northern Great Basin in the present or recent geologic
past.

The foregoing comments necessitate a more detailed consideration of
the probable migrational history of the genus in western North America.
It appears from present distributions of member species and by analogy
with ecologically similar species which have left a paleontological record
that Menziesia has migrated southward from an originally boreal dis-
tribution with the increasingly temperate climate in this region during
the first half of the Tertiary. In the Ericaceae, generic distinctions on
the basis of pollen morphology are difficult, and direct evidence of
Menziesia’s migrational routes are lacking (Hansen, 1947; 1955; Heus-
ser, 1960). The following discussion assumes that Menziesia has a simi-
lar migrational history to other boreal species whose pollen records have
been studied.

In Wisconsin Pleistocene times, glaciers formed in the mountains of
British Columbia, which led to the development of more massive pied-
mont glaciers and finally to a virtually continuous ice sheet that covered
all of western Canada and the northern half of Washington and parts of
Idaho and Montana. Alpine valley glaciers and piedmont glaciers also
formed in the mountain ranges much farther to the south (Flint, 1945;
1957). The interior of Alaska and the more westerly portions of the
Yukon Territory were not covered by the ice sheet. This area must have
acted as a refugium for many boreal species during the Wisconsin glacial
period. The western North American ice sheet was thickest and _ per-
sisted the longest in central British Columbia where it had no direct
outlets (Flint, 1957); Menzzesia is today absent from this area. It seems
likely that this is due not only to the long persistence of Wisconsin ice,
but to the warming and drying trend which immediately followed the
melting of the glaciers and culminated about 6,000 years ago (Hansen,
1947; 1955) making the area climatically unsuitable for Menziesia.

It also seems likely that during glacial periods Menziesia occupied
much area in the southern Cascades and the Great Basin where it has
not been able to persist. Certain present distributions support this hy-
pothesis. Menziesia commonly co-occurs with Chamaecyparis noot-
katensis (Lamb.) Spach along the coasts of British Columbia and
Alaska. The two also co-occur at the southernmost montane locality
known for Menziesia, in the Oregon Cascades (J. C. Hickman 492-4,
ORE.; J. C. Hickman 492-5, ORE). Several collections of Chamaecy-
paris have also been taken from an evidently relictual population of

10 MADRONO [Vol. 19

large trees in the Aldrich Mountains of east-central Oregon (A. Cron-
quist 7646, DS; O. V. Mathews, 1940 DS). These collections may indi-
cate that cool, moist conditions prevailed during the Pleistocene in much
of what is now semi-arid region between the Cascade and Rocky Moun-
tains, and that Menziesia could have been widely distributed through
this area.

The interrelationships among the populations have also been analyzed
by computing phenetic similarity (fig. 3). On the lower clustering levels
the complexity of patterns is the most striking characteristic. This is
not true for the Rocky Mountain areas, however, indicating that they
constitute the least variable grouping. Other small clusters show great
geographical diversity in their members: OTU’s cluster first with a
member of the same area in only one third of the instances. It is the
higher clustering levels that should be expected to indicate possible sub-
specific taxonomic divisions. The Rocky Mountain populations and the
bulk of the Cascade populations cluster in a large group, parallel to the
large group of coastal forms with which the remainder of the Cascade
individuals are more closely allied. Here the homogeneity of the Rocky
Mountain cluster does not imply that these plants constitute the most
distinct grouping. It is not surprising that the Cascade and Rocky
Mountain materials are phenetically similar since these two areas prob-
ably have greater environmental similarities than has either with the
coastal area. That the Coastal cluster contains numerous OTU’s from
the Cascade area (and the converse) suggests the possibility of greater
or more recent gene flow between them. This is supported by the pres-
ent geographical continuity between the coastal and Cascade areas at
Manning Park and the possibility of more southerly connections in the
recent geologic past as suggested by Detling (1954; 1958).

CONCLUSIONS

The variation pattern in Menziesia ferruginea is complex. Univariate
and multivariate analyses show that no single character nor set of char-
acters studied can be used to separate individuals into geographically or
ecologically coherent categories. Thus, the erection of subspecific taxa is
inappropriate and inadequate for describing this pattern. It must rather
be explained in terms of migrational history, past and present gene flow,
and adaptations to existing environments. These factors have led to an
overlapping and partially discordant complex of morphological clines
from north to south and from the coast to the Rocky Mountains. We
suggest that in future works botanists consider MW. ferruginea Smith var.
glabella (Gray) Peck to be a later synonym for WM. ferruginea Smith.

ACKNOWLEDGMENTS

The authors would like to thank the late L. E. Detling for his advice
throughout the course of this work and S. A. Cook for his continuing
interest and counsel. Richard W. Holm, Duncan M. Porter and Thomas

1969] HICKMAN & JOHNSON: MENZIESIA 11

A. Ebert kindly read and commented on the manuscript. The materials
used in this study came from the following herbaria: DS, OSC, ORE,
UBC, WTU, UC, and MICH. We would like to express our gratitude
to Adam F. Szcawinski and Marion Ownbey, for their kind cooperation.
Computer time for the analysis of variance was provided by the Univer-
sity of Oregon Statistical Laboratory and Computing Center. Multi-
variate analysis were done at the Stanford University Computer Center
using programs modified from NT-SYS developed by F. James Rohlf.
The work has been supported in part by Grant 2G-365-R1 from the U.S.
Public Health Service-National Institutes of Health to Stanford Uni-
versity.

Department of Botany, Washington State University, Pullman
Department of Biological Sciences, Florida State University, Tallahassee

LITERATURE CITED

Caper, J. A., and R. L. Taytor. 1956. New taxa and nomenclatural changes with
respect to the flora of the Queen Charlotte Islands, British Columbia. Canad. J.
Bot. 43:1387-1400.

Der Linc, L. E. 1948. Concentration of environmental extremes as the basis for vege-
tation areas. Madronfo 9:169-185.

. 1954. Significant features of the flora of Saddle Mountain, Clatsop Coun-
ty, Oregon. Northw. Sci. 28:52—60.

. 1958. Peculiarities of the Columbia River Gorge flora. Madrofio 14:160-—
128

EuriicuH, P. R., and A. H. Eurtiicu. 1967. The phenetic relationships of the butter-
flies. I. Adult taxonomy and the non-specificity hypothesis. Syst. Zool. 16:301-
37,

EuriicH, P. R., and R. W. Horm. 1964. In A. Montagu, ed. The concept of race.
Collier-Macmillan, London.

Flint, R. F., et al. 1945. Glacial map of North America. Special Pap. Geol. Soc.
Amer. 60:1-37.

. 1957. Glacial and Pleistocene geology. John Wiley, New York.

Gray, A. 1878. Synoptical flora of North America. Vol. 2. Iveson, Blakeman, Tay-
lor, New York.

Hansen, H. P. 1947. Postglacial forest succession, climate, and chronology in the
Pacific Northwest. Trans. Amer. Philos. Soc. 37:1—130.

. 1955. Postglacial forests in south-central and central British Columbia.
Amer. J. Sci. 253:640—658.

HeEusseEr, C. J. 1960. Late-Pleistocene environments of North Pacific North Amer-
ica. Special Publ. Amer. Geog. Soc. 35:1—308.

Hitcucocyu, C. L., A. Cronquist, M. Ownsey, and J. W. THompson. 1959. Vas-
cular plants of the Pacific Northwest. IV. Univ. Washington Press, Seattle.

Jounson, M. P., and R. W. Horm. 1968. Numerical taxonomic studies in the genus
Sarcostemma. (In press).

Peck, M. E. 1941. A manual of the higher plants of Oregon. Binfords and Mort,
Portland.

SM1TH, J. E. 1791. Plantae Icones Ined. pl. 56.

SOKAL, R. R., and P. H. A. SngatTH. 1963. Principles of numerical taxonomy. W. H.
Freeman, San Francisco.

THE DISTRIBUTION OF PINACEAE IN AND NEAR
NORTHERN NEVADA

WILLIAM B. CRITCHFIELD and GORDON L. ALLENBAUGH

INTRODUCTION

More than 50 years ago G. B. Sudworth (1913) observed: ‘Contrary
to popular belief, our present knowledge and published records of the
geographic range of North American trees is still very incomplete.” This
statement remains true today for much of the interior West, particularly
the semi-arid Great Basin. Although the botanical exploration of this
region began over a century ago, information about the distribution of
native trees is still sketchy and inaccurate. This is true even of the
Pinaceae, although most of the trees in this family are readily identifiable
and are among the most conspicuous elements of the vegetation.

One reason for this lack of information is the unique topography of
the Great Basin, which is made up of many smaller basins with interior
drainage. Tree growth is mostly confined to the numerous isolated moun-
tain ranges, called basin ranges. These ranges, usually oriented north
and south and typically much longer than wide, occupy about half the
total area of the Great Basin (Fenneman, 1931). The intervening valleys,
which sometimes contain playas or playa lakes, are usually treeless.
Many of the basin ranges are rather inaccessible, and plant collections
from some of them are few or nonexistent.

Throughout much of the Great Basin three pines, Pinus monophylla,
P. flexilis, and P. aristata, are among the dominant elements of the mon-
tane vegetation (Billings, 1951). Toward the north and northwest, how-
ever, they disappear and other members of the Pinaceae appear as out-
liers of the coniferous forests that border the northern Great Basin. This
transitional area is the region covered by this paper (fig. 1). It includes
much of the northern Great Basin and a physiographically similar portion
of the Snake River drainage (southeastern Idaho and adjoining parts of
Oregon and Nevada). This paper reports several extensions of species
ranges, summarizes what is currently known about the distribution of
the Pinaceae in this region, and reviews the events that may have
brought about the present “insular” distribution of these conifers.

SOURCES OF INFORMATION

The first detailed maps of these conifers were those of Sudworth
(1913; 1916; 1918). His original working maps, which are on file at the
Washington Office of the U. S. Forest Service, have been helpful in
establishing the sources of his information. Revised versions of Sud-
worth’s maps were published by the Forest Service in 1938 (Munns,
1938). Recently, revised distribution maps of all of the Pinaceae of this
region have been published (Fowells, 1965; Critchfield and Little, 1966).

1969] CRITCHFIELD & ALLENBAUGH: PINACEAE 13

The Idaho ranges of these species have also been mapped by Johnson
(1966).

The principal descriptions of the distribution of the Pinaceae in this
region are those of Billings (1954) and Little (1956). Parts of the region
were covered by Sudworth (1908) and Holmgren (1942).

Unpublished sources of information include field observations and
specimens in several herbaria. Our collections are not cited in the text,
but the localities that we visited are indicated in Fig. 1. Specimens from
these localities are in the conifer herbarium of the Institute of Forest
Genetics, Placerville, California (IFGP: Critchfield, 1966), and some of
them are duplicated in these herbaria: US, USFS, and MSC. Several
government agencies which have supplied us with information and col-
lections are cited in the text by these abbreviations: Forest Service, U. S.
Department of Agriculture (FS); Bureau of Land Management, U. S.
Department of the Interior (BLM); and Nevada State Fish and Game
Commission (NFG).

The contributions of several individuals and agencies are referred to
in the text. We also wish to thank J. R. Griffin, E. L. Little, Jr., and J. L.
Jenkinson for their help.

THE DISTRIBUTION OF SPECIES

ABIES CONCOLOR (Gord. & Glend.) Lindl. White fir. Both California
white fir and Rocky Mountain white fir extend into widely separated
parts of this region. The former is sometimes called A. lowiana (Gord.)
A Murr. or A. concolor var. lowiana (Gord.) Lemm. to distinguish it
from Rocky Mountain white fir, which extends from the southern Rocky
Mountains into the eastern and southern parts of the Great Basin. These
two geographically separated strains differ in morphology, but their status
as separate taxa 1s not universally recognized.

It has been known for many years that California white fir grows in
the Warner Mountains of northeastern California and southern Oregon
and in the nearby Hart Mountain area of southern Oregon (Sudworth,
1908). White fir is one of the commonest trees in the mixed-conifer
forest of the Sierra Nevada and southern Cascade Mountains, and the
forests of the Warner Mountains are an impoverished eastward extension
of this vegetation type.

East of the Warner Mountains, several unreported outliers of white fir
grow in a range of low mountains in the northwestern corner of Nevada.
These mountains do not have a generally accepted name now, but in the
past they have been called the East Warner Mountains. This range is
separated from the Warner Mountains by Surprise Valley and from the
mountains to the east by Long and Coleman Valleys.

Near the northern end of the range, just west of Coleman Valley and
a few miles south of the Oregon border, are two fairly extensive creek-
bottom stands of white fir about two miles apart (fig. 1A: 41°57.7’ N
Lat, 119°50.0’ W Long, and 41°56.4’, 119°49.5’). They are growing at

[Vol. 19

MADRONO

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1969 | CRITCHFIELD & ALLENBAUGH: PINACEAE 15

5500-6800 ft along spring-fed streams which flow eastward into Coleman
Valley (BLM). J. C. Fremont’s first expedition may have passed within
sight of these stands when it entered Nevada through Coleman Valley
early in 1844. The first and only botanists known to have visited them
are F. V. Coville and J. B. Leiberg, who collected white fir in this vicinity
in 1896 (Coleman Valley, Coville & Leiberg 119, US).

Twenty-five miles to the south is a third and much smaller outlier
growing around a spring at the head of Forty-nine Canyon (fig. 1B:
41°35.2’ N Lat, 119°54.9’ W Long, 6000 ft elev.). This grove, also re-
ported to us by BLM, is within sight of Nevada State Highway 8A. It
consists of 10 large trees and several times as many seedlings and sap-
lings. The largest trees range up to 4 ft in diameter at breast height, and
are at least 200-300 years old.

These outliers of California white fir are separated by more than 200
miles from the northwestern limits of Rocky Mountain white fir (fig. 1).
The latter is present in the higher ranges of east-central and southern
Nevada (Billings, 1954; Little, 1956), and extends north into southern
Elko County. There it grows on Pilot Peak (NFG, BLM), in the Toana
Range (NFG; A. H. Holmgren, pers. comm.), in the Pequop Mountains
(NFG), and on Spruce Mountain (Zavarin 723-737, IFGP). A single
stand in the northern Ruby Mountains (fig. 1C) is located at an eleva-
tion of about 8500 ft in Seitz Canyon, at 40°39’ N Lat, 115°28’ W Long
(CES).

Older distribution maps erroneously show white fir in many of the
mountain ranges of northern Elko Co., south-central Idaho, and north-
western Utah (Sudworth, 1916; and Munns, 1938). The most recent
map of the species still shows it on one of these ranges—the South Hills,
in Idaho (Fowells, 1965). The presence of white fir north of southern
Elko Co. has not been verified, and this error may have originated from
confusion between this species and A bes lasiocar pa.

ABIES GRANDIS (Dougl.) Lindl. Grand fir. The typical form of grand
fir, which ranges from northern California to western Montana, does not
enter or even approach the Great Basin. In the southern part of its range,
however, grand fir appears to intergrade with white fir through a broad
zone extending from northwestern California to western Idaho. This
intermediate form extends into the northwestern Great Basin. The only
representative of the Pinaceae in the southeastern corner of Oregon, it is
present in a single locality on the west slope of Steens Mountain (fig. 1D).

Although this stand is not shown on distribution maps or listed in
floras, its existence has been known for many years. F. V. Coville visited
it in 1896 (Coville 598, US). An early geological report on Steens Moun-
tain probably referred to this stand in the following statement: “A few
pines, firs, junipers, and cottonwoods grow in the deep canyons on its
west slope” (Russell, 1903). In another geological report Waring (1909),

Fic. 1. The distribution of Pinaceae in northern Nevada and adjacent areas.
Circled letters A to L are localities mentioned in the text.

16 MADRONO [Vol. 19

commenting on the remarkable absence of timber on this range, noted
as one of the exceptions ‘‘a small group of firs in one canyon.” In 1938,
O. V. Matthews collected wood samples of the fir, which he called A.
concolor (Anon, 1944).

The Steens Mountain fir grows along the banks of Big and Little Fir
Creeks and on the north-facing slopes of their canyons. The stand ex-
tends a short distance below the point where they join to form a stream
known as Mud or Fir Canyon Creek. The main stand ranges in elevation
from 5900 to 6200 ft, and covers an estimated 30-40 acres (42°47’ N Lat,
118°43’ W Long). About a mile to the south is an outlier consisting of
a few trees (BLM).

The Steens Mountain stand is nearly equidistant from A. concolor in
south-central Oregon and the intermediate populations of the Blue Moun-
tains to the north. We have grouped it with A. grandis because it is
morphologically similar to the Blue Mountains fir, which at present is
commonly classed with A. grandis (Fowells, 1965).

ABIES LASIOCARPA (Hook.) Nutt. Subalpine fir. This high-elevation
tree extends westward from the Wasatch Mountains into the Great Basin
at least as far as the Deep Creek Range, just southeast of the region
considered here (McMillan, 1948). It is absent from the isolated ranges
in the northern Great Basin, but in the northern part of Elko Co. it is
one of the commonest trees in the mountains that form the divide between
the Great Basin and the Snake River drainage. It grows in the Bull Run
and Jarbidge Mountains, and in the Independence Mountains it extends
at least as far south as the Jack Peak area at 41°30’ N Lat (FS). Sub-
alpine fir is also a common tree on two ranges north of this divide—the
Silver City Range and the South Hills (fig. 1). From the South Hills this
fir extends eastward to the Wasatch and Rocky Mountains.

Abies lastocarpa may be the fir that Sereno Watson encountered in
the extreme northwestern corner of Utah on King’s 1867—68 Fortieth
Parallel Expedition. Watson crossed the northern end of the Goose Creek
Mountains, which extend southwest from this corner of Utah into Ne-
vada. In some of the moist canyons, he found a tree that he called A.
grandis (Watson, 1871). His description does not fit any of the western
firs, but A. lasiocarpa is the only fir on the nearby ranges, including the
Albion Mountains (FS) and South Hills to the north and the Raft River
Mountains to the east (Preece, 1950).

A report of A. lasiocarpa on Steens Mountain (Bailey, 1936) is prob-
ably a mistake. Bailey’s table 7 lists it in the Hudsonian zone of this
range, but in the text he notes that the aridity of Steens Mountain elimi-
nates all of the trees characteristic of the Hudsonian zone.

PICEA ENGELMANNII Parry. Engelmann spruce. This species—the only
spruce of the northern Great Basin—is common north and east of this
region. It extends west from the Rocky Mountains to the Raft River
Mountains (Preece, 1950) and the Albion Mountains (FS). It stops

1969 | CRITCHFIELD & ALLENBAUGH: PINACEAE 17

short of the South Hills in southern Idaho, although it is often shown
there (Munns, 1938; Johnson, 1966; Fowells, 1965). Farther south,
another salient extends westward from the Wasatch Mountains to the
Deep Creek Mountains (McMillan, 1948) and the nearby high ranges
of eastern Nevada (Billings, 1954; Little, 1956).

In the region treated here, Engelmann spruce is definitely known in
only one locality, the head of Thorpe Creek in the northern Ruby Moun-
tains (FS). This stand is located at an elevation of 9-10,000 ft at 40°42’
N Lat, 115°20’ W Long (fig. IE). There may be other small outliers in
this range, however. Sereno Watson, the first botanist to visit the Ruby
Mountains, noted that “in some of the high western canyons there is a
dense growth of Abies [Picea] engelmannii’”’ (Watson, 1871). He traveled
extensively in the southern and central Ruby Mountains, and his routes
are shown on the map accompanying his account. Since the northernmost
point that Watson visited was nearly 10 miles south of Thorpe Canyon,
the stands of Engelmann spruce that he found may have been overlooked
since 1868.

Watson called the Ruby Mountains the “East Humboldt Mountains.”
The later change of names has caused a great deal of confusion, since two
other ranges in northern Nevada are sometimes called the East Hum-
boldt Mountains at present. One of them, a northward extension of the
Ruby Mountains, was called the Clover Mountains by Watson. The
other range, located east of the Humboldt Mountains in Pershing County,
is called the East Humboldt or East Range. (Watson referred to it as the
Pah-Ute Mountains.) A Pershing County outlier of P. engelmanni shown
by Munns (1938) can probably be attributed to this change in place
names.

Engelmann spruce is also sometimes shown in the Silver City Range
(Munns, 1938; Fowells, 1965). This southern Idaho occurrence has not
been verified, and is probably in error.

PSEUDOTSUGA MENZIESII (Mirb.) Franco. Douglas fir. Of the two
generally recognized geographic races of this species, only the Rocky
Mountain Douglas fir (var. glauca (Beissn.) Franco) extends into this
region. The Pacific Douglas fir (var. menziesii) reaches its eastern limits
west of the Warner Mountains.

Rocky Mountain Douglas fir, like Engelmann spruce, extends west
from the Wasatch and Rocky Mountains in two salients. In the north it
reaches the Raft River Mountains (Preece, 1950) and the Albion Moun-
tains (FS). Like Engelmann spruce, Douglas fir stops short of the South
Hills, although it is generally mapped there (Sudworth, 1918; Fowells,
1965; Johnson, 1966). The southern salient extends westward from the
Wasatch Mountains to the Deep Creek Range (McMillan, 1948) and
the higher ranges of east-central Nevada (Billings, 1954; Little, 1956).
Unlike Engelmann spruce, Douglas fir is not known to extend north
into Elko Co., although Sudworth (1918) and Munns (1938) showed it

18 MADRONO [Vol. 19

in the Ruby Mountains on the basis of an unpublished report accompany-
ing Sudworth’s original working map of this species (‘““Ruby Mountains
at Harrisons Pass, scarce’’).

The only known stands of Douglas fir in this region are in southwestern
Idaho. It is a common tree in the Silver City Range, sometimes consid-
ered part of the Owyhee Range. Two other sizable but previously un-
reported outliers are present in the Owyhee Range to the south (BLM;
fig. 1F, G). Around South Mountain (42°45’ N Lat, 116°55’ W Long)
this species is distributed rather widely, with a much smaller patch
(100-200 acres) about 12 miles to the southeast (42°38’, 116°42’).

Douglas fir has also been reported in the mountains of northern Elko
Co. (Billings, 1954; Little, 1956; Fowells, 1965), but we consider this
occurrence doubtful. These reports are based on specimens collected on
Cobb Creek and Merritt Mountain, south and northeast of Mountain
City (Nichols & Lund 373, 453, RENO). However, Forest Service and
other local informants are unanimous in stating that they have never
seen Douglas fir growing in nature anywhere in these mountains.

PINUS MONTICOLA Doug]. Western white pine. This pine is widely
distributed north and west of the Great Basin. It enters the region con-
sidered here only at its western edge, in the Warner Mountains. It was
first encountered on the higher peaks of this range by C. Hart Merriam
in 1896 (Sargent, 1897).

PINUS ALBICAULIS Engelm. Whitebark pine. This pine is characteristic
of high elevations, extending to timberline, in the high mountains west
and north of this region and in the Warner Mountains. It is also un-
expectedly common in several mountain ranges of northern Nevada. Its
presence there was not reported until recently, although it has occasion-
ally been collected in this region during the past 70 years. One reason that
P. albicaulis has been generally overlooked is its close similarity to P.
flexilis, which is much more widely distributed in the Great Basin region.
The two species differ greatly in their cones, but the rather fragile cones
of P. albicaulis are nearly always destroyed by birds or small mammals
as they approach maturity or soon thereafter. Vegetatively these two
white pines are much alike; a reported difference in the number and
distribution of resin canals in the needles of trees in southern Montana
and western Wyoming (Ericson, 1964) does not hold true in Canada
(Brayshaw, 1965) or northern Nevada.

The presence of whitebark pine in the higher parts of the Warner
Mountains has been known since C. H. Merriam found it there in 1896
(Sargent, 1897). Sudworth (1913) and Munns (1938) did not record
it elsewhere in this region. Their maps show it in two localities to the
east, the Albion Mountains of Idaho and the Wasatch Mountains east
of Salt Lake City, but we have been unable to verify either of these
occurrences. Holmgren (1942) did not list whitebark pine in his flora
of northeastern Nevada. Billings (1954) reported ‘‘vegetative specimens
(probably authentic) ... from... Pine Mountain in Humboldt County.”

1969 | CRITCHFIELD & ALLENBAUGH: PINACEAE 19

We have not been able to locate a Pine Mountain in Humboldt Co., but
there is a Pine Mountain in northern Elko Co. (41°46’ N Lat, 115°37’
W Long). Authentic P. albicaulis has been collected there (Hitchcock
1177, US), and this may be the locality to which Billings referred.

The first report that whitebark pine is widely distributed in northern
Nevada was that of Little (1956). He recorded it in the Pine Forest
Range of Humboldt Co., and the Jarbidge Mountains, Ruby Mountains,
and Pine Mountain, all in Elko Co. In addition to these areas, it is now
known to occur in the East Humboldt Range and the Bull Run Moun-
tains, both in Elko Co. (fig. 1).

Whitebark pine is the only pine of the Pine Forest Range. This stand
is far removed from any other stand of this species—at least 70 miles
east of the Warner Mountains and nearly twice that distance west of
the nearest stands in northern Elko Co. Presumably this is the pine re-
ferred to in an early geological report: ““The Pine Forest Mountains are
covered over a limited area with a forest of yellow pine, from which this
range derives its name” (Russell, 1885). Whitebark pine was collected
here in 1896 (Streator 1015, US) and 1901 (Griffiths & Morris 225, US),
but these collections were overlooked for more than 50 years. The pine
of the Pine Forest Range was identified as “Pinus flexilis (?)” by Tay-
lor (1912), a zoologist who did extensive field work in this area. His
identification was based partly on foliage, ‘“‘no cones being at hand,” and
partly on geographic and zonal considerations. One of Taylor’s photo-
graphs of this pine was included in Hall’s comprehensive treatment of
the mammals of Nevada, captioned ‘limber pines in the Pine Forest
Mountains” (Hall, 1946, plate 2b). The identification of this pine as
P. albicaulis by Little (1956) is the first mention in the botanical and
forestry literature of this or any other pine in these mountains. The Pine
Forest stand was visited a few years ago by A. Cronquist, who noted that
the pines are confined to the granitic rocks that make up the core of this
range (pers. comm.).

In the Jarbidge Mountains of northern Elko Co., P. albicaultis is the
only common pine; P. flexilis is present but rare. In this region P. albz-
caulis reaches much lower elevations than it does at the same latitude in
California. It grows from as low as 6400 ft in the valley of the Jarbidge
River to above 10,000 ft in the Jarbidge Mountains to the east. It is
known to extend as far northeast as Pole Creek, at about 41°55’ N Lat,
115°15’ W Long (Nelson & McBride 2070, US, A). Its range in north-
ern Elko Co. has recently been extended as far west as the Bull Run
Mountains, where it was collected by S. A. Scott (FS) at 41°42’ N Lat,
116°08’ W Long, 8600 ft elevation (Scott, 1965, IFGP). This species
has not yet been reported from the Independence Mountains, a higher
southward extension of the Bull Run Mountains.

South of the Humboldt River P. albicaulis appears in the connected
Ruby and East Humboldt Ranges. Around Angel Lake, at 8400 ft in the
East Humboldts, it is intermingled with P. flexilis. In the Ruby Moun-

20 MADRONO [Vol. 19

tains we have seen it at the head of Lamoille Canyon, a U-shaped glaci-
ated valley which penetrates deeply into the northern part of the range.
Below 8500 ft we saw only P. flexilis, but above that elevation P. albi-
caulis is the more common of these two pines. Both species are still
present at 10,000 ft on the crest of the range south of Lamoille Canyon.
P. albicaulis extends at least as far south as Green Mountain, at 40°23’
N Lat (FS), but it has not yet been reported in that part of the Ruby
Mountains south of Harrison Pass. At Green Mountain it reaches its
southermost known limit in the region east of the Sierra Nevada.

PINUS FLEXILIS James. Limber pine. This pine, one of the character-
istic trees of the basin ranges, is present on the higher mountains
throughout most of Nevada except the extreme western part (Billings,
1954; Little, 1956). Holmgren (1942) reported that it is frequent in
the higher mountains of Elko Co. Most distribution maps (Sudworth,
1913; Munns, 1938; Little, 1949) show it in the mountains of Elko Co.
and in the Warner Mountains of California.

Limber pine is widespread in the eastern part of this region. It was
collected in the Ruby Mountains by Sereno Watson nearly a century
ago (““East Humboldt Mountains,” Watson 1113, US). Holmgren (1942)
noted limber pine on Spruce Mountain, at the southern end of the Pequop
Mountains (fig. 1). Here, according to R. D. Wright (pers. comm.), it is
mixed with bristlecone pine (P. aristata) in an extensive forest that ex-
tends from 9000 ft to the top of the mountain.

In the northern part of Elko Co., limber pine is present in the Inde-
pendence Mountains (Kinmnaman 21, FS), Bull Run Movntains (Scott,
1965, IFGP), Pine Mountain (Hitchcock 1176, US), and the Jarbidge
Mountains, where it is quite rare. We found only two small groves in
the valley of the Jarbidge River, one growing on a rocky outcrop on the
canyon wall (41°48’ N Lat, 115°24’ W Long), and the other on the
bank of the river (41°55’, 115°257).

Elsewhere in Elko Co. limber pine is present in most of the higher
mountains (fig. 1). For this information we are indebted to L. W. Hos-
kins (NFG, pers. comm.).

North and east of this corner of Nevada the distribution of limber
pine is sporadic. It is absent from the South Hills, but is frequent at
higher elevations in the Raft River Mountains (Preece, 1950), and is
present in the Alb‘on Mountains (Johnson, 1966). In the Silver City
Range of southwestern Idaho, only two trees of this species have been
found (fig. 1H). They are growing near the top of War Eagle Moun-
tain at 8100 ft elevation, 42°59’ N Lat, 116°40’ W Long (W. H. Baker,
pers. comm.).

In the western half of this region, limber pine is present only in the
Santa Rosa Range of eastern Humboldt Co. It does not appear to have
been reported from this range before, although it is not uncommon. It
grows in scattered patches near the crest of the mountains.

Limber pine is shown in the Warner Mountains on many distribution

1969] CRITCHFIELD & ALLENBAUGH: PINACEAE 21

maps of this species. This error can be attributed to an early misidentifi-
cation of some other five-needled pine, probably western white pine. Ex-
cerpts from a 1903 report by Filibert Roth on the Warner Mountains
Forest Reserve, which are on file at the Washington Office of the Forest
Service, mention the appearance of limber pine with lodgepole pine at
7000 ft. It is not likely that Roth confused whitebark and limber pines,
since he noted elsewhere that whitebark pine is common above 7500 ft.
Western white pine, the only other white pine present in the Warners,
is a more likely candidate for this persistent but erroneous extension of
limber pine’s range into northern California.

PINUS ARISTATA Engelm. Bristlecone pine. Like limber pine, bristle-
cone pine is a characteristic tree of the basin ranges, but it is neither as
common nor as widespread as limber pine. From the southern Rocky
Mountains and the mountains of southern Utah it extends west to the
Deep Creek Range (McMillan, 1948) and the higher mountains of east-
ern and central Nevada (Billings, 1954; Little, 1956). In northern
Nevada it is known to occur on Sherman Mountain, at the southern end
of the Ruby Mountains (fig. 1J; Little, 1956), and on Spruce Mountain
in southern Elko Co. (Holmgren, 1942). On these two mountains bristle-
cone pine reaches its northwestern known limits. It was first collected in
the Ruby Mountains— presumably on Sherman Mountain—by Watson
in 1868 (‘‘East Humboldt Mountains,” Watson 1112, US), and it has
since been collected on Sherman Mountain by others (Hitchcock &
Martin 5686, POM). On Spruce Mountain, the upper slopes are occu-
pied by an extensive forest of this species and limber pine.

Bristlecone pine was also mapped in the East Humboldt Range of
Pershing Co. by Munns (1938). This erroneous range extension can be
attributed to Watson’s specimen from the ‘““East Humboldt Mountains,”
later called the Ruby Mountains.

PINUS MONOPHYLLIA Torr. & Frem. Singleleaf pinyon. This species is
the commonest member of the Pinaceae throughout much of the Great
Basin. Mixed with juniper (usually Juniperus osteosperma (Torr.) Little),
it occupies a broad woodland belt on the lower slopes of the mountains
in the central and southern Great Basin. In north-central Nevada, it
reaches its known northern limits south of the Humboldt River (fig. 1),
although older maps often show it north of the river in Elko Co. (Sud-
worth, 1913; Munns, 1938; Little, 1949). East of the Nevada-Utah
border it extends much farther north to the Raft River Mountains
(Preece, 1950) and southern Idaho (Johnson, 1966). Its range in Ne-
vada is shown in detail by Critchfield and Little (1966).

PINUS CONTORTA Doug]. Lodgepole pine. Two geographic races of this
widespread and variable species enter this region at places more than
300 miles apart. The open-cone Sierra Nevada-Cascade race, which has
been called P. contorta ssp. murrayana (Balf.) Critchfield (Critchfield,
1957), grows in the Warner Mountains. The Rocky Mountain race (P.
contorta ssp. latifolia (Engelm.) Critchfield), which often has serotinous

22 MADRONO [Vol. 19

cones, occurs in the mountains of southern Idaho as far west as the South
Hills, where it is a common tree. In the South Hills area lodgepole pine
stops a few miles short of the Nevada border (FS), although it is some-
times shown in the northeastern corner of that state (Munns, 1938).
Nor does this species extend south from Idaho to the Raft River Moun-
tains of northwestern Utah (Preece, 1950; FS), although it is shown
there on the most recent maps (Fowells, 1965; Critchfield and Little,
1966).

PINUS JEFFREYI Grev. & Balf. Jeffrey pine. This primarily California
species extends into this region only in the Warner Mountains. Although
Sudworth (1908) recorded Jeffrey pine ‘“‘at the sources of the Pitt [Pit]
River,” which originates in the Warners, it was not shown in the north-
eastern corner of California on older distribution maps (Sudworth, 1913;
Munns, 1938; Little, 1949). The species is locally common in the south-
ern Warner Mountains, and recent maps all record it there (Haller,
1962; Fowells, 1965; Critchfield and Little, 1966).

PINUS WASHOENSIS Mason & Stockwell. Washoe pine. This close rela-
tive of ponderosa pine is distributed along the western edge of the Great
Basin in a few scattered localities from Lake Tahoe north. It enters the
region that we are concerned with only in the Warner Mountains (Haller,
1961), where it intergrades with ponderosa pine. Haller, (1965) has
suggested that hybridization between the Pacific and Rocky Mountain
races of ponderosa pine may have played a role in the origin of this
doubtfully distinct species.

PINUS PONDEROSA Laws. Ponderosa pine. One of the most wide-ranging
trees in western North America, this species is unexpectedly absent from
a large area in the center of its range, including nearly all of the northern
Great Basin, northern Utah, southern and eastern Idaho, western Wyo-
ming, and southwestern Montana. The many occurrences shown in this
central region by Munns (1938) all appear to be in error. Baker and
Korstian (1931) attributed this large gap in the range of ponderosa to a
deficiency of moisture in the early part of the growing season.

The Rocky Mountain race of ponderosa pine (P. ponderosa var.
scopulorum Engelm.) does not enter the region considered here, although
it is present not far south in the mountains of eastern and southern Ne-
vada (Billings, 1954; Little, 1956). The Pacific race extends east to the
Warner Mountains and the Hart Mountain area of southern Oregon
(Sargent, 1897; Sudworth, 1908). It is also present in northwestern
Nevada in two localities that have not previously been reported.

One of these Nevada outliers (fig. 1K) is within a few miles of the
ponderosa stands in and near the Warner Mountains. In the extreme
northwestern corner of the state (41°59.5’ N Lat, 119°58.0’ W Long),
a scattered stand of this species grows along Twenty-mile Creek at an
elevation of about 5000 ft (BLM). It extends downstream a short
distance into Oregon.

The other and much smaller outlier in northwestern Nevada is at least

1969 | CRITCHFIELD & ALLENBAUGH: PINACEAE 1S

20 miles from the nearest known ponderosa pine (fig. 1L). This grove,
located on the west slope of Bald Mountain in the Sheldon National
Antelope Refuge, was reported to us by O. V. Deming of the Fish and
Wildlife Service, U. S. Department of the Interior (pers. comm.). It is
growing at an elevation of 6500 ft at 41°50.0’ N Lat, 119°38.5’ W Long.
It consists of one large tree (43 inches dbh, about 50 ft high) at least
300 years old, three smaller trees from 9 to 35 ft high and ranging in
age up to about 70 years, and a large number of scrubby seedlings. Ex-
cept for a few seedlings, the entire colony is confined to an outcropping
of whitish rhyolitic sand which appears to be extremely infertile. The
isolation of this grove and the distribution of tree ages suggest that all
of the younger trees in the colony are descendants of the single old tree.

DISCUSSION

The Pinaceae has a long history in the northern Great Basin. The four
genera represented in this region today, Abies, Picea, Pseudotsuga, and
Pinus, were present here 40 million years ago. All are elements of the
Eocene flora of the Copper Basin, near the Jarbidge Mountains in north-
ern Elko Co., Nevada (Axelrod, 1966). Several contemporary species
are represented by closely similar fossil species in this conifer-rich flora:
Abies grandis or A. concolor (A. sonomensis Axelrod), Picea engelmannii
(P. lahontense MacGinitie), Pseudotsuga menziesii (P. sonomensis
Dorf), Pinus aristata (2?) (P. crossiti Knowlton), and P. ponderosa (P.
harneyana Chaney & Axelrod). During the ensuing Miocene epoch,
several of these fossil species were also widespread on the Columbia
Plateau north of the Great Basin (Chaney, 1959).

The highly discontinuous and sporadic distribution of the Pinaceae in
this region today may have been influenced by events of the much more
recent Pleistocene epoch. During Pleistocene times the climate of the
Great Basin was periodically cooler and wetter than it is now, and these
climatic cycles are widely believed to have caused major changes in
plant distribution (Morrison, 1965). Lake Lahontan, a Pleistocene lake
that covered much of northwestern Nevada during the pluvial periods,
may have acted as a barrier to plant migration. The southern shoreline
of the lake during the last pluvial period coincides rather closely with the
present northern limits of Pinus monophylla (cf. Morrison, 1965, fig. 1;
cf. Critchfield and Little, 1966, map 16). The northern arms of the lake,
which extended to the Oregon border, may have restricted east-west mi-
gration. Cordilleran elements of the Pinaceae are confined to the region
east of the lake. Their westernmost representative is Pinus flexilis in the
Santa Rosa Range, just east of the Lake Lahontan shoreline. West of the
shoreline, in northwestern Nevada, the outliers of the Pinaceae all have
Pacific affinities with the possible exception of the Pinus albicaulis stand
in the Pine Forest Range.

A lake the size of Lahontan could not have been more than a partial
barrier to east-west migration across the Great Basin, however. It can-

24 MADRONO [Vol. 19

not entirely account for evidences of long-term separation between eastern
and western elements of the Pinaceae in this region. Four of the most
widespread species considered here have U-shaped ranges that border the
northern Great Basin on the east, north, and west. All four, Adbzes con-
color, Pseudotsuga menziesiu, Pinus ponderosa, and Pinus contorta, are
differentiated into well-defined geographic races on opposite sides of the
Great Basin. It is improbable that these races met without mixing during
the pluvials, since the eastern and western races of Pinus ponderosa
and of Abies concolor have proved to be highly compatible in crosses
made at the Institute of Forest Genetics, Placerville, California. Nor is
it likely that these races have evolved as recently as the last pluvial,
which is estimated to have been about 20,000 years ago (Martin, 1963).
Both the long generation interval of these trees and the geographic
extent of their races argue against this possibility. The alternative is that
the Great Basin long antedates the late Pleistocene as a barrier between
Pacific and Cordilleran segments of these conifers.

This conclusion is hard to reconcile with recent evidence of drastic
vertical and latitudinal plant migrations in the Great Basin and the
Southwest during the late Pleistocene. This evidence, much of it from
analyses of fossil pollen in the Southwest, has recently been reviewed by
Martin and Mehringer (1965). It supports the view that the disjunct
distributions of the montane conifers considered here are remnants of
former continuous distribution across the intervening basins.

A different interpretation of comparable patterns of distribution has
been advanced by Wells (1966) and Wells and Berger (1967). Their
investigations of late-Pleistocene macrofossils preserved in wood rat
middens in west Texas and the southern Great Basin have failed to un-
cover any evidence of substantial downward displacement of high-mon-
tane conifers during the last pluvial period. They attribute the disjunct
and sporadic distribution of these species to long-distance transport of
propagules, rather than former continuity. This view of Pleistocene
vegetational history provides some support for our interpretation of the
Great Basin as a long-term barrier in the evolutionary history of these
coniferous species.

Two range extensions in this region have been reported since this arti-
cle was prepared. S. A. Scott (FS) has collected Pinus albicaulis in the
Jack Peak area (41°30’ N Lat), extending the range of this species to the
Independence Mountains (S. A. Scott, 1967, IFGP). C. W. Ferguson of
the University of Arizona has found an susisine stand of Pinus aristata
near the summit of Pearl Peak in the Ruby Mountains (Deseret News,
Salt Lake City, Utah; Dec. 12, 1968), 7-8 miles north of the stand on
Sherman Mountain.

Pacific Southwest Forest and Range Experiment Station,
Forest Service, U.S. Department of Agriculture, Berkeley, California
University of California, Berkeley

1969] CRITCHFIELD & ALLENBAUGH: PINACEAE 25

LITERATURE CITED

Anon. 1944. Oregon travelogue written in wood. Forest Log 14(3): 7.

AXELROD, D. I. 1966. The Eocene Copper Basin flora of northeastern Nevada. Univ.
Calif. Publ. Geol. Sci. 59.

BaILey, V. 1936. The mammals and life zones of Oregon. North Amer. Fauna 55.
U.S. Dept. Agric., Washington, D. C.

Baker, F. S., and C. F. Korstian. 1931. Suitability of brushlands in the Intermoun-
tain Region for the growth of natural or planted western yellow pine forests.
Techn. Bull. U.S. D. A. 256.

Bituincs, W. D. 1951. Vegetational zonation in the Great Basin of western North
America. In Les bases écologiques de la régénération de la végétation des zones
arides. Int. Union Biol. Sci. Ser. B. No. 9.

. Nevada trees. Bull. Univ. Nevada Agric. Ext. Serv. 94 (ed. 2).

BraAysHAw, T. C. 1965. Comments on “Field identification of whitebark and limber
pines based upon needle resin canals.” J. Forest. (Washington) 63:705—706.
CuHaney, R. W. 1959. Miocene floras of the Columbia Plateau. Part I. Composition

and interpretation. Publ. Carnegie Inst. Wash. 617.

CRITCHFIELD, W. B. 1957. Geographic variation in Pinus contorta. Publ. Maria
Moors Cabot Found. Bot. Res. 3.

. 1966. A new conifer herbarium. Taxon 15:217-218.

. and E. L. Littie, Jr. 1966. Geographic distribution of the pines of the
world. U.S. D. A. Misc. Publ. 991.

Ericson, J. E. 1964. Field identification of whitebark and limber pines based upon
needle resin canals. J. Forest. (Washington) 62:576—-577.

FENNEMAN, N. M. 1931. Physiography of western United States. McGraw-Hill, New
York.

FoweE tts, H. A., (compiler). 1965. Silvics of forest trees of the United States. U. S.
D. A. Agric. Handbook 271.

Hatt, E. R. 1946. Mammals of Nevada. Univ. Calif. Press, Berkeley.

Hatter, J. R. 1961. Some recent observations on ponderosa, Jeffrey, and Washoe
pines in northeastern California. Madrono 16:126-132.

. 1962. Variation and hybridization in ponderosa and Jeffrey pines. Univ.
Calif. Publ. Bot. 34:123-166.

. 1965. Pinus washoensis in Oregon: taxonomic and evolutionary implica-
tions. Amer. J. Bot. 52:646.

Hormcren, A. H. 1942. A handbook of the vascular plants of northeastern Nevada.
Utah Agric. Exp. Sta., Logan.

Jounson, F. D. 1961. Native trees of Idaho. Bull. Univ. Idaho Agric. Ext. Ser. 289.

LitTLe, E. L., Jr. 1949. Important forest trees of the United States. In Trees. The
Yearbook of Agriculture. Dept. Agric., Washington, D. C.

. 1956. Pinaceae, Rutaceae, Meliaceae, Anacardiaceae, Tamaricaceae, Cor-
naceae, Oleaceae, Bignoniaceae of Nevada. Contrib. Fl. Nevada 40. Beltsville,
Maryland.

Martin, P. S. 1963. The last 10,000 years. A fossil pollen record of the American
Southwest. Univ. Arizona Press, Tucson.

.. and P. J. MEHRINGER, JR. 1965. Pleistocene pollen analysis and bio-
geography of the Southwest. Jn H. E. Wright, Jr., and D. G. Frey, (editors).
The Quarternary of the United States, Princeton Univ. Press, New Jersey.

McMirtan, C. 1948. A taxonomic and ecological study of the flora of the Deep
Creek Mountains of central western Utah. M. S. thesis, Univ. Utah, Salt Lake
City.

Morrison, R. B. 1965. Quaternary geology of the Great Basin. In H. E. Wright, Jr.,
and D. G. Frey (editors). The Quaternary of the United States. Princeton
Univ. Press, New Jersey.

Munns, E. N. 1938. The distribution of important trees of the United States.
U.S. D.A. Misc. Publ. 287.

26 MADRONO [Vol. 19

PREECE, S. J., JR. 1950. Floristic and ecological features of the Raft River Moun-
tains of northwestern Utah. M. S. thesis, Univ. Utah, Salt Lake City.
RussELL, I. C. 1885. Geological history of Lake Lahontan. U. S. Geol. Surv.
Monog. 11.
. 1903. Notes on the geology of southwestern Idaho and southeastern Ore-
gon. U.S. Geol. Surv. Bull. 217.
SARGENT, C. S. 1897. The silva of North America. Vols. 11, 12. Houghton, Mifflin,
Boston.
SupworTH, G. B. 1908. Forest trees of the Pacific Slope. U. S. Forest Serv., Wash-
ington, D. C.
. 1913. Forest Atlas. Geographic distribution of North American trees.
Part 1, U.S. Forest Serv., Washington, D. C.
. 1916. The spruce and balsam fir trees of the Rocky Mountain region.
US) D. AT Bull (1015-23) 327.
. 1918. Miscellaneous conifers of the Rocky Mountain region. U. S. D. A.
Bull. (1915-23) 680.
Taytor, W. P. 1912. Field notes on amphibians, reptiles and birds of northern
Humboldt County, Nevada (with a discussion of some of the faunal features
of the region). Univ. Calif. Publ. Zool. 7:319-436.
Warinc, G. A. 1909. Geology and water resources of the Harney Basin region,
Oregon. U.S. Geol. Surv. Water-Supply Paper 231.
Watson, S. 1871. Vol. 5, Botany. In C. King. Report of the United States geo-
logical exploration of the fortieth parallel. Washington, D. C.
Wetts, P. V. 1966. Late Pleistocene vegetation and degree of pluvial climatic
change in the Chihuahuan Desert. Science 153:970-975.
. and R. BERGER. 1967. Late Pleistocene history of coniferous woodland
in the Mohave Desert. Science 155:1640-1647.

LAURA M. LORRAINE, 1904-1968

ROXANA S., FERRIS

To be a good botanical collector does not mean that one must be a
professional botanist. The aesthetic enjoyment of seeing nature undis-
turbed and a desire to know something about the plants of meadows,
streams and forests stimulated Laura Lorraine to collect flowers to satisfy
this curiosity. During her college years at Stanford University, where she
received her Bachelor’s and Master’s degrees in Romance Languages, she
took a course on the classification of flowering plants with Dr. L. R.
Abrams, a course designed for both botanical and non-botanical students.
From this background of laboratory and field work, she acquired a most
rewarding hobby.

Our many botanical excursions together were profitable as well as
pleasant and extensive collections were added to the Dudley Herbarium
at Stanford. The specimens were collected in Northern California or the
Sierra Nevada. One of the pack trips in the Sierra yielded a new Lewisia
—Lewisia sierrae, collected at the headwaters of the south fork of the
San Joaquin.

1969] FERRIS: LAURA LORRAINE 27

Laura Lorraine was born in Michigan, December 12, 1904 and died in
Sebastopol, California, February 5, 1968. After receiving her master’s
degree in 1926, she became a teacher at the Analy High School in Sebas-
topol, and with the exception of a 3-year leave of absence, held that post
until her death. The leave was spent in New York as Executive Director
of the National Business and Professional Women. Her administrative
abilities were also evidenced during her long tenure at Analy High School
by her work with the California Scholarship Federation, at the state as
well as local level.

Dudley Herbarium, Stanford University

REVIEWS

The Population Bomb. By Pau R. Eur ticu. xiv + 223 pp., Ballantine Books,
New York. 1968. $.95.

An argument of extreme importance for biologists and the future of mankind is
raging within the rapidly-growing profession of futurology. One pole is represented
by Kahn and Wiener, who argue in The Year 2000: A Framework for Speculation
on the Next Thirty-Three Years that despite great increases in the world population,
a trouble-free world is the most likely prospect for the next few decades. The oppo-
site pole is represented by the book under review. A very large number of experts is
lined up on both sides, and it is important to understand why reasonably well-
informed people can have such remarkably different points of view. We will consider
the basic issues in Ehrlich’s book and explain why his critics err when they take
issue with him.

1. The most basic contention Ehrlich makes is that there will be large-scale star-
vation in the world in about a decade, involving considerably more than the present
3% million deaths per annum from this cause. Related to this point, he contends
that family planning has been a failure, the impressive new strains of the cereal
grains may lead to a variety of serious problems resulting in lower production than
expected, and the maximum amount of animal protein we can get from the world’s
oceans is only about double the present annual catch. The optimists point out, on
the other hand, that world rice production rose 12% in 1967, the take from the
oceans increased sharply in Japan and Norway last year, and family planning can
work, because the birth rate per capita is dropping sharply in the U. S. and else-
where. In this bewildering barrage of facts, claims, and counterclaims, who is right?
In general, there are three explanations for the discrepancies: successes tend to be
well publicized, whereas typically only the real experts know about failures or
omens of doom; there is a widespread tendency to underestimate the complexity of
many problems, so that important factors are ignored in making predictions; and
finally, many of the “experts” in this whole complex of problems are in fact only
experts on one aspect. Physicists may not realize how little they know about epi-
demiology, and agronomists may overestimate their grasp of the current situation in
fisheries, for example. In the case of the cereal grains, while the success with new
strains is well known, the international crisis developing because of pesticide resist-
ance in cotton pests is not. In general, few people grasp the basic ecological facts
about pesticides: they generally produce a less serious long-term effect on the popu-
lations of pests than on the populations of parasites and predators that would
normally control the pests, because of higher initial pest densities; as spray calendars
draw on a greater variety of chemicals, cross-resistance to a wide variety of chemi-
cals develops in the pests, so that there is rapid selection for pests that can withstand
anything that can be used against them; and no matter what the details are of a
contro] program, the long-term consequences of pest buildup are likely to be worse
where the largest possible acreage is planted to the same species and variety. One
of the best-informed books available on the future for cereal grains is Paddock and
Paddock’s Famine 1975!, for which the title gives the plot.

Two lines of argument support Ehrlich’s contention that oceanic production
cannot be enormously increased: a theoretical argument and one based on the
history of exploited marine resources. The former is that much of the world’s oceans
are aquatic deserts because of inadequate upwelling of minerals, and further, since
we typically harvest predators, there is great thermodynamic conversion inefficiency
in harvesting food pyramids several steps removed from incident solar energy. Also,
it would take too many calories per calorie obtained to make seiving plankton out
of sea water a profitable activity in most parts of the oceans. The historical argu-
ment is that given the current economic demand for animal protein and the long
experience exploiting most major oceanic stocks, how is it that many of these have
shown an actual decline in yield over the last several decades if we have only begun
to tap the resource ?

The magnitude of the impending population-unplanned family catastrophe can

1969] REVIEWS 29

best be grasped by studying a typical village in an underdeveloped country in detail,
as in Gilbert Etienne’s Studies in Indian Agriculture. To a considerable extent it
has been possible to prevent massive catastrophe up to the present by increasing
the proportion of all available land under cultivation. However, just about all land
that could possibly be cultivated is now under cultivation, yet the population keeps
increasing. Thus, mass famine can only be averted by increasing yield per acre, but
thermodynamic realities impose an upper asymptote on this figure.

2. Ehrlich notes that pandemics may be a great problem in the near future, a
point not made by many experts. The fact is that plague, for example, could break
out on a fearful scale in India. Precisely those conditions that allowed a resurgence
of plague in India in the 1940’s are found there again: large numbers of rats per
person moving freely through warehouses full of American grain shipments, with
inadequate efforts being made to kill the rats or board up holes. Plague can explode
with such speed that it could overwhelm public health organizations.

3. Ehrlich notes that we are seriously polluting the planet. The magnitude of
this problem is not well known. Smog from Los Angeles is having a serious effect
on plants far to the east of the surrounding mountains, and some authorities state
that it is now causing enormous losses in agricultural productivity. Emphysema
death rates are among the most rapidly rising variables on earth.

In general, it appears that more familiarity with existing data would show that
it is Ehrlich’s critics who err, not Ehrlich—Kennety E. F. Watt, Department of
Zoology, University of California, Davis.

Rocky Mountain Flora. By Witt1aAM A. WEBER. vii + 437 pp., 346 fig. Univer-
sity of Colorado Press, Boulder. 1967. $9.40.

“This book, a revision of the Handbook of Plants.of the Colorado Front Range,
culminates twenty years of intensive field and laboratory studies of the Rocky
Mountain flora.” It is a field guide to the “Ferns, conifers, and flowering plants of
the Southern Rocky Mountains from Pikes Peak to Rocky Mountain National Park
and from the plains to the Continental Divide. . . . Over 1,500 kinds of plants
[1,400 species, cf. p. 2] are keyed and classified. The book is small enough to be
carried in a rucksack [but not a pocket], and only a hand magnifier is needed to
make the necessary examinations.”

The introductory pages include a tantalizing review of why the vegetation of
the Front Range is more than a “green blur.” Weber points out that the flora of the
Southern Rockies includes some taxa that are circumpolar; reoccur in the mountains
of Central Asia; or are Tertiary relicts. The paragraphs on “Plant Geography” lead
one to expect an important digest, but instead one is abandoned with a parsimonious
list of examples on p. 6 and a reference to an earlier paper (Weber, W. A. 1965.
Plant geography in the southern Rocky Mountains. Jn H. E. Wright, Jr., and D. F.
Frey (editors). The Quaternary of the United States. Princeton Univ. Press, New
Jersey.) which will not be available to many for whom this book is intended. This
list, which could have kindled a lot of interest, is disappointing inasmuch as extra-
territorial occurrences in the text frequently are neglected. None of the three en-
demics mentioned on p. 6 is clearly indicated in the text as not occurring elsewhere;
one of these, Aletes acaulis, also occurs in New Mexico and Texas and must have
been a mistake for A. anisatus. The selection of Aralia racemosa as an example of an
“Eastern Woodland-prairie” cognate ought to have been explained as this species is
not otherwise included in this book.

The remainder of the book is organized like its predecessor, as a continuous series
of keys without the interjection of descriptions and the frills of endless measure-
ments and literature citations. The first key leads to helpful categories such as Para-
sites (here including saprophytes), Aquatics, Vines, Monocots, Woody Dicots, and

30 MADRONO [Vol. 19

Herbaceous Dicots. The families (and their genera) are arranged alphabetically
within major groups, with the monocots appearing last. This is very practical for
rapid finding of family and genus. The genera of Compositae are strictly alpha-
betical, but the grasses are arranged by tribe. Short commentaries on field obser-
vations are frequent with conscientious attention to ecology and distribution within
the Front Range. Occasional synonyms are given and there are references to the
second edition of the Handbook when there has been a change in name.

The illustrations by C. F. Yocom are an asset. The introduction and glossary use
78 of the 346 figures; thus, about 18 percent of the taxa are illustrated. The supple-
mental dissections or blow-ups shown beside the main drawings, which could have
been helpful to beginners, are nowhere explained.

Weber has succeeded in presenting a handy and attractive two-fisted means for
finding names of plants in the Front Range. The area where generally applicable is
substantially greater than that of the Front Range; but both the area and the style
are similar to those of the author’s earlier Handbook and it would have seemed
appropriate for this to have been the illustrated third edition without change of title.
This volume distills much personal experience and is a welcome addition to the
books on plants of the Rocky Mountains ——WALLace R. Ernst, Smithsonian Institu-
tion, Washington, D. C.

Taxonomy of Flowering Plants. 2nd ed. By C. L. Porter. ix + 472 pp., 400 plates
(311 individual and sets of line drawings, 88 black & white photographs, 1 color
photograph). W. H. Freeman and Company, San Francisco. 1967. $6.75.

The second edition of this popular text, now in a more readable print, is basic-
ally the same as the first edition, with few significant changes, but with numerous
small refinements. Since Mooring’s discriminating review of the contents and format
of the first edition (Madrono 16:171-172) could apply equally appropriately to the
second edition, this brief review pertains primarily to the refinements in the second
edition.

In Part I, which deals with History, Principles and Methods, a brief discussion of
chemical and numerical taxonomy has been added to the chapter on Concepts of
Taxa. 63 additional entries are found in the lists of references at the end of chap-
ters; charts have been improved in format, and boldface type has been substituted
for italics wherever definitions occur. Part II, which covers “Selected Orders and
Families of Monocotyledons” includes several additional examples of certain taxa
plus three subclass descriptions. In Part III, which deals with “Selected Orders and
Families of Dicotyledons,” further examples of taxa, along with keys to the fam-
ilies of the Ranales and the “Tribes of the Asteraceae (Compositae),’ have been
added. A floral diagram has been corrected, and a qualifying statement on the
Apetalae has been inserted.

Throughout the book, illustrations have been renumbered in groups with parts
a, b, c, etc., instead of each individual illustration being numbered consecutively ;
asian pack and white photographs have been added, and the clarity of most. of
the photographs has been improved.

The second edition, like the first, contains few errors, and, despite minor reserva-
tions about the author’s continued use of Fabaceae, ieee etc., instead of tra-
ditional family names, impresses this reviewer as being an excellent text, if not the
best available, for introductory taxonomy courses, especially those of less than a
year’s duration. The clear floral diagrams and line drawings will also be found very
useful in lengthier introductory courses, but instructors would probably want to
supplement the material on history and principles in such instances——KINGSLEY R.
STERN, Department of Biological Sciences, Chico State College, Chico, California.

1969] NOTES AND NEWS on

Handbook of Northwestern Plants. By Heten M. Gitkey and LA Rea M.
Dennis. 505 pp., illustrated. Oregon State University Bookstores, Inc., Corvallis,
Oregon 97331. 1967. $7.00.

Those of us who were botanically weaned on Gilkey’s “Handbook of Northwest
Flowering Plants” will welcome this edition of a most useful guide to the more con-
spicuous vascular plants of the Pacific Northwest. In this recent revision, Dr. Gilkey
has been ably aided by La Rea Dennis, assistant curator of the Oregon State Uni-
versity Herbarium. The keys are easy to use and the illustrations are good. The
concise descriptions of the plants are often accompanied by comments that make
the book interesting browsing. However, although I agree that the flower of
Calypso bulbosa has a “delicate lovely fragrance,” I still remain to be convinced
that it is the stems and leaves of Lysichitum americanum that are responsible for
its skunk-like odor rather than its “pleasantly fragrant” flowers.

The present edition of the book represents a substantial revision over earlier ones.
It is about 100 pages longer than the previous edition; this increased length is due
not only to the use of larger type, but also to the inclusion of additional taxa. The
book now includes vascular cryptogams, although earlier versions did not. This
expanded coverage accounts for the altered title. Illustrations of various species also
have been added, and the reproduction of most illustrations is superior to that in
earlier editions. Nomenclature changes are evident throughout the work: the former
Baeria maritima is now found as Lasthenia minor subspecies maritima; Cacalio psis
is a Luina; and in many other genera recent monographic work has been utilized.
Errors seem to be few. For example, although Oxalis stricta is keyed out, a descrip-
tion of the species is missing from the text.

The authors have done a good job in selecting species to be included in this
handbook, since it is not intended to be comprehensive. Readers are told to use
other references for identification of grasses and sedges. Weeds are particularly
prominent in the book, perhaps because they are likely to be picked up by amateurs
or agriculturalists. Although this book will appeal to amateur botanists or novices,
it will be of use to professionals in the region as well. Its small size makes it truly
a handbook; its simplicity of style and ease of usage insure its wide adoption by
northwesterners interested in naming the plants around them.—RoOBERT ORNDUFF,
Department of Botany, University of California, Berkeley.

NOTES AND NEWS

Zor. A small reside of back numbers of the biological journal, Zoe, published
by T. S. and Katharine Brandegee from 1890 until 1908 is available from the
Herbarium, Department of Botany, University of California, Berkeley. No charge
will be made except for postage. From Vol. I, numbers 1-4, 6, 8-10, and 12 are
missing; from Vol. IJ, numbers 1 and 2 are missing; Vols. II] and IV are complete;
and from Vol. V, number 1 is missing.

NEW DISTRIBUTION RECORD FOR CLAYTONIA NEVADENSIS FROM NORTHWESTERN
CALIFORNIA.—Chambers (Leafl. West. Bot. 10:1-8. 1963) reviewed the known dis-
tribution of Claytonia nevadensis Wats. citing specimens from a number of collec-
tions in the Sierra Nevada, the Mt. Lassen area and Steens Mountain in southeastern
Oregon. In the summer of 1967 I collected this species in Trinity Co., Calif., appar-
ently a new westward distribution_record. The plants were found growing in dense
clumps on wet, gravelly soil below. the permanent ice field on the north side of
Thompson Peak at an elevation of 7800 ft. There were several colonies observed in
the immediate area. The specimens collected are deposited in the Herbarium of
Humboldt State College (HSC), Ferlatte 907, August 11, 1967 —Wi111AM J. FEr-
LATTE, Division of Biological Sciences, Humboldt State College, Arcata, Calif.

32 MADRONO [Vol. 19

NEw PuBLICATIONS

Evolution of the Fern Genus Osmunda. By C. N. Mitter, Jr. Contributions from
the Museum of Paleontology, University of Michigan, 21:139-203. 1967.

The Book of Grass, An Anthology on Indian Hemp. Edited by GrorcE ANDREWS
and SIMON VINKENOOG. xiv + 242 pp. Grove Press, New York. 1967. $5.00.

The Lupines of Canada and Alaska. By Davin B. DUNN and JoHNn M. GILLETT. 89

pp. Canada Department of Agriculture, Research Branch, Monograph No. 2.
1966.

Key to the Native Trees of Canada. By T. C. BraysHAw. xix + 43 pp., illus. Bull.
125, Department of Forestry. Queen’s Printer, Ottawa. 1961. $0.50.

The Ancient Bristlecone Pine Forest. Edited by Russ and ANNE JOHNSON. 44 pp.,
illus. Chalfant Press, Inc., Bishop, Calif. 1966. $1.25.

Agrostology. By W. Epwin Boor. vi + 222 pp., illus. The Endowment and Re-
search Foundation, Montana State University, Bozeman, Montana. 1964.
Flora of the Cabeza Prieta Game Range. By NoRMAN M. Simmons. Journal of the

Arizona Academy of Sciences 4(2) :93-104. 1966.

John Torey, A Story of North American Botany. By ANDREW DENNY Rockers III.
xiii + 352 pp. (Facsimile of the edition of 1942.) Hafner Publishing Co., 31 E.
10th St., New York. 1967. $7.50.

The Wild Flowers of California. By Mary ELizBETH Parsons, with a new table of
changes in nomenclature by Roxana S. Ferris. cvi + 425 pp. (Facsimile of the
edition of 1907.) Dover Publications, New York. 1966. $2.25, paper.

Plants and Civilization. By HERBERT G. BAKER. vii + 183 pp., illus. Wadsworth
Publishing Co., Belmont, Calif. 1965.

A History of Botany in West Virginia. By WHELDON Boone. xi + 196 pp., illus.
McClain Printing Co., Parsons, West Virginia. 1965. $6.00. A very interesting
little book full of biographical information about the professional and amateur
botanists who have worked in West Virginia.

Place Names of Shasta County. By GERTRUDE A. STEGER, revised by HELEN HINCK-
LEY JONES. 71 pp. La Siesta Press, Glendale, California. 1966.

Notes on the Vegetation Zones of Western Canada, with Special Reference to the
Forests of Wells Gray Park, British Columbia. By LEENA HAmMeEtT-Autt. Annales
Botanici Fennici 2:274-300. 1965.

Plants of the White Mountains, California and Nevada. By Ropert M. Lioyp and
RicHArRD S. MITCHELL. iv. + 60 pp., mimeographed. Department of Botany
and White Mountain Research Station, University of California, Berkeley. Re-
vised edition, 1966.

An English-Classical Dictionary for the Use of Taxonomists. By RoBert S. Woops.
xiv + 331 pp. Pomona College, Claremont, California, 1966.

A Dictionary of the Flowering Plants and Ferns. By J. C. WItts, revised by
H. K. Atry SHaw. xxii + 1214 + liii pp. Cambridge University Press. 7th
edition, 1966.

Flora of Peru. Solanaceae. By DONOVAN S. CorRELL. Botanical Series Field Museum
of Natural History 13, part V-B(2) :271-458. 1967.

Skyline Landscape of the San Francisco Peninsula Cities. By KATHRYN STEDMAN.
36 pp. The Council for Foothill Planning and Research, P.O. Box 11511, Palo
Alto, California. 1966.

Skyline Scenic Parkway, San Francisco to Monterey, California. 24 pp. Loma Prieta
Chapter, Sierra Club. 1966.

Exploring Our Baylands. By Diane R. Conrapson and Howarp KInc. 60 pp. Palo
Alto Chamber of Commerce, Palo Alto, California. 1966.

A WEST AMERICAN JOURNAL OF BOTANY

A quarterly journal devoted to the publication of botanical research,
observation, and history. Back volumes may be obtained from the Sec-
retary at $12.00 per volume. Single numbers of Volumes 1 and 2 may be
obtained at $1.00 and of Volumes 3 through 18 at $1.50. Some numbers
are in short supply and are not available separately.

The subscription price of Madrofo is $6.00 per year ($4.00 for stu-
dents). If your institution does not now subscribe to Madrono, we would
be grateful if you would make the necessary request. Since there is no
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bership in the California Botanical Society on the basis of his institu-
tion’s subscription. Address all orders to: Corresponding Secretary,
California Botanical Society, Department of Botany, University of Cali-
fornia, Berkeley, California 94720.

INFORMATION FOR CONTRIBUTORS

Manuscripts submitted for publication should not exceed an estimated
20 pages when printed unless the author agrees to bear the cost of the ad-
ditional pages at the rate of $20 per page. Illustrative materials (includ-
ing “typographically difficult” matter) in excess of 30 per cent for papers
up to 10 pages and 20 per cent for longer papers are chargeable to the
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the author assume the complete cost of publication.

Shorter items, such as range extensions and other biological notes,
will be published in condensed form with a suitable title under the general
heading, “Notes and News.” |

Institutional abbreviations in specimen citations should follow Lanjouw
and stafleu’s list (Index Herbariorum. Part 1. The Herbaria of the World.
Utrecht. Fifth Edition, 1964).

Abbreviations of botanical journals should follow those in Botanico-
Periodicum-Huntianum (Hunt Botanical Library, Carnegie-Mellon Uni-
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Membership in the California Botanical Society is normally considered
a requisite for publication in MADRONO.

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A WEST AMERICAN JOURNAL OF BOTANY

VOLUME 20, NUMBER 2 APRIL, 1969

Contents

THE XEROPHYTIC CUCURBITA OF NORTHWESTERN
MEXICO AND SOUTHWESTERN UNITED STATES,
W. P. Bemis and Thomas W. Whitaker

Some New Species, NEw CoMBINATIONS, AND
NEw ReEcorps oF RED ALGAE FROM THE
Paciric Coast, /sabella A. Abbott

A New NAME FOR A SPECIES OF POLYPODIUM FROM
NORTHWESTERN NortH AMERICA, Frank A. Lang

THE PyGmMy Forest-PopsoL ECOSYSTEM AND ITS
DUNE ASSOCIATES OF THE MENDOCINO COAST,
H. Jenny, R. J. Arkley, and A. M. Schultz

New REcorRDS OF MYXOMYCETES FROM OREGON. L.,
Dwayne H. Curtis

Reviews: ARTHUR CRONQUIST, The Evolution and
Classification of Flowering Plants
(Lincoln Constance); Eric Huxtéen, Flora
of Alaska and Neighboring Territories
(Kenton L. Chambers)

S£? 03 1969

Ligmanicc. 6

-JBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established
1916. Subscription price $6.00 per year ($4.00 for students). Published quarterly in
January, April, July, and October by the California Botanical Society, Inc., and
issued from the office of Madrofo, Herbarium, Life Sciences Building, University of
California, Berkeley, California. Orders for subscriptions, changes in address, and un-
delivered copies should be sent to the Corresponding Secretary, California Botanical
Society, Department of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS

LyMAN BENSON, Pomona College, Claremont, California
Kenton L. CHAMBERS, Oregon State University, Corvallis
Joun F. Davinson, University of Nebraska, Lincoln
WALLACE R. Ernst, Smithsonian Institution, Washington, D. C.
ARTURO GOMEZ PompPaA, Universidad Nacional Autonoma de México
EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia
Mitprep E. Maruias, University of California, Los Angeles
ROBERT ORNDUFF, University of California, Berkeley
Marion OwnBEY, Washington State University, Pullman
Duncan M. Porter, Missouri Botanical Garden, St. Louis
REED C. Rotiins, Harvard University, Cambridge, Massachusetts
IrA L. Wiccrns, Stanford University, Stanford, California

Editor — JoHn H. THomas
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCasKILL
P. O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Harry Thiers, Department of Ecology and Systematic Biology, San
Francisco State College. First Vice-President: Grady Webster, Department of Bot-
any, University of California, Davis. Second Vice-President: Ernest C. Twisselmann,
Cholame, California. Recording Secretary: John West, Department of Botany, Uni-
versity of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Depart-
ment of Botany, University of California, Berkeley. Treasurer: June McCaskill, De-
partment of Botany, University of California, Davis.

THE XEROPHYTIC CUCURBITA OF NORTHWESTERN
MEXICO AND SOUTHWESTERN UNITED STATES

W. P. Bemis and THomMAs W. WHITAKER

The genus Cucurbita is indigenous to the American continents. The
center of origin probably is the tropical and semitropical regions of south-
ern Mexico (Whitaker and Bemis, 1964). In the hot, arid deserts of
northwestern Mexico and southwestern United States there occur a group
of xerophytic species, possibly derivatives of populations from mesic
ancestors that became adapted to climatic changes toward greater aridity.
These species essentially are isolated geographically from other species of
Cucurbita, and are well adapted to the highly specialized habitat to
which they are currently restricted.

The restricted xerophytic species are C. cylindrata Bailey, C. cordata
Wats., C. palmata Wats., and C. digitata Gray. Although a fifth species,
the wide ranging and variable C. foetidissima HBK., is truly xerophytic,
it is only distantly related to the four restricted species. Another species,
C. pedatifolia Bailey from central Mexico, has some strong xerophytic
characters, and may represent a transitional stage to the xerophytic con-
dition. This paper is concerned with some of the biological characteristics
of the restricted species that fit them for their specialized environment
and their genetical relationships to each other and to other species of
Cucurbita.

RANGE AND HABITAT

These species are confined to the extreme southwestern portions of the
United States and adjacent Mexico. This area comprises some of the
hottest and most arid locations on the North American continent. The
general area is characterized not only by low average rainfall but by rela-
tively prolonged precipitation-free periods; 6 to 10 consecutive months
without rainfall are not unusual. At Bahia de Los Angeles in Baja Cali-
fornia, Mexico, where C. cordata has been collected, a rain-free period
of 23 consecutive months has been recorded (Hastings, 1964).

The xerophytic species must be well adapted to their rugged environ-
ment in order to survive. These species generally grow in loose, gravelly,
well-drained soils below 4,000 feet elevation. The banks or the flood
plains of dry, sandy washes are favorite habitats. Such habitats are
normally free of stringent competition from other species, and because of
their location are likely to receive relatively more moisture than other
locations in this uncongenial environment.

Figure 1 shows the range or collection sites of the four species. Cucur-
bita cylindrata is found only in Baja California, mostly in Baja Cali-
fornia del Sur, or in about the middle portion of the peninsula. Cucurbita
cordata has been found only around Bahia de Los Angeles in Baja Cali-

Maprono, Vol. 20, No. 2, pp. 33-80. August 11, 1969.
33

34 MADRONO [Vol. 20

CALIFORNIA
ARIZONA
ta
se
oe
digjtata
\
VN9itoy,
AN
f) MEXICO
6
4 oat
2". :
8
4.
1 = C. cordata S'\¢

2-8 C. cylindrata

Fic. 1. Distribution of Cucurbita palmata, C. digitata, C. cordata and C. cylindrata.

fornia del Norte. It may be that C. cylindrata and C. cordata are sym-
patric in the central portion of Baja California. This suggestion, however,
requires confirming data from more thorough botanical exploration of
the peninsula.

The ranges of C. palmata and C. digitata are more extensive. Cucurbita
palmata extends from the northeastern portion of Baja California through
California into the San Joaquin Valley and lower portions of the Salinas
Valley and eastward to near the Colorado River. Cucurbita digitata ex-
tends from northern Sonora, Mexico, into southern Arizona and New

1969 | BEMIS & WHITAKER: CUCURBITA 35

C. palmate
C. cordata

C. digitata eee aes _ ©. eytindrata

Fic. 2. Typical shape of mature leaves of Cucurbita palmata, C. cordata, C.
digitata and C. cylindrata.

Mexico. Cucurbita digitata also is found at higher elevations in northern
Baja California. The ranges of C. digitata in southern Arizona and north-
ern Baja California are separated by C. palmata. These two species are
sympatric at the periphery of their ranges and in these locations inter-
specific hybridization between them was observed (Bemis and Whitaker,
1965).

MoRPHOLOGICAL DIFFERENCES

A numerical taxonomic study was conducted by Rhodes, e¢ al. (1967)
in which 93 plant characters were scored for 21 different species of
Cucurbita. The four xerophytic species clustered together at a significant
level in the eight different statistical techniques that were employed. This
indicates a relatively close morphological relationship between them
when compared with other species of Cucurbita. There are, however,
morphological differences among the four xerophytic species.

36 MADRONO [Vol. 20

The most obvious differences among these four species are the shapes -
of the leaf blades. The first 2 to 4 true leaves of the seedlings (juvenile
leaves) are similar in appearance in all four species. The older leaves,
however, are quite different (fig. 2). In Cucurbita digitata the narrow
lanceolate lobes extend almost to the base of the leaf blade; in C. palmata
the lobes are broad, and about twice as long as they are wide; in C.
cordata the leaf lobes are again broad extending about one-third the
depth of the blade; and finally in C. cylindrata the lobes are deep, narrow
and obtuse.

There are many other subtle differences between these four species.
Among the more marked dissimilarities are the size and shape of the
large storage roots. Cucurbita digitata has a large turnip-shaped taproot;
in C. palmata the storage root is stout, fusiform, and bifurcated or even
trifurcated a few inches below the crown. Cucurbita cordata and C.
cylindrata each have large fusiform roots. Dittmer and Talley (1964)
suggest that the roots of the xerophytic Cucurbita are adapted for extract-_
ing the moisture from the soil to a depth of not greater than 4 feet. Root
penetration is such that there is no possibility of obtaining moisture
from deep subsurface water tables. Excluded from this source of moisture,
the root system by means of highly developed laterals is well adapted for
gathering and storing moisture from the upper 4 feet of soil, and retain-
ing it for an indefinite period of time.

The small gourd-like pepos produced on the vines are similar in shape
for each of the xerophytic species, being nearly round. All are distinctly
striped but their color patterns differ as follows:

C. cylindrata Dark green not yellow at maturity
C. cordata Gray green not yellow at maturity
C. palmata Diffuse green mottle yellow at maturity
C. digitata Clear green mottle yellow at maturity

The fresh weight of the pepos range from 150 to 330 grams with C.
cylindrata usually having the smallest pepos of the four species. The
pepo rind is thin, but relatively hard. The narrow band of white flesh is
fibrous and when exposed to the desert environment dries into a network
of fibers. The pepos are five-carpelate except for C. digitata which has
three to five with a mean of 4.5 + .8 carpels. This characteristic of five
carpels (consistent for C. cordata, C. cylindrata and C. palmata) is
unique in the genus inasmuch as all of the other species of Cucurbita
are predominantly tri-carpellate. The five carpel character is basically
recessive but not completely so because the F, hybrids between C.
moschata « C. digitata and C. moschata * C. palmata produce pepos
with carpel numbers of 3.4 + .6 and 3.6 + .8, respectively. Cucurbita
moschata Duch. ex Poir. is consistently tri-carpellate (Bemis, 1963).

A single pepo often contains more than 500 seeds. The seeds of C.
cordata and C. cylindrata are smaller than those of C. digitata and C.

1969 | BEMIS & WHITAKER: CUCURBITA By

palmata. Room dry weight of 100 seeds for the species is 2.8 + .3,
2.8 + .7 and 5.4 + .5,4.9+ .7 grams, respectively. The seed shape and
seed coat color are similar for all four species.

The rate of pepo enlargement is rapid for these xerophytic species.
The mean diameter of the ovary of C. palmata at the time of pollination
is 1.6 cm. The rate of enlargement follows a typical sigmoid growth
curve which begins to decrease in rate about the 8th day after pollina-
tion. The mean maximum pepo diameter of 7.0 cm. was reached on the
12th day after pollination. The rate of seed development (weight),
however, is slower than the rate of pepo enlargement. Maximum seed
weight is not reached until 34 days after pollination (Ba-Amer, 1967).

REPRODUCTION

Observations in the field, experimental garden, and greenhouse sug-
gest that the vegetative method of reproduction in the xerophytic Cucur-
bita is the most common. Each node of a runner is capable of producing
adventitious roots, especially if covered with soil. In time the inter-
vening internodes decay. The net result is a group of vegetatively pro-
duced plants surrounding a mother plant, and the entire community
forms a roughly circular colony. For example, from a single plant of
C. foetidissima in the experimental garden, a colony of about 705 indi-
vidual plants was vegetatively produced within the course of three grow-
ing seasons.

Our observations in the field suggest that plants of the xerophytic
Cucurbita regularly produce quantities of ripe fruit with numerous viable
seeds, but few seedlings and even fewer mature plants originate from
this source. The seeds are an attractive and nutritious source of food for
desert animals, particularly rodents. It is evident that only a few seeds
survive their depredations. Those that do survive and become young
seedlings are vulnerable to browsing mammals, insect attacks, and
drought. Thus sexual reproduction is erratic, undependable, and only
infrequently results in a mature plant.

The seeds exhibit delayed dormancy apparently associated with the
relatively tough seed coat. Some seeds from a mature fruit, properly
ripened, will germinate readily, while others do not unless the seed coat
is removed. This delayed dormancy may have significance for the survival
of the species, but the phenomenon is not well understood.

These observations suggest that the xerophytic Cucurbita are adapted
to cope reproductively with the rugged environment mainly through a
modified version of conventional vegetative reproduction. Such a system
allows only limited genetic variation.

POLLINATORS
Paul D. Hurd, Jr., and his colleagues have made important mono-
graphic studies of squash bees of the genera Peponapis and Xenoglossa
(Hurd and Linsley, 1964; 1966). Females in species of these genera are

38 MADRONO [Vol. 20

specifically dependent upon particular species of Cucurbita for their |
pollen nutrition. Thus species limitation of these insects for pollen be- |
comes a powerful taxonomic tool for studying species relations in the
Cucurbita. Hurd informs us that it is indeed unusual to find a species-to- |
species relationship of plants and insects within a single genus of plants. _
This relationship permits a new approach to the systematics of Cucurbita. |
Already some exciting results are commencing to appear. |

Squash bees restricted to Cucurbita palmata and C. digitata for their |
pollen supplies are Peponapis timberlaket, Xenoglossa angustior and X.
strenua. Hurd has no data for the Baja California species, C. cordata and |
C. cylindrata, but he judges from the distribution of squash bees in this |
area that the same bee species as those dependent upon C. palmata and
C. digitata will also need C. cordata and C. cylindrata for their pollen
economy, with one exception. For those in the southern half of the
peninsula, X. angustior would be replaced by the endemic X. mustelina.
As would be expected, a number of species of squash bees are dependent
upon the widely ranging C. foetidissima as follows: Peponapis pruinosa,
P. smithu, Xenoglossa angustior, X. kansensis, X. patricia and X. strenua.

These observations reinforce support for a close relationship between
the four xerophytic species from California, Arizona, and Baja California,
and the wide divergence of C. foetidissima from this group.

COMPATIBILITY

The four xerophytic species of Cucurbita hybridize readily in all com-
binations with little if any decrease in fertility in the F, plants. The F,
plants have great vegetative vigor, are self-fertile, and fertile in back-
crosses to the parent species.

Where these species are sympatric they tend to hybridize, usually
along the perimeter of their range as we have shown for Cucurbita pal-
mata and C. digitata (Bemis and Whitaker, 1965). It is probable that
future botanical exploration in Baja California will uncover a similar
situation where the ranges of C. cordata and C. cylindrata overlap in
about the central portion of the peninsula.

The chromosome number of these four xerophytic species is m = 20.
This number is consistent for all species of Cucurbita examined thus far.
The homology of their respective chromosome complements was verified
by cytological examination of C. cylindrata, C. digitata, C. palmata and
their reciprocal F, hybrids (Groff & Bemis, 1967a). Occasional uni-
valents were observed, but their frequency was the same whether they
occurred in a species, or a species hybrid. Hybrids with C. cordata were
not examined cytologically, but the fertility of the hybrids suggests that
its chromosome complement is homologous with the other three species.

Extensive attempts to hybridize these species with other Cucurbita
yielded few or no seeds. The only success recorded was with Cucurbita
moschata, a cultivated species, when it was used as the pistillate parent
in matings with C. digitata and C. palmata. The partially developed

1969 | BEMIS & WHITAKER: CUCURBITA 39

embryos from these matings were artificially cultured in order to produce
the hybrid plants (Bemis & Nelson, 1963). The hybrids were vegeta-
tively vigorous, but completely sterile. Sixty-three pollen mother cells
at MI were examined from the F, hybrid C. moschata C. digitata
and the frequency of univalents was 36.4 + 2.9, 2n = 40 (Groff and
Bemis, 1967b). These observations suggest an almost complete lack of
chromosome homology between these xerophytic species and C. moschata.

BIOCHEMISTRY

Some significant biochemical information relating to the four xero-
phytic species of Cucurbita has been recorded by Chisholm and Hopkins
(1966), and confirmed and expanded by Bemis, ef al. (1967). In a
chemical analysis of the seeds of C. digitata and C. palmata Chisholm and
Hopkins found the conjugated fatty acid, punicic acid, in relatively large
amounts (17% —C. digitata; 11%—C. palmata). They state: “C. digi-
tata and C. palmata are believed to be the only plants native to North
America that have been shown to produce punicic acid.”

Bemis, e¢ al. (1967) added C. cordata with 8.99@ punicic acid, and a
recent analysis of C. cylindrata seeds shows this taxon to have 21.4%
punicic acid. Cucurbita foetidissima had only .36% punicic acid, while
in 12 other species of Cucurbita the punicic acid ranged from .05% to
33%. This unique occurrence of punicic acid in significant amounts in
the seeds of the four xerophytic Cucurbita and its virtual absence in
seeds of other Cucurbita, including the xerophytic C. foetidissima, again
signifies the close relationship of this group of species.

DISCUSSION

The xerophytic Cucurbita species, C. cylindrata, C. cordata, C. digitata
and C. palmata represent an extreme evolutionary divergence in the
genus. If our assumption is correct that the center of distribution of
Cucurbita is the tropical and semitropical region of southern Mexico,
then the xerophytic Cucurbita are terminal ecotypes adapted and appar-
ently restricted to the hot, dry environments of the deserts in western
North America. The evolution of the xerophytic Cucurbita from the
mesophytic progenitors probably is recent. Evidence for this is that the
chromosome number has remained constant, the monoecious character
of the plants and the simple inflorescence also have remained constant;
but more important is the limited compatibility that still exists with the
cultivated species, C. moschata.

The morphological characters of the xerophytic Cucurbita are either
unique for the species or are extreme modifications of certain characters.
Among the unique characters are the consistent five-carpelate nature of
the pepos with the exception of C. digitata which still has some variation.
The other species of Cucurbita are predominantly tri-carpellate. The pis-
tillate and staminate flowers differ from those of mesophytic species. The
pistillate flowers have a pronounced tubular corolla which encloses a long

40 MADRONO [Vol. 20

Style with a five-lobed compound stigma. The staminate flowers are also
tubular enclosing a long fused anther column.

The flowers are conveniently adapted to protect the solitary squash
bees which inhabit the flowers on the day they blossom. The symbiotic
relationship between species of Cucurbita and certain of the squash bees
is unusual. The bees are dependent upon the Cucurbita for their pollen
nutrition. This need for food from a specific group of plants restricts the
distribution of the insects.

The large storage roots of the xerophytic species represent an extreme
modification of the tuberculate root of transitional species such as C.
pedatifolia. The development of the tuberous roots has resulted in the
perennial nature of the xerophytic species and the predominantly vegeta-
tive mode of reproduction, which in turn restricts the genetic variation
because of reduced sexual reproduction. Likewise the relatively high
amounts of punicic acid in the seeds found only in the four xerophytic
species suggests a close genetic relationship.

The wide ranging C. foetidissima, while having xerophytic properties,
is not adapted to the extremes of environment of the four species dis-
cussed here. Differences in morphological and biochemical characters,
specific pollinators, and incompatibility suggest that C. foetidissima and
the other four xerophytic species diverged at an early date in the evolu-
tion of the genus.

The evidence presented in this report probably would lead the trained
taxonomist to consider these four taxa as subspecies, if the similarities
are considered more significant than the differences between them. In
our opinion the subspecies category would be satisfactory.

SUMMARY AND CONCLUSIONS

1. Cucurbita cordata, C. cylindrata, C. palmata, and C. digitata are
adapted to the hot arid regions of the Sonoran desert, primarily through
a modification of their root systems.

2. Modification of the root system has led to a change in growth habit
from annual to perennial, and to an efficient system of vegetative repro-
duction which in effect restricts genetic variation.

3. These four species are cross compatible and have many morpho-
logical similarities.

4. Two of the four species are specific pollen hosts for three species
of squash bees, Peponapis timberlakei, Xenoglossa angustior, and X.
strenua. Host specificity of squash bees is a highly sophisticated measure-
ment of relationship among Cucurbita species.

5. Certain morphological and biochemical characters unique in the
genus separate them from other species of Cucurbita. The extreme nature
of the habitat they occupy suggests they are terminal ecotypes.

6. Limited compatibility with C. moschata relates these species to the
annual, mesophytic, cultivated members of the genus.

1969 | BEMIS & WHITAKER: CUCURBITA 41

ACKNOWLEDGMENTS

We are heavily indebted to Paul D. Hurd, Jr., Department of En-
tomology, University of California, Berkeley, and Harlan Lewis, Division
of Life Sciences, University of California, Los Angeles, for offering much
helpful criticism and many pertinent comments.

Department of Horticulture, University of Arizona, Tucson

Crops Research Division, Agricultural Research Service,
U.S. Department of Agriculture, La Jolla, California

LITERATURE CITED

Ba-AM_Er, M. A. 1967. The effect of fruit maturity on seed development in certain
xerovhytic species of Cucurbita L. M. A. thesis, Univ. Arizona, Tucson.

Bemis, W. P. 1963. Interspecific hybridization in Cucurbita II. C. moschata Poir.
* xerophytic species of Cucurbita. J. Heredity 54:285-289.

—_—., J].M. Berry, M. J. Kennepy, D. Woops, M. Moran, and A. J. DEUTSCH-
MAN, JR. 1967. Oil composition of Cucurbita. J. Amer. Oil Chem. Soc. 44:429-
430.

. and J. M. Netson. 1963. Interspecific hybridization within the genus
Cucurbita I. Fruit set, seed, and embryo development. J. Arizona Acad. Sci.
2:104—-107.

. and T. W. Wuiraker. 1965. Natural hybridization between Cucurbita
digitata and C. palmata. Madronfio 18:39-47.

CrisHoim, M. J., and C. Y. Hopxins. 1966 Kamlolenic acid and other conjugated
fatty acids in certain seed oils. J. Amer. Oil Chem. Soc. 43:390-392.

Dittmer, H. J., and R. P. Tarrey. 1964. Gross morphology of tap roots of desert
cucurbits. Bot. Gaz. (Crawfordsville) 125:121-126.

GrorF, D., and W. P. Bemis. 1967a. Bivalent frequencies in interspecific hybrids
from three xerophytic species of Cucurbita. Manuscript in preparation.

————., and W. P. Bemis. 1967b. Meiotic irregularities in Cucurbita species hy-
brids. J. Heredity 55:109-111.

Hastincs, R. J. 1964. Climatological data for Baja California. Univ. Arizona Inst.
Atmos. Physics Techn. Rep. 14.

Hurp, P. D., Jr. and E. G. Linstey. 1964. The squash and gourd bees—genera
Peponapis Robertson and Xenoglossa Smith—inhabiting America north of
Mexico. (Hymenoptera: Apoidea). Hilgardia 35:375-477.

. and E. G. Linsey. 1966. The Mexican squash and gourd bees of the genus
Peponapis (Hymenoptera: Apoidae). Ann. Entomol. Soc. Amer. 59:835-851.
Ruopes, A. M., W. P. Bemis, T. W. Wuitaker, and C. G. CarMer. 1968. A nu-

merical taxonomic study of Cucurbita. Brittonia 20:251-266.

Wuitaker, T. W. and W. P. Bemis. 1964. Evolution in the genus Cucurbita. Evo-

lution 18:553-559.

SOME NEW SPECIES, NEW COMBINATIONS, AND NEW
RECORDS OF RED ALGAE FROM THE PACIFIC COAST

ISABELLA A, ABBOTT

The algae described in this paper come from two Pacific Coast areas
where intensive collections have been made, the Monterey Peninsula,
California (Smith, 1944; Hollenberg and Abbott, 1966) and Coos Bay,
Oregon (Doty, 1947). This paper forms a part of a continuing study
by the author of red algae, particularly from the Monterey area, and of
studies of other new taxa with George J. Hollenberg (Hollenberg and
Abbott, 1965; 1966; 1968).

Three species of Delesseriaceae (Ceramiales) are here newly de-
scribed: Nitophvllum dotyi from Oregon, N. cincinnatum from Cali-
fornia and Cryptopleura rosacea from California. Nitophyllum hollen-
bergu (Delesseriaceae) is a transfer from Myriogramme. In the Gigar-
tinales, Ozophora J. Agardh (Phyllophoraceae) is reinstated with O.
californica J. Agardh as the type species, and O. latifolia and O. norrisu
as newly described. Chondrus ocellatus Holmes (Gigartinaceae), known
from northern Japan, is reported from Sunset Bay, near Cape Arago,
Oregon, and is the first species of this genus to be reported from the east-
ern Pacific since Kylin (1928) removed Chondrus affinis Harvey to
Rhodoglossum.

Specimens without an herbarium abbreviation are at the Hopkins
Marine Station. Abbreviations for herbaria are the standard ones and the
following ones: GMS, Gilbert M. Smith algae herbarium at the Hopkins
Marine Station, and MSD, collection of Maxwell S. Doty, University of
Hawaii. Collecting numbers, except where preceded by a name, are those
of the author. Color names which are capitalized in the description are
those of Ridgeway (1915).

Nitophyllum hollenbergii (Kylin) Abbott, comb. nov. Myriogramme
hollenbergi Kylin, Acta Univ. Lund. 27(11): 32, pl. 11. 1941.

Nitophyllum is distinguished from Myriogramme on very technical
erounds. Vegetatively they are for the most part monostromatic and lack
veins of any kind except perhaps basally. One of the reproductive dif-
ferences emphasized by Kylin (1956) is the terminal carpospores of
Nitophyllum, whereas those of Myriogramme are in chains.

The lectotype of M. hollenbergi, Hollenberg 2854, GMS, which is cysto-
carpic shows carpospores that are terminal.

Other reference in the literature to Mvriogramme hollenbergu are
Smith (1944), Dawson (1962), and Norris and West 1966).

Distribution: British Columbia, Vancouver I.; Washington, near Smith
I., Partridge Bank off Whidbey I. Oregon, Tegula Bay, south of Cape
Arago, Doty 25764, MSD. Several new collections from Monterey, Cali-

— 1969] ABBOTT: RED ALGAE 43

fornia, were made in the vicinity of the previously known localities
(Smith, 1944). Dawson (1962) has reported this species from La Jolla,
California, and Isla Magdalena, Baja California.

Nitophyllum cincinnatum Abbott, sp. nov. Fig. 1. Thallus (in algis
corallineis) epiphyticus, membranaceus, per hapteron parvum, et secundo
gradu per extensiones paxilliformes cellularum thalli inferiorum affixus,
usque ad 6 cm alt., lobatus, lobis in circulis cristatis aggregatis, omni
lobo cuneato, latitudine apicum aequa altitudinis thalli aut dimidio brev-
iore; interdum prolifero, marginibus crebre corrugatis, corrugatienibus
crispatis, et undulatis et fimbriatis. Thallus monostromaticus, sine venis
microscopicis, rubro-purpureus. Cystocarpi 1.0-1.5 mm diam., gonim-
oblastus sporas unicas terminales, 25-28 « 12-15 u habens. Sori tetra-
sporangiales c. 0.5 mm diam., longiores quam lati, in centro altiores quam
ad margines.

Thallus epiphytic on corallines, membranous, attached by a small
holdfast and secondarily by occasional peg-like extensions of the lower
thallus, up to 6 cm high, lobed, lobes occurring in circular to trumpet-
shaped groups, each lobe cuneate, the width of the tops equal to or one-
half the height, occasionally proliferous, with densely ruffled margins,
the ruffles crisp, both undulate and fimbriate. Thallus monostromatic,
with no microscopic veins. Color when fresh reddish-orange (Russet
Vinaceous to Sorghum Brown), reddish purple when dry (Deep Corinth-
ian Purple). Cystocarps 1.0 to 1.5 mm across, gonimoblast with single ter-
minal spores, 25-28 by 12.5 u. Tetrasporangial sori about 0.5 mm in
diameter, longer than wide, and center higher than at the edges.

Holotype: California, Monterey County, cast ashore at the south end
of Carmel Beach, Abbott 2029, May 12, 1961, GMS, 6 specimens tetra-
sporangial, 2 cystocarpic on | sheet.

Other specimens: California, San Mateo Co., attached at 15 ft lepth
off Moss Beach, 4130, on Calliarthron; Monterey Co., at 20-25 ft depth
off Whaler’s Cove, Pt. Lobos State Reserve Park, 4135, on Calliarthron;
Monterey Co., 4138, US; 4129, WTU; 4141, UC; 4134, UCSB; 4130,
MSD; 41731 MSD.

Nitophyllum cincinnatum is named for the deep ruffling and curling
of the thallus. When first collected, it seemed to match the description
of N. mirabile Kylin (1925) from the Friday Harbor region, but an
examination of specimens of that species showed that the deep ruffling
and size of the new species were different from the northern species.
Nitophyllum mirabile, further, is a relatively flat blade with undulate
margins, whereas NV. cincinnatum is funnel or trumpet shaped, resem-
bling in form the medusoid genus Haliclystus. As described, it is the only
species of Nitophyllum of this shape on this coast, NV. mirabile being a
broad flat blade with ruffled margins, NV. northii strap-shaped without
ruffles, and N. hollenbergii small broad blades, without a modified
margin.

44 MADRONO [Vol.20 |

Fics. 1-4. Nztophyllum sp. and Cryptopleura rosacae: 1. habit of Nztophyllum
cincinnatum showing cystocarps on the crisp, ruffled lobes of the thallus, x 34; 2.
Nitophyllum dotyi, the type specimen which is a cystocarpic thallus, x 34; 3-4,
Cryptopleura rosacea; 3, habit of Abbott 4190, the type specimen, showing the lobes
arranged in clumps which are characteristic of this species, x 1%; 4, three segments
in detail, showing dense ruffling on margins, X 1.

Nitophyllum dotyi Abbott, sp. nov. Fig. 2. Thallus 8 cm alt., mem-
branaceus, roseus, in aliquot lobos primarios divisu, omni lobo ad dimi-
dum altitudinis loborum primariorum vicissim diviso, marginibus undu-
latis proliferisque. Thallus omnio monostromaticus, nisi ad basem, sine
venis micro-aut macroscopicis. Cystocarpi 300-500 p lat., carposporae
terminales, ellipticae ad obovatas, 1.5-2.0 plo longiores quam latae.
Tetrasporangia spermatangiaque non visa.

Thallus 8 cm tall, membranous, rose-red (Rocelin Purple to Deep
Helebore Red), divided into several primary lobes, each lobe divided
again to one-half the depth of the primary lobes, the margins undulate
and proliferous. Monostromatic except basally, without microscopic
veins. Cystacarps 300-500 u wide, carpospores terminal, 45-60 by 30 u,
elliptical to obovate, up to twice longer than broad. Tetrasporangia and
spermatangia not seen.

1969 | ABBOTT: RED ALGAE 45

Holotype: Oregon, Coos Co., Lighthouse Beach, Cape Arago, Ethel /.
Sanborn, July 11, 1926, UC552380. Regretfully known only from the
holotype, this species can be placed without question in this genus, as
each procarp contains only one carpogonial branch and one group of
sterile cells, the carpospores are terminal, and the thallus is without veins.

It differs from the other Nitophyllum species on this coast by being
larger, more lobed, and more strongly proliferous. In fact, this species
resembles some of the proliferous forms of Hymenena setchellii more
than it does any species of Nitophyllum, but lacks the microscopic veins
present in Hymenena, and is a more delicate thallus.

Nitophyllum dotyi is named in honor of Maxwell S. Doty of the Uni-
versity of Hawaii in recognition of his major contribution to the knowl-
edge of the Oregon marine algal flora, and the marine flora of the Pacific
Coast.

Cryptopleura rosacea Abbott, sp. nov. Figs. 3, 4. Thalli 5-10 cm alt.
lobos taeniaformes flabellato et ramosos, qui segmenta taeniaformia 2—3
cm long. ferunt, habentes; margines segmentorum crebre crenati, omni
crena c. 0.5 cm long. cacumina obtusa ad spathulata habente. Partes
inferiores venas macroscopicas non perspicuas, partes superiores venas
microscopicas praebentes. Partes thalli inferiores polystromaticae, partes
superiores et margines monostromaticae. Cystocarpi per loborum super-
ficiem sparsi, carposporas terminales habentes.

Thalli 5—10 cm tall, in a crisp clump, with branched flabellate, ribbon-
like lobes bearing ribbon-like segments 2—3 cm long, with densely scal-
loped margins, each scallop about 0.5 cm long, segments with blunt to
spathulate tips. Thallus a deep rose-red (Indian Lake to Dahlia Crim-
son). Lower portions with indistinct macroscopic veins, upper portions
with microscopic veins. Lower portions of thallus polystromatic, upper
portions and margins monostromatic. Cystocarps few in the main seg-
ments, 1.0-1.5 mm wide, somewhat flat; gonimoblasts with terminal
carpospores.

Holotype: California, Monterey Co., cast ashore at the south end of
Carmel Beach, 4 bbott 4190, May 5, 1965, GMS.

Other specimens: From the type locality, 4191, May 19, 1965, UC;
4192, June 19, 1965; 4193, April 25, 1965; 4194, May 7, 1966.

In general form, this newly described species of Crvptopleura resem-
bles the illustration of C. dichotoma Gardner (1927) but Gardner’s de-
scription of size of thallus, width of blades, branching of segments, char-
acter of the margins and color of the thallus is different from these in
C. rosacea, the latter species being taller, having wider blades, branches
flabellate, with ruffly margins, and of a brighter color. Since C. dichotoma
is known only from the type specimen, no further comparisons can be
made.

Cryptopleura rosacea is smaller and a more splender species than C.
lobulifera or C. brevis, as well as differing in color, width of segments

46 MADRONO [Vol. 20

0 A
WWao8a5
SS

Fics. 5-14. 5-12, Ozophora species; 5, habit of Ozophora clevelandi (the type
specimen of Phyllophora clevelandiz), showing prominent stipes and linear blades
characteristic of this species, x 14; 6, tetrasporangial nemathecia of O. clevelandii
showing chains of tetrasporangia arranged in a nemathecium on the surface of the
thallus; 7, monosporangia of O. clevelandii, arranged in a nemathecium on the sur-
face of the thallus; 8, spermatangial sorus of O. latifolia, occurring on special leaf-
lets; 9, spermatangial leaflets which occur in clusters near the midline of the blades
of the thallus of O. clevelandii (similar in other species of Ozophora), X* 1; 10,
cystocarpic papillae of O. clevelandii, * 1; 11, habit of the type specimen of O.
latifolia, showing broad flabellae dichotomously branched, and an inconspicuous
stipe, both characteristic of this species, x ¥%; 12, habit of cystocarpic thallus of
the type specimen of O. norrisii, showing repeated branching, the last orders of
branches with slender, delicate stipes which ar characteristic of this species. Basal
portion missing, X %; 13-14, Chondrus ocellatus; 13, erect, tufted, little-branched,
furrowed thalli known as C. ocellatus {. parvus which resemble Rhodoglossum af-
fine, X 7%; 4, broad, short, blade-like form close to C. ocellatus {. ocellatus and re-
sembling depauperate specimens of Jridaea flaccida, X 7%.

1969 | ABBOTT: RED ALGAE 47

and density of ruffling of the margins. It appears to be a subtidal species,
as it has not been collected intertidally.

OzopHora J. Agardh (1892) with O. californica as the only species,
was described from the Golden Gate (San Francisco) California, and
was transferred by Kylin (1931) to Phyllophora. Dawson (1961) re-
stricted the northern specimens described by Smith (1944, as PAyllo-
phora clevelandiu Farlow) to P. californica, reserving for the specimens
from Santa Barbara south the name of P. clevelandi. Examination of
the type specimen of O. californica in the Agardh herbarium shows the
cystocarpic material to be identical with the more recently collected ma-
terial in the northern California area. Richard E. Norris of the Univer-
sity of Washington some years ago called to my attention the peculiar
spermatangial leaflike proliferations of this entity. Detailed studies of
these and other reproductive structures show that these specimens can-
not be allied with Phyllophora where the spermatangia occur in cavities
on the surface of the thallus, and where the cystocarps are borne on leaf-
like proliferations. It therefore seems advisable to present a more ade-
quate description of Ozophora.

Thallus erect, with one to several fronds arising from a disc-like hold-
fast, with or without several cylindrical stipes, upper portions producing
irregularly linear to wedge-shaped blades with stipes, or branching dich-
otomously in blade-like segments; if not stipitate, then main axis and
subsequent branches blade-like, sometimes with proliferous stipitate
bladelets and ultimately forming broad, blunt tips. New growth and re-
generation common in various parts of the thallus, at first appearing as
small ear-like lateral proliferations. Medulla parenchymatous, with a
narrow 2-3 layered cortex. Spermatangia in thin cordate leaflets clustered
on the surfaces of the thallus, toward the mid-line, or marginal, the
spermatangia occurring in a colorless superficial band on the surface of
the leaflets. Cystocarps in simple cylindrical or fusiform proliferations
(papillae) the cystocarps bulging out the median portion on the prolif-
erations. Carpospores small, in clusters separated by sterile filaments.
Asexual thalli with monospores (undivided tetraspores?) in superficial
blisters or in small marginal warts. Tetrasporangia in surface nemathecia,
the sporangia in chains, cruciately divided.

Ozophora is clearly related to Phyllophora on the basis of the vegeta-
tive structure, having a parenchymatous medulla and narrow cortex, and
in reproduction, having tetrasporangia in superficial nemathecia, the
tetrasporangia (fig. 6) borne in chains. These characters are also shared
with Petroglossum Hollenberg (1943). Petroglossum differs from PAyl-
lophora in having spermatangial sori continuous with the surface of the
thallus and although spermatangial sori are continuous (fig. 8) in
Ozophora, they are in special (fig. 9) leaflets, whereas in Phyllophora,
in contrast, spermatangia are in pit-like cavities. Petroglossum and
Phyllophora bear cystocarps in leaflets; Ozophora has cystocarps (fig. 9)

48 MADRONO [Vol. 20

in papillae. If the asexual sori of the three species of Ozophora are
sometimes monosporangial (assuming that these are not undivided tetra-
sporangia), these are suggestive of the modified life cycle shown by Ahn-
feltia plicata in the Phyllophoraceae.

Ozophora clevelandii (Farlow) Abbott, comb. nov. Figs. 5, 6, 7, 9,
10. Phyllophora clevelandu Farlow, Proc. Amer. Acad. Arts 2: 368. 1875.
Ozophora californica Agardh, Analecta algologica, 82. 1892. Phyllophora
california (Agardh) Kylin, Acta Univ. Lund. 27(11): 34, pl. 20, fig. 50.
1931. Phyllophora submaritimus Dawson, Allan Hancock Found. Publ.
Occas. Pap. 8: 6, figs. 17, 18. 1949.

Thalli 10-28 cm high, frequently overgrown by encrusting bryozoans,
hydroids, and barnacles, several clindrical wiry stipes (fig. 5) up to half
the height of the thallus arising from a disc-shaped, woody holdfast, the
linear or spathulate blades occasionally furcate, on short stipes if on the
second or third order of branches, blades 0.5 to 2.0 cm wide, up to 15 cm
long, occasionally proliferous (regenerative) at the tips, which are other-
wise blunt. Cystocarps on papillae (fig. 10) borne on the central portion
of both surfaces of the blades, sometimes spreading to the margins but
not marginal; spermatangial leaflets in clusters (fig. 9) at the center of
the blades; monosporangia (fig. 7) (undivided tetrasporangia?) in blis-
ter-like nemathecia on the surface of the blades; tetrasporangia (fig. 6)
in superficial nemathecia, in chains, cruciately divided.

Holotype of Ozophora californica: California, San Francisco, from the
Golden Gate, no. 25365, LD, a fragmented thallus which is cystocarpic.
Another specimen, no. 25364, LD, from Unalaska, is said to be this spe-
cies but it is too fragmentary to identify.

Holotype of Phyllophora clevelandu; California, San Diego Co., San
Diego, Daniel Cleveland, Dec. 1874, FH.

Other specimens: California, Marin Co., cast ashore at Duxbury Reef,
4216, Dec. 27, 1967; Santa Cruz Co., cast ashore at Davenport, Hair &
Nicholson, July 16, 1965; Monterey Co., Hollenberg 3934, July 16,
1939; cast ashore at Moss Beach, 4177; 4178, MSD; 4179; 4209; cast
ashore 3 miles north of San Simeon, 4180, GMS, UC, WTU; 4181, GMS,
MSD, UCSB; San Luis Obispo Co., cast ashore at Shell Beach, 4183,
June 4, 1966; 4184, Oct. 22, 1967, GMS, MSD.

Additional references to Ozophora clevelandii are Smith (1944) and
Dawson, 1949. Additional references to Phyllophora californica are
Dawson (1958; 1961).

Distribution: from Duxbury Reef, Marin Co., through central Cali-
fornia, Channel Islands, and reported by Decor (1961) from several
Baciae Mexico localities.

Ozophora clevelandii is usually collected in the drift, and there usu-
ally as fragments. Such specimens usually resemble various forms of
Prionitis, particularly P. andersonii, and without experience with these
two taxa would be easy to confuse with the latter. Likewise, O. latifolia

1969 | ABBOTT: RED ALGAE 49

(described below) is also found cast ashore, but except for possible con-
fusion with Rhodymenia pacifica, is more easily defined as a separate
entity from other red algae. In part, these associations are a reflection
of the main differences between these two species of Ozophora, namely,
that the conspicuous cylindrical stipes (fig. 5) with broadly linear blades
of O. clevelandii remind one of Prionitis and the broad foliar segments
of O. latifolia (with basal portions lacking) remind one of Rhodymenia
pacifica. The intact thalli of O. latifolia (fig. 11) obtained by dredging
or diving show only very short stipes less than 2 cm high, whereas stipes
may be one-half the total height of the thallus in O. clevelandi. Because
more collections are now available than when Dawson (1949, 1961) drew
up his descriptions of the southern California and Pacific Mexico PAyl-
lophora species, it is clear that on vegetative grounds Phyllophora sub-
maritimus Dawson is the same as Ozophora clevelandu although no fer-
tile material of the southern taxon is known.

The holotype of Phyllophora clevelandi (fig. 5) bears that designa-
tion in the hand and initials of F. S. Collins. It is represented by two
bleached, sterile specimens, obviously parts of the same thallus. On the
same sheet is another specimen, different from these two and bearing
tetrasporangia (fig. 6) in chains. Two packets are also on this sheet, one
from Santa Cruz collected by C. L. Anderson is also tetrasporangial; the
second, from San Francisco, collected by N. L. Gardner is cystocarpic.
The tetrasporangial nemathecia are oval and on the surface of the thal-
lus, and resemble the nemathecia which bear monosporangia of the more
recently collected thalli. The cystocarps of the Gardner specimen are in
fusiform proliferations or papillae similar to those of the type specimen
of Ozophora californica. Although sterile, the type specimen of Phyl-
lophora clevelandit is of the shape, size, and structure of the specimen
illustrated by Smith (1944), which is accepted here as identical.

Phyllophora clevelandu is described by Dawson (1961) as having
tetrasporangia on small leaflets borne on the surface of the blades; since
these are borne in a different place than the reported monospores and
tetrasporangia borne in nemathecia on the surface of the thallus in
Ozophora, it is concluded that some of his specimens are different from
O. clevelandi, and thus probably not Ozophora.

Ozophora latifolia Abbott, sp. nov. Figs. 8, 11. Thallus usque ad 30
cm alt., cum aut sine stipite brevi cylindrico; axis principalis complana-
tus, laminiformis, 2.5—3.0 cm lat. irregulariter ad regulariter dichotome
flabellate ramosus, aut segmenta prolifera stipitata 7-10 cm long., 1.0-
2.0 cm lat., e margine axis principalis producta, habens. Sectiones trans-
versae complanatae, medullam achromaticam parenchymatiformem et
filamenta corticalia photosynthetica, e 3—4 stratis constantia, habentes.
Spermatangia in proliferationibus cordatis laminiformibus, in fasciculis in
superficie segmentorum ordinatis, spermatangiis superficie proliferation-
um complanatis. Cystocarpi in proliferationibus e superficiei laminarium

50 MADRONO LVol. 20

cylindricis ad fusiformes ad teretes, simplicibus aut semel-ramosis facti,
massa Carposporarum media, inflationem parvam efficiente. Thalli asex-
uales monosporici, monosporae (tetrasporae no divisae?) in excrescentiis
marginlibus, aut in nematheciis pustuliformibus in segmentis factae.

Thallus up to 30 cm tall, brick red when fresh, drying to a rusty-red,
with or without a short, cylindrical stipe less than 2 cm high, main axis
flattened and blade-like, 2.5 to 3.0 cm wide, irregularly to regularly
dichotomously flabellately branched (fig. 11), or with stipitate prolif-
erous segments 7-10 cm long and 1.0 to 2.0 cm wide produced from the
margin of the main axis. Cross sections flattened, with a colorless
parenchyma-like medulla and photosynthetic cortical filaments of 3-4
layers. Spermatangia in pink cordate leafllike proliferations 1-2 mm high
and 1-3 mm broad, arranged in clusters on the surface of the segments,
the spermatangia forming a continuous, colorless band (fig. 8), on the
surface of the leaflets. Cystocarps borne in simple or once-branched
cylindrical to fusiform and terete proliferations (papillae), 1 to rarely
2 mm high, from the surface of the blades, and occasionally fringing the
margins; Carpospore mass median in section, and making a small swell-
ing in the proliferation, internally filling the entire center of the medulla.
Carpospores small and ovate, about 10 by 7 u arranged in small clusters
separated by sterile filaments. The asexual thalli monosporangial, mono-
sporangia (undivided tetrasporangia? ) produced in marginal outgrowths,

or in blister-like nemathecia on the segments (branches of the second’

order).

Holotype: California, Monterey Co., dredged at 50-60 ft depth, north
of Coastguard breakwater off Monterey, Abbott 4172, July 31, 1964,
GMS, spermatangial and cystocarpic plants on 1 sheet.

Other specimens: Oregon, Lincoln Co., from Seal Rocks, Doty 2664.
California, San Mateo Co., at 10 ft depth off Pigeon Pt, 4146; Monterey
Co., 29 specimens variously distributed to: GMS, MSD, UC, UCSB, US,
WTU; San Luis Obispo Co., Shell Beach, 3 specimens distributed to:
GMS, MSD, WTU; Santa Barbara Co., Santa Barbara, Peattie 28,
SBM.

Thalli of O. latifolia are taller, the segments longer and broader and
more branched than those of O. clevelandiu. The chief differences are in
the possession of longer and more prominent stipes and linear blades of
the latter species.

Ozophora latifolia is named for its broad axis, blades and thallus
segments.

Ozophora norrisii Abbott, sp. nov. Fig. 12. Thalli subaestuales, usque
ad saltem 20 cm alt. (thallo altissimo fracto), hapteron discoideum
parvum, stipitem delicatum, et axem principalem foliarem habentes; axis
principalis axe secondarios latitudine quasi aequos, ad tertiam quar-
tamque ordinem conferte ramosos, pinnatim irregulariterque efficiens.

1969 ] ABBOTT: RED ALGAE Sit

Laminae proliferae simplices ad spathulatas ad divaricate furcatas, in
omni ordine ramificationis, 1-3 cm long., ad 1 cm lat., in stipitibus deli-
catis tenuibusque sitae. Cystocarpi in papillis secundum margines
orientibus siti; foliola spermatangialia marginalia; monosporangia in
nematheciis pustuliformibus in superficie laminarum orientibus sita.

Thalli up to at least 20 cm tall, from a small discoid holdfast and a
delicate stipe, with a foliar main axis which gives rise pinnately, flabel-
lately, and irregularly to secondary axes of nearly the same width, and
branching closely to the third and fourth order. Simple to spathulate to
divaricately forked proliferous bladelets on all orders of branching, 1-3
cm long, up to 1 cm wide, on delicate slender stipes. Cystocarps on
papillae borne along the margins; spermatangial leaflets marginal; mono-
sporangia (undivided tetrasporangia?) in blister-like nemathecia on
blade surfaces.

Holotype: Washington, San Juan Co., dredged in 40 ft depth Par-
tridge Pt., west of Whidbey I., Norris 4952, July 27, 1964, GMS-holo-
type, WTU.

Other specimens: Washington, San Juan Co., dredged off Salmon
Bank, southwest of San Juan I., Norris 4803, July 16, 1964, WTU,
spermatangia; Norris 5167, Feb. 13, 1965, cystocarpic; at type locality,
Norris, 5214, July 6, 1965, cystocarpic, spermatangial, sporangial, dredged
in 30-35 ft.

Ozophora norristi has a thinner thallus than the other two species in
the genus. It also branches more profusely, the third and higher orders of
branches being distinguished by very delicate stipes. Furthermore, sper-
matangial leaflets are marginal in location. In general form, it is similar
to Petroglossum pacificum Hollenberg (1943, fig. 4) but 3-4 times the
size of this species, and lacks the crustose base of Petroglossum pacifi-
cum. With O. latifolia, it shares a short, inconspicuous stipe.

Ozophora norrisi is named in honor of Richard E. Norris of the Uni-
versity of Washington who first called attention to the peculiar repro-
ductive structures of this genus.

CHONDRUS OCELLATUS Holmes f. PARVUS Mikami, Sci. Pap. Inst.
Algol. Res. Fac. Sci. Hokkaido Imp. Univ. 5: 233, pl. 3, fig. 1. 1965. Fig.
13,14.

Thalli saxicolous, olive green to brown, tufted in two growth forms,
one (fig. 13) 0.5 to 1 cm high from a short, stout holdfast and stipe,
upper portion of flabellae dichotomously branched once or twice, fur-
rowed with blunt, obtuse tips, but otherwise smooth, with edges raised at
the margins; the other (fig. 14) type foliar, with almost no stipe to stipes
of 1 cm high, the blades expanding and 2.5 to 3 cm broad, with broad
rounded tips, and up to 5 cm high.

Monoecious. Spermatangia produced laterally near the tips of the
cortical filaments, less than 2 u wide. Cystocarps up to 1 mm wide, scat-
tered over the median portions of the blades, internally with no ‘“Faser-

a2 MADRONO [Vol. 20

hulle” (special medullary filaments which surround the cystocarp).
Tetrasporangia arising from accessory branches of the medullary fila-
ments, 28 by 35 u associated in flat to ovate internal sori.

Distribution: Oregon, Coos Co., at 3.5—4 ft tide level, below Endo-
cladia zone, Sunset Bay, near Cape Arago, Norris 5359, July 20, 1967,
GMS, MSD, WTU. The Japanese locality for this form is Shimonoseki,
Yamaguchi Prefecture, south central Japan. Forma ocellaltus is found
throughout Honshu and its type locality is Shimoda, Shizuoka Pre-
fecture.

Chondrus, as understood by modern workers, is not known on the Pa-
cific Coast of North America, previous reports having been shown to be
species of Rhodoglossum and other genera. Therefore, to find a repre-
sentative of Chondrus, and to be able to ally it with a known species is
rather a surprise. Although these specimens, on casual inspection, re-
semble Rhodoglossum affine and depauperate /ridaea specimens, close
study of the reproductive structures shows conclusively that this is a
species of Chondrus. The two strongest characteristics shown are: no
“Faserhulle” which is present in the 3 other genera of Gigartinaceae on
this coast, namely, Rhodoglossum, Gigartina and Iridaea; and tetra-
sporangia arising from accessory branches of the medullary filaments,
which only /ridaea of the other genera shares. The specimens show a
remarkable resemblance to those illustrated in Plate 3, fig. 1 of Mikami
(1965), and some of them grade into what Mikami (1965) considers to
be f. ocellatus, being somewhat taller and more robust than the average
for f. parvus.

ACKNOWLEDGEMENTS

I very much appreciate the loan of specimens from Richard E. Norris,
University of Washington; Maxwell S. Doty, University of Hawaii; the
Agardh herbarium at Lund University, the Farlow Herbarium, and the
University of California, Berkeley. Grateful acknowledgment is also
made for financial support under Contract N-0014-67-A-0112 with the
Office of Naval Research. It is a pleasure to acknowledge my continued
indebtedness to George J. Hollenberg for the sharing of his collections,
experience and knowledge with me.

Hopkins Marine Station, Stanford University, Pacific Grove, California

LITERATURE CITED

AGARDH, J. G. 1892. Analecta algologica. Lund.
Dawson, E. Y. 1949. Contributions toward a marine flora of the southern California
Channel Islands. Allan Hancock Found. Pub. Occas. Pap. 8:i-ili, 1-56.
. 1958. Notes on Pacific coast marine algae VII. Bull. S. Calif. Acad. Sci.
57:6-80.
. 1961. Marine red algae of Pacific Mexico Pt. 4. Gigartinales. Pacific Nat-
uralist 2:191-341.

1969 | LANG: POLYPODIUM 53

. 1962. Marine red algae of Pacific Mexico. Pt. 7. Ceramiales: Ceramiaceae,
Delesseriaceae. Allan Hancock Pacific Exped. 26:1-—106.
Doty, M. S. 1947. The marine algae of Oregon Part II. Rhodophyta. Farlowia 3:
159-215.
GarpDNER, N. L. 1927. New Rhodophyceae from the Pacific Coast of North America.
II. Univ. Calif. Publ. Bot. 13:235-273.
Ho.LienBERG, G. J. 1943. New marine algae from southern California. II. Amer. J.
Bot. 30:571-579.
. and J. A. Appotr. 1965. New species and new combinations of marine
algae from the region of Monterey, California. Canad. J. Bot. 43:1177-1188.
., and . 1966. Supplement to Smith’s Marine Algae of the Mon-
terey Peninsula. Stanford Univ. Press.
.» and . 1968. New species of Marine Algae from California.
Canad. J. Bot. 46: 1235-1251.
Kyun, H. 1925. The marine red algae in the vicinity of the biological station at
Friday Harbor, Washington. Acta Univ. Lund. 21(9) :1-87.
—. 1928. Entwicklungsgeschichtliche Florideenstudien. Acta Univ. Lund.
2A (4): 1-27.
. 1931. Die Florideenordnung Rhodymeniales. Acta Univ. Lund. 27(11) :1-

48.
. 1941. Californische Rhodophyceen. Acta Univ. Lund. 37( ?) :1-51.
. 1956. Die Gattungen der Rhodophyseen. Gleerups, Lund.

Mikami, H. 1965. A systematic study of the Phyllophoraceae and Gigartinaceae
from Japan and its vicinity. Sci. Pap. Inst. Algol. Res. Fac. Sci. Hokkaido Imp.
Univ. 5:181-285.

Norris, R. E., and J. West. 1966. Notes on marine algae of Washington and south-
ern British Columbia. Madrono 18:176-178.

Ripcway, R. 1915. Color standards and Color Nomenclature. Washington, D.C.

SmitH, G. M. 1944. Marine algae of the Monterey Peninsula, California. Stanford
Univ. Press.

A NEW NAME FOR A SPECIES OF POLYPODIUM FROM
NORTHWESTERN NORTH AMERICA

FRANK A. LANG

A biosystematic study of the Polypodium vulgare complex in the Pa-
cific Northwest (Lang, 1965) has shown that there are three cytotypes
present in the area from Alaska south along the Pacific Coast to central
California and east to the Rocky Mountains that are apparently involved
evolutionally with each other.

One, represented by P. glycyrrhiza D. C. Eaton, is uniformly diploid
(n = 37) throughout its range and is morphologically, ecologically, and
geographically distinct from another species in the area, P. hesperium
Maxon.

Cytological investigations on P. hesperium have shown that this species
is composed of two cytotypes, one diploid (n = 37) and one tetraploid
(n = 74) (Evans, 1963; Knobloch, 1962; Lang, 1965; Lloyd, 1963,
Manton, 1950).

54 MADRONO [ Vol. 20

The distinctive coastal species P. scouleri Hook and Grev. is also pres-
ent in the study area but is not considered here as it does not appear to
have played a role in the present problem.

The two cytotypes included in P. hesperium are quite distinct and
separable on a number of points (Table 1). It is felt that these differ-
ences are such as to warrant their recognition as distinct species.

TABLE I. CHARACTERS DISTINGUISHING THE Two CYTOTYPES OF P. HESPERIUM

Tetraploid Diploid
Chromosome number n— 74 n = 37
Frond shape oblong oblong
Segment shape acute to obtuse obtuse
Sorus shape oval circular
Sorus location median marginal (near margin

than midrib)

Paraphyses very rare common
Rhizome pruinose — +
Rhizome taste licorice acrid
Scale strip absent te
New fronds summer spring
Geographical interior (mostly east of Western mountains
distribution Cascade Mountains)

Since the two cytotypes included in P. hesperium are recognized as
two distinct species, it thus becomes necessary to establish to which cyto-
type the epithet “es perium properly belongs, the diploid or the tetraploid.
Polypodium hesperium Maxon is the oldest available name which must
be used for one or the other of the two cytotypes.

The type specimen of P. hesperium was compared morphologically
with a range of both diploid and tetraploid specimens from throughout
the Northwest, and chromosome determinations were made on topotype
material. This comparison makes it clear that the tetraploid cytotype
agrees very closely with the type specimen of P. hespertum Maxon.
Among other features in common, both have oval sori located midway
between the costa and segment margin, and a sweet licorice-like rhizome,
as mentioned by Maxon (1900) in his original description. The diploid
cytotype, on the other hand, has circular sori near the segment margin
and an acrid tasting rhizome. The geographical distributions give fur-
ther evidence that the holotype of P. kesperium is tetraploid since, as far
as is known, the diploid cytotype does not occur in the area of its type
locality. A collecting trip to the type locality of P. hesperium in Coyote
Canyon, Lake Chelan, Washington, yielded several isolated colonies of
Polypodium. They were essentially similar in morphology and all plants
on which chromosome counts were made proved to be tetraploid (n =
74). On the basis of the evidence there is little doubt that the tetraploid
should bear the name P. hes perium Maxon.

The restriction of P. hesperium to the tetraploid cytotype apparently
leaves the diploid cytotype without a name. There are, however, several

1969 | LANG: POLYPODIUM 55

possible names in the literature which should be considered.

One epithet that might apply is P. amorphum Suksdorf, the type speci-
men of which was collected at the base of a shady cliff in Dog Creek
Canyon, Skamania Co., Washington, in the region of the Columbia River
Gorge. On the basis of a photograph in Frye (1934), examination of the
type specimen and Suksdorf’s description, this species has some features
in common with the diploid cytotype, viz., a thin slender pruinose
rhizome with typical scales and circular marginal sori with paraphyses.
In morphology of the frond, P. amorphum is very variable with mostly
semicircular frond segments and bifurcate frond tips. The diploid cyto-
type does not agree with it in these features.

In the fall of 1967 and again in the spring of 1968, I made two trips
to Dog Creek to collect topotype material of P. amorphum. Dog Creek
flows through a narrow percipitous canyon in a series of waterfalls, some
up to 30 ft. high. The first trip involved a search of the lower reaches of
the canyon from its mouth at the Columbia River to a point one-half
mile upstream. Here further upstream search became impossible because
of a high waterfall with no safe way around it. The only Polypodium
found on this part of the stream was P. glycvrrhiza. Several additional
attempts by others (Slater, 1964; pers. comm.) at the lower end of the
canyon to re-collect P. amorphum have also failed.

The upper part of the canyon was reached on the second trip after a
steep four and one-half mile hike up Dog Mountain and down a tribu-
tary stream to Dog Creek. Here it was discovered that Dog Creek and
the tributary meet at the botom of an inaccessible box canyon after a
vertical fall of about 75 feet. Specimens of the diploid cytotype were seen
growing, out of reach, from the underside of the overhanging cliffs. None
of these plants appeared to be similar to P. amorphum. A search upstream
from the head of the box canyon yielded only P. glycvrrhiza.

Since Suksdorf was about 70 years old when P. amorphum was first
collected in 1925 and was under a doctor’s order not to make extended
collecting trips (Weber, 1944), it seems likely that he collected his type
specimen from the lower end of the canyon. Weber (pers. comm.) con-
curs with this evaluation and states that Suksdorf definitely did not have
an assistant to collect for him in the field. In making up specimens for
distribution, Suksdorf returned to the type locality three times; in doing
so he probably collected most of the existing plants, any survivors having
subsequently died out. Polypodium amorphum seems to be an example
of the “sports” that occasionally occur in various fern species with its
unusual frond segments and bifurcate frond tips. In all probability P.
amorphum is no longer an extant taxon due mainly to the zeal of its
author.

Article 71 of the 1966 edition of International Code of Botanical
Nomenclature states that a name must be rejected if it is based on a
monstrosity. If one accepts that a monstrosity, in the botanical sense, is
a plant that deviates greatly from the natural form or character, is ab-

56 MADRONO [Vol. 20

normal, or is malformed, then P. amorphum must be rejected, since P.
amorphum does deviate greatly from natural form (compare the phto-
graph of the type of P. amorphum in Frye (1934) Fig. LVIII, with the
drawing of the diploid cytotype of P. hesperium, Fig. LVII, 3 and 4, in
Frye). If one accepts that it is merely a sport of the diploid cytotype of
P. hesperium s.\., it is in any event abnormal in morphology when com-
pared to most members of the P. vulgare complex.

One of the problems with Article 71, as pointed out by Davis and Hey-
wood (1965), is that it fails to define a monstrosity. The definition given
above is based on the 1966 edition of Webster’s New Collegiate Dic-
tionary. One of the examples of the application of Article 71 given in the
rules refers to the orchid genus Uropedium, a peloric form with the third
sepal (labellum) resembling the other two. The generic name, Urope-
dium, according to the rules, is based on a monstrosity and “‘must there-
fore be rejected.”’ The monstrosity is now referred to as Phragmipedium
caudatum (Lindl.) Rolfe.

If P. amorphum was a plant that represented a morphological extreme
along a more or less continuous line of variation in the diploid cytotype
of P. hesperium s.l., then it would be difficult to reject the name under
Article 71. Article 7, note 1, states that the nomenclatural type is not
necessarily the most representative element of a taxon; it is merely that
element with which the name is permanently associated. An exception to
this is when a name is based on a monstrosity (Benson 1962).

Since P. amorphum shows a very clear morphological discontinuity
from the rest of the diploids in P. hesperium s.1., apparently no worse
than the example of Uropedium, Article 71 makes the rejection of the
epithet amorphum mandatory if the Code is to be followed. Of all the
specimens examined in this study, none have approached the morphology
of P. amorphum. In all probability the form arose only once and is now
extinct.

Shivas (1961) has rejected the epithet cambricum for the European
diploid P. australe Fee on the basis that P. cambricum L. is based on a
monstrosity.

I accept a similar view that P. amorphum is based on a monstrosity
and must be rejected under Article 71.

Several varietal names should be considered on the basis of Recom-
mendation 60a of the Code. P. vulgare var. columbianum Gilbert can-
not be used for the diploid cytotype since it is apparently a synonym
of the tetraploid P. hesperium. All of the plants examined from the gen-
eral area of the type locality, the Arrow Lakes in British Columbia, ap-
pear to be tetraploid; the diploid has not been found in the interior.
Photographs of the type specimen indicate that it was probably tetra-
ploid, also they agree closely with the type specimen of P. hesperium and
with known tetraploid plants. In any case, the new combination P. col-
umbianum for the diploid would be illegitimate since it would be a later
homonym of P. columbianum Baker.

1969] LANG: POLYPODIUM 57

Clute (1910) described Polypodium vulgare var. perpusillum from
Mount Lemon, Arizona and gave the following brief description: ‘Fronds
one to four inches long, one-half to three-quarters of an inch wide de-
minishing below, pinnules oblong, obtuse, about eight pairs; sori medium
size, numerous, near margin than midrib.” This description more or less
fits the diploid cytotype, especially the sori being near the margin than
the midrib. Unfortunately, Clute’s type specimen has not been located
nor has topotype material been available for examination. The descrip-
tion is not precise enough to state with certainty that it is the same as
the diploid cytotype of P. hesperium s.1.

In view of this the writer does not feel justified in taking up Clute’s
varietal epithet of perpusillum for the diploid cytotype, and so the new
name Polypodium montense is proposed. Quantitative measurements are
presented in following manner to display best their variability. Using
stipe length as an example the shortest stipe measured was (8 mm long)
(80% of all stipes measured were from 28 to 100 mm long, the average
stipe length being 58 mm long) (the longest stipe was 142 mm long).

Polypodium montense F. A. Lang, nom. nov. Polypodium amorphum
Suksdorf, Werdenia 1: 16, 1927. Holotype: Dog Creek Canyon, near
Cooks, Skamania Co., Washington, Suksdorf 11667, WS.

Rhizoma repens, amarum, 3-5 mm per medium, saepe pruinosum, pa-
leaseum; squamae rhizomatum atrobrunneae vel castaneae, saepe axe cen-
trali cellularum fuscatarum, anguste ovatae vel ovatae, saepe constrictae
supra fundo, usque 5 mm longae, plerumque apicibus capillaribus,
crasse dentatis, cellulis grandibus circa 25 numero trans squamam paulo
supra fundo; frondes circa 130 mm longae, maxima longitudine circa 300
mm; stipites graciles, (8) (28-58-100) (142) mm longi; laminae cor-
iaceae vel membranaceae, oblongae, (18) (46-81-122) (190) mm
longae, (11) (17-24-30) (45) mm latae; laciniae oblongae vel abo-
vatae, apicibus obtusis vel raro acutis, marginibus integris vel cernulatis,
(5) (9-13-17) (25) mm longae, (3) (4-6-7) (12) mm latae, latitudine
longitudini collata ut una pars duabus partibus confertur (1.2) (1.8-2.3-
3.0) (3.6); venis liberis, semal atque iterum; hydathodi parvae, rotun-
dae, paucis cellulis; sorus circularis, propior margini quam costae;
paraphyses multae; chromosomatum numerus n = 37, 2n = 74. Holo-
type: Cheakamus River, British Columbia, Lang 211, UBC.

Rhizome creeping, acrid, 3-5 mm in diameter, often pruinose, pale-
aceous; rhizome scales dark brown to castaeus, often with a central
strip of darkly colored cells, narrowly ovate to obovate, often constricted
above point of attachment, to 5 mm long, usually with a capillary tip,
margin coarsely toothed, cells large, ca. 25 in number across scale just
above point of attachment; frond averaging 130 mm long; max. ca. 300
mm long; stipe slender, (8) (28—58—100) (142) mm long; blades cori-
aceous to membraneous, oblong (18) (46-81—122) (190) mm long, (11)
(17-24-30) (45) mm wide; segments oblong to obovate, tips obtuse to

[ Vol. 20

MADRONO

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Ar Ce
—

nearer the margin than the costa; paraphyses many;

)

sorus circular

chromosome number n = 37, 2n

Sal

1969 | LANG: POLYPODIUM

New fronds produced from late April to June; found growing in rock
crevices in mountains from central Coast Range in British Columbia
south through the Cascade Mountains in Washington to Oregon and the
Sierra Nevada Range in California, in the Olympic Mountains and Wen-
atchee Mountains of Washington and the Northern Coast Range of
Oregon, usually at high elevations but descending to bottoms of river
valleys.

The geographical distribution of P. montense is based on specimens
from the following herbaria, UBC, UC, V, and WTU. Figure 1 shows the
distribution of P. montense, P. hesperium and P. glycyrrhiza in North-
western North America. Polypodium montense is apparently absent from
the Mount Hood region of Oregon south to the Sierra Nevada Moun-
tains in California.

A few herbarium specimens from the mountains of Northeastern Col-
orado, the Laramie Hills of southeast Wyoming and some of the high
mountains of Arizona appear to be P. montense. The majority of speci-
mens from the Rocky Mountain Region, however, are apparently the
tetraploid P. hes perium.

Polypodium montense is closely related to P. virginianum L. of East-
ern North America, sharing many features in common with it, but dif-
fering in its obtuse frond segments and geographical distribution. The
possible role of P. montense in the parentage of the tetraploid cytotype
of P. virginianum has already been discussed (Lloyd and Lang, 1964).

I would like to express my thanks to T. M. C. Taylor for his invalu-
able aid, to William Reid of the University of Washington for his help
with the Latin description, and to Southern Oregon College for partial
financial support.

Department of Botany, University of British Columbia, Vancouver
Present address: Department of Biology, Southern Oregon College, Ashland

LITERATURE CITED

Benson, L. 1962. Plant taxonomy. Ronald Press, New York.

CiuTe, W. N. 1910. Two new polypodies from Arizona. Fern Bull. 18:96—98.

Davis, P. H., and V. H. Heywoop. 1965. Principles of angiosperm taxonomy. Oliver
and Boyd, Edinburgh.

Evans, A. M. 1963. New chromosome observations in the Polypodiaceae and Gram-
mitidaceae. Caryologia. 16:671-677.

Frye, T. C. 1934. Ferns of the Northwest. Metrcpolitan Press, Portland, Oregon.

Kwosiock, I. W. 1962. Tetraploid Polypodium vulgare var. columbianum from
Arizona. Amer. Fern J. 52:65—68.
Lanc, F. A. 1965. A cytotaxonomic study of the Polypodium vulgare complex in
Northwestern North America. Ph.D. thesis, University of British Columbia.
Lioyp, R. M. 1963. New Chromosome numbers in Polypedium. Amer. Fern J. 53:
99-101.

————., and F. A. Lana. 1964. The Polypodium vulgare complex in North Amer-
ica. Brit. Fern Gaz. 9:168-177.

Manton, I. 1950. Problems of cytology and evolution in the pteridophyta. Univer-
sity Press, Cambridge.

60 MADRONO [Vol. 20

Maxon, W. R. 1900. Polypodium hesperium, a new fern from Western North Amer-
ica. Proc. Biol. Soc. Wash. 13:200.

Suivas, M. G. 1961. Polypodium in Europe and America. II. Taxonomy. J. Linn.
Soc., Bot. 58:27-38.

SLATER, J. R. 1964. Fern distribution in Washington State. Occas. Pap. Dept. Biol.,
Univ. Puget Sound 27:243-256.

WeBER, W. A. 1944. The botanical collections of Wilhelm N. Suksdorf, 1850-1932.
Res. Stud. State Coll. Washington. 11:54-121.

THE PYGMY FOREST-PODSOL ECOSYSTEM AND ITS
DUNE ASSOCIATES OF THE MENDOCINO COAST

H. Jenny, R. J. ARKLEY, and A. M. SCHULTZ

INTRODUCTION

Along the Mendocino coast some twenty irregular patches of pygmy
forest, dominated by stunted cane-like cypresses (Cupressus pygmaed)
and dwarfed bishop (Pinus muricata) and Bolander pines (Pinus con-
torta ssp. bolanderi), are surrounded by belts of tall bishop pines and
shore pines (Pinus contorta) and by luxurious regional forests contain-
ing giant redwoods (Sequoia sempervirens) and Douglas firs (Pseudo-
tsuga menziesi). This striking forest differentiation, marked by floristic
endemism, has fascinated botanists ever since Bolander’s early explora-
tions over a century ago. |

Bishop and Bolander pines, but not the shore pine, are closed-cone
pines. Mason (1934) approached the problem of the origin of the coastal
closed-cone pine forests from a broad point of view. Looking for natural
barriers that would conserve pines, he ruled out—correctly we think—
local climates, topographic constellations and especially country rock, as
there are no serpentines, quartzites or other rock extremes. Mason then
searched for a water barrier and assumed, in analogy with the fossil and
living forests on Santa Cruz I. off Santa Barbara, that the coastal strips
used to be Tertiary pine-populated islands that later united with the
mainland and preserved their unique flora against infiltration from the
continental forest. Just how the aggressive invaders from the regional
redwood and Douglas fir forest were kept at bay during hundreds of
thousands of years could not be explained. A good account of Mason’s
ideas is given by Cain (1944) and more recently by Langenheim and
Durham (1963).

The possible role of soils in the floral discontinuities of the greater Fort
Bragg area came into focus with the work of the Mendocino County Soil
and Vegetation Survey during the late forties, and by the subsequent
studies of Gardner and Bradshaw (1954) and Mason’s student Mc-
Millan (1956; 1964).

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST 61

NEED FOR AN ECOSYSTEM CONCEPT

When it was discovered that pygmy forest grows on podsol soil, known
locally as Blacklock soil, having a white, bleached surface horizon and
an iron-cemented hardpan below it, naturalists indulged in an apparent
circulus vitiosus. On field trips the professors of botany would tell their
students that the podsol soil is the cause of the unusaul assortment of
plant species, whereas the visiting professors of pedology (soil science)
would attribute—in the light of classical podsol theory—the soil horizon
features to the acid-producing vegetation. While it is true that a species
individual responds to its soil niche, it is also true that it modifies that
niche, which, in turn, reacts upon the individual. A broader approach is
called for, the joint development of soil, vegetation and animal life with
their mutual interrelated feedbacks (Jenny, 1961). It is embodied in the
concept of the ecosystem.

ECOSYSTEMS RELATED TO LAND ForMs

During Pleistocene times, when the continental glaciers formed and
melted, the world-wide sea level sank and rose. The rising sea cut terraces
into the prevailing graywacke sandstone rock. The retreating sea covered
these platforms with beach sands, gravels and clays. Tectonic forces ele-
vated the terraces. In this light, the higher terraces in the Mendocino
area most likely are the older terraces.

Detailed field work between Navarro River and Fort Bragg led Gard-
ner (1967) to assign terrace levels at altitudes of 100 ft., 175 ft., 300 ft.,
425 ft. and 650 ft., corresponding to first, second, third, fourth and fifth
terraces. These are the major terraces, according to our observations.
The measurements do not refer to the actual terrace surfaces but to the
hidden, buried “‘nickpoints” where terrace floor and sea cliff meet. In-
variably, the pygmy forests and their associated extreme podsols are ex-
tensive on the three upper terraces (fig. 1).

The nearly level terraces are dissected by rivers that flow from the
inland graywacke mountains westward to the sea. Hill and canyon slopes
are continually being rejuvenated by a combination of slow geologic
erosion (back cutting) and sandstone weathering. The slopes, mostly
steep, are covered with impressive regional forests rich in redwoods and
Douglas firs.

Besides these terraces, mountains and canyons a fourth landscape fea-
ture assumes prominence, the sand dunes. Wind is presently blowing
graywacke-type sand from the beach up on to the adjacent higher first
terrace, thereby placing fresh dune sand upon older, weathered and
plant-covered beach deposit. What is happening today apparently hap-
pened in the past, for extensive sand dunes rest on all terraces. On the
higher plateaus they have undergone intensive weathering and produced
Noyo soils, but dune size and shape are largely preserved. Most impor-
tant, the dunes on the lower terraces carry redwood and Douglas fir,
those on the higher mainly bishop pines.

62 MADRONO [Vol. 20

' Wave cut
Platforms

pEEEE] Dunes
EEEEsd Beach deposits
\Gelean ITI Sondsto

[HTT] Sondstone

Sea

Fic. 1. Schematic arrangement of four marine terraces (1, 2, 3, 4) Fort Bragg area,
with a young dune on second and very old dune on fourth terrace. Gr. = grassland,
Rw, Df = redwood-Douglass fir forest, Bi = bishop pine forest, Py = pygmy for-
est. Horizontal distance is 3 miles, vertical distance 500 ft. above sea level.

To sum up, there is, then, a remarkable and fortunate mineralogical
uniformity in the soil parent materials which are the initial states of the
various forest soil ecosystems. They are either weathering graywacke
sandstone or sandstone-derived beach materials and dunes. And though
the land-form features fail to be uniquely reflected in vegetation discon-
tinuities, they nevertheless bring out clearly the convergence of narrow
endemism and dwarfism upon the higher and hence older land surfaces
(vetusta surfaces). Their strongly podsolized soils provide niches for
closed-cone pines and these niches also govern size and shape of the trees,
tall bishop pines on Noyo soil and puny dwarfs on Blacklock soil.

THE Pycmy ForeEst-PopsoLt ECOSYSTEM

Among the various forest soil ecosystems along the coast the pygmy-
podsol type has received most attention.

Vegetation mosiac. Extreme pygmy forest is species-poor and space-
unsaturated, with as much as 25 per cent of the ground area bare or
covered with colonies of lichen. Slender cypresses and gnarled and twisted
bishop-and Bolander pines, many decades old and some passing the cen-
tury mark, are only 1.5-3 m tall. Their trunk thicknesses do not exceed
the diameter of a human wrist or arm. |

Bolander pine was recognized as a distinct taxon, Pinus bolanderi Parl.
by McMillan (1956). Critchfield (1957) designated it as a subspecies of
P. contorta. Its leaves are narrowed, devoid of resin canals and the cones
are heavier and asymmetrical. Unlike P. contorta, they are serotinous
(closed-cone), a feature confirmed by plantings at the Institute of For-

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST 63

est Genetics, Placerville, California (personal communication by W. B.
Critchfield). According to E. G. Linsley (personal communication) the
cones harbor a longhorned beetle (Paratimia conicola) not observed on
P. contorta but known from closed-cone pines in the dry interior Coast
Range foothills (P. attenuata). Bolander pine is an advanced stage of
ecotypic differentiation.It is an “edaphic” ecotype conforming to Tures-
son’s definition.

Likewise dwarfed are the prominent ericaceous shrub companions, like
Ledum glandulosum (Labrador-tea), Rhododendron macrophyllum)
(rose-bay), Gaultheria shallon (salal), the two Arctostaphylos (manza-
nita) species, nummularia and columbiana, and Vaccinium ovatum
(huckleberry). Trees and shrubs exhibit die-back symptoms and fungus-
gall infestations suggestive of specific nutrient deficiencies. Indeed,
chemical analysis of pine needles registers deplorable shortages of potas-
sium, calcium, magnesium, and phosphorus (analyses by A. Ulrich).

The dwarf extremes are interspersed with clusters and thickets of
taller pines and cypresses, in the 6—12m range, still with no signs of red-
wood or Douglas fir, though these giants prosper a short distance away.
Occasionally, a statuesque bishop pine as high as 21m towers amidst the
dwarfs.

Soil profile features. For an explanation of this living mosaic we must
take a look at the underlying Blacklock podsol soil (fig. 2), a Typic
Sideraquod. It is easier said than done. It takes two people with a sharp
auger two to three hours to penetrate the dark-gray, 4-inch thick surface
layer (Al horizon), the 14-inch-thick bleached, white A2 horizon, and
the concrete-like hardpan B-horizon (Bmir) which occupies the depth
interval of about 18-30 inches. Below the pan is rusty, mottled sand or
sandy loam, weakly cemented in places. At a depth of 5-10 ft. unaltered
sandy beach material (C-horizon) is reached. It rests at 10-20 ft. on the
impervious, sea-cut platform of hard sandstone.

The Blacklock surface soil is extremely acid, pH 2.8-3.9, one of the
most acid soils known anywhere. It is low in available nitrogen and phos-
phorous, demonstrated already by McMillan (1956), and in potassium
and micronutrients, as ascertained by elaborate pot tests in the green-
houses at Berkeley. The supply of the nutritionally important exchange-
able calcium (Ca), magnesium (Mg) and potassium (K), expressed as
milliequivalents in 100 g oven-dry soil, is exceedingly low. Above the
hardpan the sum of these bases (Ca + Mg + K) is less than 1 meg/
100g.

In stratified and finer textured beach materials of the old terraces a
dense clay pan with up to 61% clay may occur instead of the iron ce-
mented hardpan. It too acts as an effective impediment to root penetra-
tion. The bases are likewise low and mineral acidity is high. We are
naming this soil Aborigine. A multitude of brown iron streaks and
patches tint and mottle its clays.

64 MADRONO [Vol. 20

Fic. 2. Pygmy forest—podsol ecosystem. Cane-like Cupressus pygmaea crowing on
Blacklock soil showing surface humus horizon on white bleached A2 horizon which
rests on iron hardpan. Water table is at 66 cm depth. Photo R. A. Gardner.

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST 65

Altitude of ferraces

Nov Jan March Moy July Sept
Oct DEG Feb April June Aug

Fic. 3. Fluctuating water tables in Blacklock soil. Profile horizons indicated on
left. During fall the water table is below 10 ft. After winter rains start a perched
water table p above the hardpan is formed. The deep ground water table rises slowly
to maximum height in February. During prolonged rains it may reach the hardpan.
Mean annual precipitation is 38 inches.

Water regimen. If a 10 foot test hole is dug in the fall season, no free
water is encountered. In early November, following the first 5-8 inches
of winter rains, water begins to pile up on the basal rock plane and a
rising groundwater table is set in motion. Long before it reaches the
upper strata, water accumulates above the hardpan layer creating a sec-
ond, perched water table (fig. 3) that floods the entire surface soil. In
late spring the surface water table disappears by evapotranspiration and
seepage. The soil down to the hardpan dries out, hardens and imparts
extreme xeric conditions. In depressions and low places wetness persists
throughout the rainless summer, giving rise to small sphagnum bogs. By
October Ist, the descending ground water table has receded below the 10
ft.mark. Judging from about 40 permanent installations the rate of
descent varies substantially among sites. Its correlation with the vegeta-
tion pattern has not yet been undertaken.

Podsolization process. As said, during the rainy season the surface soil
is terribly wet and water stands in puddles and ponds. Its color is coffee
brown from dissolved acid humus substances, their acidity originating
from the carboxyl groups of pine needles and ericaceous leaves. The
chelates of the humus combine with iron and other metals made acces-
sible by weathering and render them mobile. Prior to and during hard-
pan formation the metal chelates (Fe, Mn) percolated and diffused into

66 MADRONO [Vol. 20

the subsoil, thereby bleaching and impoverishing the surface soil and
leaving behind a snow white, thixotropic A2 horizon. This is podsoliza-
tion. For reasons not yet fully understood, even though European investi-
gators succeeded in imitating the process in the laboratory, iron may be
precipitated in the subsoil as colloidal iron hydroxide. Its positive charge
combines, according to a prominent theory, with the negatively charged
soil particles cementing them together to an iron hardpan (Bmir of
Blacklock). According to R. Tuxen (personal comment to Jenny) and
Bloomfield (1965), iron migration is sensitive to the floristic composition
of the plant cover. The process offers a promising challenge to an eco-
system-oriented biochemist.

Enclaves and borders. Within the pygmy forest area the taller pine
and cypress thickets previously mentioned occupy fine textured (more
clay, less sand) beach deposits but their clayey B-horizon is more perm-
eable to roots than the severe Aborigine clay pan. The base status of
lower horizons may be relatively high. The origin of this soil diversity is
still obscure. One of the lone, impressive bishop pines surrounded by
dwarfs had its root system exposed by chiseling away the hard A2 and
indurated Bmir horizons. Surprisingly, an enormous tap root penetrated
the hardpan and extended into the permeable deeper strata. Maybe there
had been a crack or blemish in the pan, or the root was endowed with
an exceptional supply of iron-dissolving chelates.

Seen from a distance, the change from pygmy to regional forest is very
sharp. Vegetation appears discontinuous. Looking at the boundary more
closely, 30-100 ft. wide transition zones (ecotones) disclose modulations
of soils and plants. Where the canyon of Jug Handle Creek cuts into the
fourth terrace (fig. 4) the drainage pattern is altered, the surface soil is
deeper and moist in summer, and the hardpan is partially or entirely ab-
sent. At another site, an old sand dune rises rather abruptly above the
pygmy plain. Its deep, weel drained soils offer a foothold to tall trees and
thereby maintain a sharp vegetational contrast.

Multiple causes. In ecological parlance “‘edaphic causes” shape the ap-
pearance of vegetation. The adjective edaphic pertains to soil and its
parent material. For Blacklock soil, what are the specific causes of en-
demism and dwarfism? Is it wetness in winter, dryness in summer, fluc-
tuating water table, or is it oxidation or reduction, or harmful nitrite
formation associated with the water regimen? Or, is it the impenetrable
hardpan, either as a physical obstacle or as an unfavorable chemical en-
vironment? Or, is it high acidity or its related aluminum toxicity, or
any one or all of the deficient nutrient elements in the spectrum of soil
fertility? There is an enormous multiplicity of “causes,” for thousands
of soil properties are interrelated among each other, and with countless
properties of the root system, and with enzymatic reactions and meta-
bolic pathways inside the plant. It is a truly multivariate statistical prob-
lem, and we are approaching it in this light. We selected operationally
soil variables that are amenable to manipulation and that are largely in-

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST 67

ft | Rw Rw Bi Bi Bo Fy 49 — 3m
>30m = 15-30m 6-l2m

20
20
:
es mum tndurated A 0
/O ae levies. partial hp

——— water table, Jan.

/00m
LOO 400 tt.

Fic. 4. Transition from pygmy (Py) forest on fourth terrace to regional forest in
Jug Handle canyon. Circles indicate profile sites; water table as of Jan. 17; tree
height in meters; Rw = redwood, Bi = bishop pine, Bo = Bolander pine. Vertical
scale magnified.

dependent of each other (non-collinear). Thus, we initiated large-scale
field experimentation on drainage with deep and shallow drains, random-
ized fertilization and breaking up of hardpan. So far, after a year’s work,
growth increments have been small, as one might expect for dwarfs.
Mean elongation of five marked twigs on each of 80 pine trees was more
than twice that of 80 cypresses. Also, elongations were larger on drained
than undrained sites.

At any rate, this formidable array of soil attributes resulting from sys-
tem evolution puts a tremendous strain on higher organisms. Redwood
and Douglas fir do not grow and cypress and pines barely survive.

GENESIS OF ECOSYSTEM AND ITS SIGNIFICANCE FOR
ENDEMISM AND DWARFISM

The staircase of terraces carpeted with beach materials and dune sand
offers beautiful illustrations of ecosystem genesis. The dune sequence
shall be taken up first.

On the lowest terrace, recent dunes, if bare, are still moving inland.
Stabilization is brought about by colonizers, including lupines and P.
contorta and P. muricata. The pines are able to endure strong salty
winds, dry crests and winter-wet depressions. Slowly soil fertility is be-
ing built up.

Further away from the coast, as near Inglenook, the dunes on the sec-
ond terrace (Lv-sites) are thousands of years old, but no C-14 dates are
on hand. Dunes on the third terrace, Nm sites southeast of Cleone, might
have been blown at the end of the last inter-glacial period or sometime

68 MADRONO [Vol. 20

thereafter. In all these young dunes oxidation has converted the orig’n-
ally drab, gray color of the recent dunes into a warm, rich brown. The
minerals have weathered moderately, clay in amounts of 10-20% has
been formed, and the exchangeable bases Ca + Mg + K are present in
full measure, especially in the surface horizons (fig. 5). Acidity is around
pH 5, considered advantageous for forest growth. All sites are covered
with magnificent forest of redwoods, Douglas fir, grand fir (Abies gran-
dis) and some western hemlock (Tsuga heterophylla). The soils abound
in total nitrogen and in mildly acid humus, as exemplified by the carbon
(C) — curve Lv in Fig. 6. These organic soil properties were not pres-
ent at ecosystem time zero, the fresh dune, rather they are feed-back de-
rivatives of the plant mantle on the one hand and the active microbial
soil population — including the crucial nitrogen-fixers — on the other.

On the fourth and fifth terraces very old dunes are clearly recogniz-
able. Sites specifically studied are labelled as Wi at 410 ft. altitude along
Willits Road, and Dr at 560 ft. at the east end of Gibney Lane. The
dunes are strongly weathered to great depth. The soils, known as Noyo,
have podsolic features with a conspicuous, bleached, whitish A2 horizon
underlain by yellow-brown, clay-rich B-horizons. The deeper subsoils
may exhibit red-white reticulate mottling, a sign of profound chemical
alteration. There is no hardpan though isolated iron concretions and
cementations may appear in its place. During winter, water tables may
be observed at depths greater than 10 feet.

The base content (Ca + Mg + K) is low (fig. 5, Wi, Dr), and soil
reaction is sour, pH being around 4. Instead of mere humus acidity, as in
Lv, Noyo’s has a strong component of aluminum-rich clay acidity, said
to be harmful to root growth. To a depth of 4 inches organic matter is
enriched, but below that surface strip it drops to low magnitudes (fig.
6):Dr)..

The dominant vegetation consists of sizable bishop pines, up to a
century old, an isolated redwood tree here and there, and dense 1.5—3m
tall underbrush of ericaceous species, joined by wax mytle (Myrica
californica ) and chinquapin (Castanopsis chrysophylla).

To summarize, the dune sequence expresses the transformation of the
inert, fresh dune into a giant-tree ecosystem with an abundance of mild
humus, a rich supply of bases, and advantageous quantities of soil acid-
ity. This forest belongs to the major climax associations of the Coast
Range (Heusser, 1960). Further soil transformations enhanced mineral
acidity, depleted the stock of bases by leaching, and drained the humus
reservoir to one-half by virtue of altered litter fall and soil microbe as-
sembly. Gradually, the habitat became Noyo soil and the lush regional
forest was displaced by endemic stands of closed-cone pine. From a util-
itarian point of view, such as a lumberman’s, the ecosystem deteriorated.
If climax is defined (Cain, 1944) as a terminal plant community which
is in dynamic equilibrium with the prevailing climate, then bishop pine
rather than redwood forest would be climax on the dune. It explains in

1969 |

2 O

JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST 69

Ca + Mq + K
5 10

me/|l00 g

eae 100

ole

200

Fic. 5. Soil-depth curves of sum of exchangeable bases (me. of Ca + Mg + K in
100 grams of soil) for sand dunes of various ages. Lv is the youngest, Dr the oldest

dune.

part the perpetuation of pine forests along the coast.

The parallel genesis of the terrace soils proper is accompanied by veg-
etation diversification of spectacular extent. New state factors entering
into play are sluggish surface drainage, seasonal high water table, and
texture extremes of beach materials ranging from sands to clays. When
these materials were deposited their exchangeable bases had been in

70 MADRONO [ Vol. 20

ZC al C 3 4
| cm

<i) Fe See

4 Dr
: Lv
20k 00
(
mele

ONG)

6G [50

Fic. 6. Soil organic matter (mainly humus) expressed as percentage of organic
carbon, in young (Lv) and very old (Dr) dunes in relation to soil depth.

chemical equilibrium with sea water which enriched their sodium content.

The first terrace supports grasslands and pine forests. Few redwoods,
Douglas firs or hemlocks are seen. It appears that these trees cannot bear
the local sea-salt and sodium challenge of air and soil. Under bishop pine
in a sandy matrix Gardner (1967) sampled near Cleone a profile having a
weakly bleached A2 horizon resting on a rusty colored sandy hardpan of
weak cementation. It may be considered a precursor to Blacklock soil,
the more so as its base content is relatively high.

East of Fort Bragg, the expansive second terrace displays wide eco-
system diversity. There are patches of tall redwoods on finer textured
soil lacking wetness and A2 horizon but possessing an iron-stained
clayey B. It might be a precursor to Aborigine soil. Nearby are tall,
slender redwoods and hemlocks on waterlogged soil with A2 and iron

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST qi

concretions. Not far from it dense mixed stands of 12-18 m tall cy-
presses and bishop pines with an occasional redwood tree grow on
bleached soil with iron nodule concretions. Last but not least, there is a
tract of Blacklock soil with dwarfy forest devoid of commercial timber
species.

On the third and higher terraces pygmy forest with pines and cypresses
of various degrees of dwarfism is associated with Blacklock and Aborigine
soils, as mentioned. In their sand fractions Gardner (1967) counted the
slowly weathering potassium feldspar crystals (F) and the highly re-
sistnet quartz grains (Q). In Fig. 7 the half circles on the vertical axis
denote F/Q ratios of recent dune and beach materials. The white dots
characterize the C-horizons of Blacklock soils on various terraces. In
spite of the scatter of points, the trend (dashed line) confirms the miner-
alogical uniformity of the parent materials. Their mean is 17.5 K-feld-
spars per 100 quartz grains.

The black dots record F/Q of the Blacklock A2 horizons. The profiles
of the older surfaces display exceedingly low ratios, less than 0.0003 for
the fifth terrace. They signify far-gone weathering and they establish
antiquity of soils, and, therewith, stability of the land forms. Moreover,
the declining curve, in approaching zero, defines a terminal steady state
condition. Barring a catastrophic change in state factors, we cannot vis-
ualize progression except perhaps for the trees getting more dwarfy.

It is tempting to view the evolution of the podsol ecosystems as an
approximation to a monotone time-sequence operating in the cool and
humid oceanic climate of the Fort Bragg area under conditions of toler-
able salt influx. The initial state, the landscape situation at the start,
comprises dune and terrace plus its biotic factor, the latter defined as
the pool of species available to the site.

Today’s biotic factor is made up of the germules offered by the re-
gional forest and the coastal scrub and grasslands with an admixture of
pine diaspores, particularly bishop-, shore- and Bolander pines. For the
old Noyo, Blacklock and Aborigine soils the initial biotic factor might
have included the Pleistocene ancestors of the pines. As the generations
of seeds sprouted and grew the ensuing tree growth and vegetation dif-
ferentiation responded to niche-creating soil development. Specifically,
the emergence of dwarfism, severe ecotypes and local endemism became
a consequence of orderly system evolution. It is not known on what soil
type or types bishop pine evolved its genetic constitution, but that of
Bolander pine presumably developed in conjunction with Blacklock and
Aborigine soil genesis.

Natural vegetation sequences of the order of magnitude here envisioned
are customarily attributed to a climatic shift. While we do recognize cli-
matic changes in the Mendocino area, we do not believe them to be
critical. Even if effective moisture had been doubled or reduced, it would
have merely temporarily accelerated or retarded the long-time podsolic
trend that molded both phenotype and genotype.

72 MADRONO [Vol. 20

100 175° 300 425 650fF

Fic. 7. Ratios of per cent K-feldspar (F) and per cent quartz (Q) in the sand
fractions of parent materials (white dots) and in Blacklock A2 horizons (black dots).
Parent material averages 17.5 feldspar to 100 quartz grains. The lower the ratios the
more weathering has occurred.

THE AGE PROBLEM OF THE PopsoL ECOSYSTEM

Heusser (1960) published a pollen profile of a boggy site in the pygmy
forest southeast of Fort Bragg. Bolander pine was prominent through-
out the span of some 6,000 years. Redwood can be traced to much earlier
periods as buried trees are frequently encountered by well drillers. A
log was found at 16 ft. depth at the base of the Nm dune which sits on
the front edge of the third terrace.

If the origin of the first terrace is correctly interpreted, its cutting was
completed at the onset of the Wisconsin glaciation, some 100,000 years
ago. If the higher terraces are also linked to glacial periods, time spans
up to one million and more years could be involved (middle or early
Pleistocene). These estimates pertain to the rock-cut terrace platforms.

1969 | JENNY, ARKLEY, & SCHULTZ: PYGMY FOREST mR)

Owing to erosion and deposition, the soils on a terrace might be
younger than the platform itself. Also, since the classic podsols of north-
ern Europe are all post-glacial, podsolization is a relatively fast process.
Still, none of the German and Scandinavian profiles and their plant cover
even approaches the extreme hardpan and dwarfism of the Mendocino
Blacklock ecosystem.

The low F/Q quotients of Fig. 7 prove advanced age of soils but they
do not elucidate the age of the profile features, specifically of the hard-
pan. In a clever piece of detective work, Gardner (1967) answered the
query for site Wi where an extensive old dune rests on the fourth ter-
race. It was blown in when the sea level stood at the third terrace.
Gardner dug a vertical shaft into the dune. At 13 ft. depth the deeply
weathered mantle faded rapidly into unaltered dune material, its slip
faces still intact. Their inclinations were identical with those of today’s
fresh dunes, and so was their orientation as to wind direction. At 20 ft.,
at the unweathered base of the dune, a light-gray horizon was underlain
by a rusty-streaked, irregularly cemented hardpan, the two strata iden-
tifying a Blacklock precursor. No wood remains showed up.

Podsolization on the high terrace must have started prior to dune de-
position, and though the process became arrested under the dune, it
continued outside, for Blacklock exists there now. Gardner (1967) de-
vised a speculative mathematical model of the weathering process that
predicted a soil age of about a million years on the highest terrace. The
order of magnitude seems plausible.

To conclude, not only the terraces themselves but their soil profiles too
possess a venerable age For how long today’s Blacklock and Aborigine
profiles have capped the terrace mantle is not known. Because of its age
and extreme features the pygmy forest-podsol ecosystem is unique in the
temperate region, and it comes as close to a terminal steady-state system
with balanced inputs and outputs as can be expected to be found in na-
ture. It deserves further intensive investigation. To do so, suitable sites
must be protected as scientific reserves. It is an urgent task that demands
highest priority (Jenny, 1960).

SUMMARY

1. Along the coast of northern California, the higher, older marine ter-
races carry pygmy forest, and the associated old sand dunes are covered
with bishop pines. The lower, younger terraces and dunes support grass-
lands, pines, and redwood-Douglas fir forest.

2. Pygmy forest consists of small cypresses (Cupressus pygmaea),
dwarfed, closed-cone pines (Pinus muricata and P. contorta ssp. bolan-
deri) and stunted ericaceous associates, growing on extreme podsol soil
having a bleached, white A2-horizon underlain by an indurated iron
hardpan. During millennia vegetation and soil have evolved together
(pygmy forest-podsol ecosystem).

3. Two sequences of podsolization are envisioned, both starting on

74 MADRONO [ Vol. 20

sandy parent materials: a. Dunes. Younger dunes, in contrast to unal-
tered recent dunes, are slightly weathered. They are high in bases and
humus, low in clay acidity and they support luxurious redwood-Douglas
fir forest. Continual weathering and leaching in the cool and humid
oceanic climate impoverishes the soil and augments acidity. Through
time the regional forest is gradually replaced by endemic bishop pines.
b. Terraces. On the level plateaus with high, fluctuating water tables the
podsolic processes are intensified, resulting in hardpans and clay pans
and soil conditions that produce dwarfism, narrow endemism and _ pro-
nounced ecotypic differentiation.

4. Geologic considerations and soil weathering indices provide age esti-
mates of several hundred thousand years. The pygmy forest and its pod-
sol soil comes as close to a terminal steady-state ecosystem as can be ex-
pected to be found in nature. Adequate preservation is urgent.

The authors acknowledge the help of A. E. Salem. Research supported
in part by the National Science Foundation, Grant GB-6057.

Department of Soils and Plant Nutrition, and School of Forestry
and Conservation, University of California, Berkeley

LITERATURE CITED

BLOOMFIELD, C. 1965. Organic matter and soil dynamics. Jn Hallsworth, E. G. and
D. V. Crawford. Experimental pedology. Butterworth’s, London.

Carn, S. A. 1944. Foundations of plant geography. Harper and Broth, New York.

CRITCHFIELD, W. B. 1957. Geographic variation in Pinus contorta. Publ. Maria
Moors Cabot Found. Bot. Res. No. 3.

GarDNER, R. A. 1967. Sequence of podsolic soils along the coast of northern Cali-
California. Ph.D. thesis, Univ. California, Berkeley.

GARDNER, R. A., and K. E. BrapsHaw. 1954. Characteristics and vegetation relation-
ships of some podsolic soils near the coast of northern California. Soil Sci. Soc.
Amer. Proc. 18:320-325.

HeusseEr, C. J. 1960. Late-Pleistocene environments of North Pacific America. Amer.
Geogr. Soc. Publ. 35.

Jenny, H. 1960. Podsols and pygmies: a special need for preservation. Sierra Club
Bull. 8-9, April-May.

. 1961. Derivation of state factor equations of soils and ecosystems. Soil
Sci. Soc. Amer. Proc. 25:385-388.

LANGENHEIM, J. H. and J. W. DuruHaAm. 1963. Quaternary closed-cone pine flora
from travertine near Little Sur, California, Madrono 17:33-51.

Mason, H..L. 1934. Pleistocene flora of the Tomales Formation. Publ. Carnegie Inst.
Wash. 415:81-180.

McMittran, C. 1956. The edaphic restriction of Cupressus and Pinus in the Coast
Ranges of central California. Ecolog. Monogr. 26:177-212.

. 1964. Survival of transplanted Cupressus and Pinus after thirteen years

in Mendocino County, California. Madrono 17:250-253.

NEW RECORDS OF MYXOMYCETES FROM OREGON. I.
DWAYNE H. CurtTIs

A moderate amount of information has appeared in the literature deal-
ing with slime molds of Oregon. At the present time, using only the
species concepts accepted by Martin (1949), 188 species of Myxomy-
cetes have been recorded from the state. The most extensive investiga-
tion, covering a period of more than twenty years, was conducted by
Peck and Gilbert (1931). They collected primarily in northwestern Ore-
gon from the western slope of the Cascades south to the Three Sisters
Mountains, and the upper northern third of the Willamette Valley in-
cluding the Coast Mountains. In 1932, Martin reported the occurrence
of a new species from Oregon. In recent years only a few species have
been added to the list. Two new species have been described by Kowalski
(1966, 1968), and I (Curtis, 1968) reported Barbevyella minutissima
Meylan from southern Oregon.

During the summers of 1966 and 1967, I collected in Crater Lake Na-
tional Park in the south, central part of the state. Most of the specimens
were obtained from moist decaying wood near the melting snow in the
months of June and July. Later in the year, the slime molds were col-
lected from duff, bark, fallen twigs and decaying wood. All my collec-
tions were taken at altitudes from 4,000 to 7,500 feet. In this paver, 8
species of Myxomycetes are listed as new to the state in the sense that
no report of their occurrence in Oregon has been previously published.
This brings the total number of slime molds found in the state to 196
species.

All collections have been deposited in the University of Iowa Herbar-
ium, Iowa City, Iowa, and specimens in the Trichiaceae, Physaraceae
and Didymiaceae have been deposited in the Crater Lake National Park
Herbarium, Crater Lake, Oregon. The numbers used for the collections
are my own and in this report indicate only those specimens given to the
University of Iowa Herbarium.

LICEACEAE
Licea pusilla Schrad. On decayed coniferous wood 0.2 miles north of
Park Headquarters, 6,500 feet, 6, June 15, 1966. A limited number of
sporangia were found in one collection. They are purplish-brown, sessile,
and dehisce by preformed lobes. This exceedingly tiny species was re-
ported by Kowalski (1966a) recently from California. Previously it had
only been found as far west as Iowa, and is considered rare.

RETICULARIACEAE
Lycogala flavofuscum (Ehrenb.) Rost. I obtained only one aethalium
on the side of a dead, barkless stump about 4 feet above the ground on
the east side of Kerr Valley, 6,500 feet, 52, July 28, 1966. The aethalium
is about 15 mm in diameter, ochraceous-grey and the spores are buff in
mass.

76 MADRONO [Vol. 20

DIANEMACEAE
Dianema Andersoni Morgan. One collection on decayed wood, about 2
miles north of Park Headquarters, 6,800 feet, 1067, July 14, 1967. The
sporangia are sessile, 0.6—0.8 mm in diameter. This Myxomycete has
been reported from Washington, British Columbia, and more recently
from California (Kowalski and Curtis, 1968). It is considered rare.

TRICHIACEAE

Trichia affinis De Bary. Five collections on bark and decayed wood,
one near White Horse Creek and Highway 62, 5,800 feet; two near Park
Headquarters, 6,500 feet, 45, 69; one on the east side of Kerr Valley,
6,500 feet; and one in the vicinity of the Vidae Falls Springs, 6,800 feet.
The sporangia are 0.5—1 mm in diameter, crowded, and bright golden-
yellow.

Hemitrichia montana (Morgan) Macbr. My collections, 67, 835, 857,
1064, indicate that this species is ubiquitous throughout the park at ele-
vations from 6,000 to 7,000 feet. It is so common that I am convinced
that perhaps it was reported by Peck and Gilbert (1931) under another
name. Hagelstein (1944) uses it synonymously with Hemitrichia clavata
(Pers.) Rost. This common species may be sessile to short stalked with
a translucent, shining peridium. The color is variable from a bright ochra-
ceous-orange to a dark greenish-olive.

PHYSARACEAE
Physarum auripigmentum G. W. Martin. Twelve collections, fre-
quently found beneath layers of decayed wood on fallen logs, at altitudes
from 4,400 to 6,800 feet, 7006, 1022, 1090, 1100, primarily from the Rim
Village area south to lower Annie Creek, found throughout the summer
months in both 1966 and 1967. The sporangia are readily recognized
since they are 0.4—0.6 mm in diameter and greenish to bright yellow.

DIDYMIACEAE

Diderma deplanatum Fries. Eleven collections on decayed wood, taken
from the Rim Village area southward to the south Park Entrance, North
Rim road and Kerr Valley, at elevations from 6,000 to 7,000 feet, 762,
955, 1031, 1105, June and July, 1967. The sporangia are white, sessile
and 1—1.5 mm in diameter. They are very fragile and lose their peridium
readily.

Diderma nigrum Kowalski. One collection, on coniferous twigs about
2 miles south of Park Headquarters, 6,200 feet, 7041, July 10, 1967. This
rare Myxomycete was recently described by Kowalski (1968). The spor-
angial dehiscence is star-shaped, revealing the white inner sporangial
walls and black capillitium.

I am indebted to Donald T. Kowalski for verifying the determinations
and for his assistance through the course of this investigation. This study
was supported in part by the Chico State College Foundation, grant
GU-1627.

1969 | REVIEWS 77

Department of Biology, Chico State College, Chico, California

LITERATURE CITED
Curtis, D. 1968. Barbeyella minutissima Meylan, A new record for the western
hemisphere. Mycologia. 60: 708—710.
HaGEIsTEIN, R. 1944. The Mycetozoa of North America. Mineola. 306 p.
KowatskI, D. 1966. A new species of Lamproderma from California. Mycologia.
58:808-810.
. 1966a. New records of Myxomycetes from California. I. Madrofo. 18:140—
142.
————. 1968. Three new species of Diderma. Mycologia. 60:595-603.
KowatskI, D. and D. Curtis. 1968. New Records of Myxomycetes from California.
III. Madrono. 19:246-249.
Martin, G. W. 1932. New Species of Slime Molds. J. Wash. Acad. Sci 22:88-92
. 1949. North American Flora. 1:1—190
Peck, M. and H. GrBert. 1931. Myxomycetes of Northwestern Oregon. Amer. ais
Bot. 19:131-147.

REVIEWS

The Evolution and Classification of Flowering Plants. By ARTHUR CRONQUIST.
x + 396 pp. Houghton Mifflin Co., Boston, 1968. Price $6.95.

In an invited paper presented at meetings commemorating the 50th anniversary
of the Botanical Society of America, held at the University of Connecticut in 1956,
this reviewer had the temerity to suggest that, “there seems to be rather general
agreement that sufficient evidence to formulate a really new, thorough-going, and
generally satisfactory phylogenetic arrangement of flowering plants is not yet avail-
able.” (Amer. J. Bot. 44: 88-92. 1957.) Later in the same meetings, however, two
new systems for at least part of the angiosperms were presented by Herbert F.
Copeland (Madrofo 14: 1-9. 1957) and Arthur Cronquist (Bull. Jard. Bot. Etat 27:
13-40. 1957), respectively, and Robert F. Thorne announced that he was working
toward the same goal (Aliso 6: 57-66. 1968). I do not recall that any of us were
then aware of the work of Takhtajan, which has subsequently assumed such major
importance.

The present volume is the outgrowth of that original Cronquist paper and is an
attempt to devise a general classification of angiosperms responsive to all presently
available pertinent information. The scope and variety of this information and its
application are impressive. It ranges from the more traditional morphology and
anatomy of the flower, fruit, and vegetative body, to pollen, embryology, and bio-
chemical characteristics. The author is especially partial to type of nectary, nuclear
constitution of pollen grains, details of ovular structure, nature of seminal food re-
Serves, distribution of vessels, and type of stomatal apparatus, among other features.
_ Cronquist emphasizes that while taxonomy is necessarily based on multiple corre-
lation of characters, a proper taxonomic system must also reflect (albeit muddily)
evolutionary relationships, and that development of taxonomy and the unraveling of
phylogeny each influences and strengthens the other. “A phylogenetic scheme which
Provides for all the available information and hangs together without serious inter-
‘nal contradictions is regarded as not only satisfactory but also something of a tri-
‘umph.” His classification is essentially one of consensus, in which he attempts to
capitalize on the various natural groupings that have been achieved in the past. It is
interesting to note how numerous these are on various levels. He asserts that if the
‘Tequirement of a strictly single (monophyletic) origin for groups is not insisted

‘upon too strictly, much of the apparent conflict between phylogeny and taxonomy

78 MADRONO [Vol. 20

disappears. He thinks the occurrence of evolutionary parallelism is itself an indi-
cator of relationships and should be taken into account. The evidence for the adap-
tive significance of many of the character combinations that distinguish orders and
families leaves him distinctly unsatisfied, and he repeatedly wonders aloud if the
unfashionable concept of “evolutionary momentum” (orthogenesis?) may not play
a role where selective impetus is obscure or undemonstrated.

He believes that angiosperms are a monophyletic group with ancestors somewhere
in the seed ferns, and that the primitive flowering plants were woody and probably
arborescent dicotyledons with magnolian/ranalian characteristics. Monocotyledons
must have been derived from aquatic dicotyledons which had lost their cambium
and hence the capacity to produce secondary growth and vessels in the normal way;
monocot leaves developed from modification of a bladeless petiole. Consistent with
his emphasis on consensus, Cronquist has adopted in major outline the system pro-
posed by Takhtajan (Taxon 13: 160-164. 1954). The angiosperms (re-christened
Magnoliphyta by Cronquist) are divided into dicots (Magnoliatae) and monocots
(Liliatae). The dicotyledons are construed as consisting of 6 subclasses—Magnoli-
idae, Hamamelidae, Caryophyllidae. Dilleniidae, Rosidae, and Asteridae—and the
monocotyledons as comprising 4 subclasses—Alismatidae, Commelinidae, Arecidae,
and Liliidae. Whereas Takhtajan admitted 61 orders of dicots and 21 orders of
monocots for a total of 82, Cronquist accepts 56 orders of the former group and 18
of the latter for a total of 74. Thorne, incidentally, eschews subclasses but recog-
nizes 19 superorders and 43 orders of dicots and 5 superorders and 11 orders of
monocots for a total of 54 orders. Although a good many differences in treatment do
in fact exist between the first two of these arrangements, and even more between
them and the last, the similarities are vastly more striking than are the differences.
As Cronquist remarks, “‘We are all—or nearly all—Besseyans.” It appears that we
may be in danger of becoming Takhtajanians, as well.

Cronquist provides keys to the subclasses, to the orders, and to the component
families. These must obviously allow for many exceptions, but they are useful. The
selective bibliography accompanying the discussion of each order should prove to be
even more useful. The writing is clear, concise, and positive, but the difficulties with
various taxonomic dispositions and the possibility of alternative choices are pointed
out frankly. The really fascinating aspect of the book is the opportunity afforded in
the running discussions of orders to find out what has happened to the groups of
one’s particular interest. If there is any danger in the treatment, it is that so many
of the long-standing controversies and indecisions seem to have been resolved so
easily and logically. It should be rewarding to see whether consensus widens or
diminishes as other books involving comparable schemes of classification appear, as|
they surely will. For the present, Cronquist has given us a very useful, well written, |
and stimulating volume in an uncrowded field of endeavor. —LINCOLN CONSTANCE,
Department of Botany, University of California, Berkeley.

Flora of Alaska and Neighboring Territories. By Eric HULTEN. xxii + 1008 PP
illustrated. Stanford Univ. Press. 1968. $35.00.

Eric Hultén’s preeminence among students of the Alaskan flora is a present-day.
example of how floristics research in a state or region tends to be dominated, for
long periods of time, by the outstanding work of a single individual. Although Pro-'
fessor Hultén’s principal interests, by his own admission, have been in the phyto-
geography of circumboreal floras, he has contributed to taxonomy such important
references as the Flora of Kamchatka (1927-1930), Flora of the Aleutian Islands
(1937, 1960), and Flora of Alaska and Yukon (1941-1950). As those who have used)
these books know, their purpose was to document scientifically the literature, col-
lections, nomenclature and distribution of arctic plants; and descriptions, illustra-
tions and keys are generally lacking. |
|

1969 | REVIEWS 79

In a charmingly personal preface to the present book, Hultén tells how he de-
cided to prepare what would be “. . . a flora of another character ... one that
would serve a larger public.” This work, Flora of Alaska and Neighboring Terri-
tories, is in all respects a great achievement. As a manual its primary purpose is that
of plant identification, and this is accomplished through concise keys, descriptions
and illustrations of all the species and most infraspecific taxa of Alaskan higher
plants (spermatophytes and vascular cryptogams). Other essentials are also in-
cluded—keys to families and genera, a glossary of terms, a list of botanical author-
ities, and bibliography. Especially remarkable, however, is the book’s content of
phytogeographical information, given in paired range maps for each taxon—a dot
map of the area covered by the manual, and an outline map of each entity’s com-
plete circumpolar range. These are Hultén’s unique contributions, derived from
more than 40 years of study of boreal floras and they add a highly useful dimen-
sion that is rarely found in regional floristic manuals.

Alaska is a very large place, and a large book is required to do justice to it. What
are the statistics of this flora? The area covered is Alaska, including the Aleutian
Islands, Yukon Territory, the northwest tip of British Columbia, and the Chukchi
Peninsula of Sibera—a total of 1,022,400 square miles. In it are, “Some 1,974 dis-
tinct, taxonomically named plants, belonging to 1,559 species, 412 genera,, and 89
families . . .’ The book also mentions hundreds of hybrids and over 200 closely re-
lated taxa occurring in neighboring boral regions. Not only are there large latitu-
dinal and altitudinal differences within this area, but the land is geologically com-
plex and includes major sections that were free of ice during the last two glacial
maxima. The importance of its central position in the migration route between the
Old and the New Worlds hardly needs mentioning. What better vantage point is
there to view, in panorama, the history and relationships of northern plant species ?

“A general condition of the flora of this region is that the morphological varia-
tion of a given taxon is greater in Alaska than in other parts of its range.’’ Hultén’s
taxonemic approach to this complex flora makes use of two traditional tools: a
conservative view of species, and a concept of subspecies as the major morpholog-
ically recognizable, geographical divisions of a species. Because the author has so
extensively revised the nomenclature of the flora, particularly at the subspecific
level, every reviewer will find much to comment on in groups he knows at first hand.
Species that we in the Pacific Northwest have recognized as distinct may run north-
ward and intergrade with others in Alaska or elsewhere in the arctic. As a result we
find, for example, Populus trichocarpa made a subspecies of P. balsamifera, ‘Beck-
mannia syzigachne” submerged in an asiatic subspecies of B. erucaeformis, and Phyl-
lodoce glanduliflora reduced to a subspecies of P. aleutica. Even more notable is
Hultén’s synonymizing of Aster foliaceus with A. subspicatus; the types of both of
these are Alaskan and rather similar, but farther south the names are applied to two
very distinct entities. Although such unions appear justified, one wonders about
other cases where evidence of intergradatien is slighted and the plants are kept as
distinct species. Tiarella trifoliata and T. unifoliata were shown by Kern to be inter-
fertile and intergrading, yet they are recognized here as species. Amelanchier florida
was placed by Hitchcock as a subspecies of A. alnifolia on the basis of their evident
intergradation, and they are not convincingly distinguished by Hultén’s descriptions
and illustrations. In Saxifraga, intergradation is admitted between S. davurica and
S. unalaschcensis, yet they are kept as species. A similar relationship seems to exist
between Hieracium triste and H. gracile, which nonetheless are not merged by
-Hultén. Very minor differences, principally of pubescence, appear to mark the two
species recognized in Romanzoffia as well as the three of Douglasia, whose geograph-
ical relationships resemble those of subspecies.

Some interesting changes in generic alignments can be noted, but forunately these
are minimal. The treatment is conservative in groups like Lycopodium, Claytonia,
and Chrysanthemum-Tanacetum. Minuartia is divided from Arenaria, Podagrostis
from Argostis, and Platanthera from Habenaria, however. Hultén’s conservative

80 MADRONO | Vol. 20

handling of apomictic and hybridizing groups is important in reducing the number
of named microspecies. In the difficult genus Salix there are 56 taxa described, in
Antennaria there are 19, in Arnica 17, and in Taraxacum only 11.

Many of the author’s nomenclatural changes appear to be derived from a rather
long paper published in Arkiv for Botanik, in 1967. Not having this paper available,
I have only noted a few of what appear to me to be unusual selections of names
for this flora. Alnus oregona should be A. rubra Bong., for example, the Betula-
Alnus rubra of Marshall being no homonym of the latter name. Stachys emersonii
is used by Hultén, although from Epling’s examination of the type it appears that
S. mexicana Benth. is an earlier name for this species. Finally, Echinopanax is used
in place of Oplopanax, although A. C. Smith, in the Flora of North America, states
that the former is a nomen nudum. The editorial work on this book is, overall, so
excellently done that it is mere nit-picking to call attention to minor errors. In a
few cases, however, illustrations seem to be significantly at variance with plant de-
scriptions; these were noticed for Ranunculus trichophyllus var. trichophyllus, where
floating leaves are not lacking from the drawing, for Amelanchier, where the key
leaf-shape differences are not apparent, and for Linnaea borealis ssp. longiflora, whose
leaves are hardly “elliptical, acute.” On the range maps, the circumpolar distribu-
tion of Papaver nudicaule is omitted, as is the occurrence on the Gaspé Peninsula
of Agoseris aurantiaca.

This review would be incomplete without the simple statement that Flora of
Alaska is a beautiful book! The difficult job of arranging keys, drawings, maps and
text has been solved with neatness and economy of space. Luxurious additions are
the physiographic maps on the end-boards and a section of superb colored plates
from photographs by the author. Among many small but helpful details in the
book are the habitat notes and information on type localities provided for all taxa,
an index of common names, and pronunciation guides for the scientific names. With
such careful attention to details evident throughout the work, two omissions stand
out quite noticeably—there are no generic descriptions and no index to synonyms.
The latter would have helped this reviewer discover how the genus Youngia came to
be lost, before he noticed that its only North American species, VY. americana, re-
sides on page 956 in the synonymy of Crepis nana.

It will be apparent to all who use this book that the Alaskan flora contains a
goldmine of biosystematic problems that have hardly been touched. Hultén has
brought to a high level of refinement the taxonomic knowledge that can be gained
from morphological and geographical evidence, but the genetic, cytological and bio-
chemical information necessary for a total synthesis of relationship is largely lacking.
The difficulties of integrating biosystematic data with existing morphological cate-
gories are already clear from what is known of the cytotaxonomy of such “difficult”
groups as Epilobium angustifolium (Mosquin), Calamagrostis canadensis (Love,
Mitchell, and others) and Achillea millefolium, sens. lat. (Ehrendorfer, Mulligan and
Basset). The flora of Alaska will amply repay careful study by generations of fu-_
ture botanists, all of whom will be indebted to Eric Hultén for this landmark in the >
botany of North America. —Kenton L. CHAMBERs, Oregon State University, Cor- |
vallis.

A WEST AMERICAN JOURNAL OF BOTANY

INFORMATION FOR CONTRIBUTORS

Shorter items, such as range extensions and other biological notes,
will be published in condensed form with a suitable title under the general
heading, “Notes and News.”

Institutional abbreviations in specimen citations should follow Lanjouw
and stafleu’s list (Index Herbariorum, Part 1. The Herbaria of the World.
Utrecht. Fifth Edition, 1964).

Abbreviations of botanical journals should follow those in Botanico-
Periodicum-Huntianum (Hunt Botanical Library, Carnegie-Mellon Uni-
versity, Pittsburgh, Pennsylvania, 1968).

_ Membership in the California Botanical Society is normally considered
_ a requisite for publication in MApRONo.

VOLUME 20, NUMBER 3 JULY, 1969

Contents

NEOGENE FLORISTIC AND VEGETATIONAL
HIsTorY OF THE PACIFIC NORTHWEST,
Jack A. Wolfe | 83

EcoLocic PLANT GEOGRAPHY OF THE PACIFIC
NortHwEST, R. Daubenmire 111

SoIL DIVERSITY AND THE DISTRIBUTION
OF PLANTS, WITH EXAMPLES FROM
WESTERN NorTH America, A. R. Kruckeberg _ 129

PHYTOGEOGRAPHY OF NORTHWESTERN NorTH
AMERICA: BRYOPHYTES AND VASCULAR
Prants, W. B. Schofield 155

Pah HS ss
of a Oo Na

SEP 1 8 1969

WL/BRARIE?

A WEST AMERICAN JOURNAL OF BOTANY > SPB

3 BLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

BOARD OF EDITORS

LyMAN BENSON, Pomona College, Claremont, California
KENTON L. CHAMBERS, Oregon State University, Corvallis
Joun F. Davipson, University of Nebraska, Lincoln
WALLACE R. Ernst, Smithsonian Institution, Washington, D. C.
ArTURO GOMEZ PomPa, Universidad Nacional Autonoma de México
EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia
Mivprep E. Maruias, University of California, Los Angeles
RoBERT OrNDUFF, University of California, Berkeley
Marion Ownsey, Washington State University, Pullman
Duncan M. Porter, Missouri Botanical Garden, St. Louis
REED C. Ro iins, Harvard University, Cambridge, Massachusetts
IrA L. Wiccins, Stanford University, Stanford, California

Editor — JoHN H. THOMAS
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCASKILL
P. O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Harry Thiers, Department of Ecology and Systematic Biology, San
Francisco State College. First Vice-President: Grady Webster, Department of Bot-
any, University of California, Davis. Second Vice-President: Ernest C. Twisselmann,
Cholame, California. Recording Secretary: John West, Department of Botany, Uni-
versity of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Depart-
ment of Botany, University of California, Berkeley. Treasurer: June McCaskill, De-
partment of Botany, University of California, Davis.

The Council of the Calfornia Botanical Society consists of the officers listed
above plus the immediate past President, Elizabeth McClintock, California Academy
of Sciences, San Francisco; the Editor of Madronio; and three elected Council Mem-
bers: Annetta Carter, Department of Botany, University of California, Berkeley;
Robert Ornduff, Department of Botany, University of California, Berkeley; and
Malcolm Nobs, Carnegie Institution of Washington, Stanford.

A SPECIAL ISSUE OF

MADRONO
A WEST AMERICAN JOURNAL OF BOTANY

PUBLISHED BY THE CALIFORNIA BOTANICAL SOCIETY, INC.

In honor of the botanists

attending the

XI INTERNATIONAL
BOTANICAL CONGRESS

UNIVERSITY OF WASHINGTON
Seattle, Washington, U.S. A.
August 24—September 2, 1969

Maprono, Vol. 20, No. 3, pp. 81-208. August 20, 1969.

NEOGENE FLORISTIC AND VEGETATIONAL HISTORY
OF THE PACIFIC NORTHWEST

Jack A. WOLFE

The Neogene fossil plant assemblages of the Pacific Northwest are
more numerous and more completely studied than the Neogene assem-
blages of any other area of North America (fig. 1). Most of the work has
been based on the study of plant megafossils—particularly leaves—which
provide a valid basis for the reconstruction of lineages and hence floristic
history. The increasing body of palynologic data, on the other hand, pro-
vides an insight into vegetational history, which is difficult to reconstruct
from megafossils that represent largely the specialized streamside and
lakeside vegetation. Both megafossil and microfossil assemblages typically
represent ligneous plants, although microfossil assemblages include more
representatives of herbaceous plants than do megafossil assemblages. This
report will thus primarily concern woody plants.

By piecing together lineages and by analyzing the vegetational types
in which the lineages have lived, it is possible to understand the develop-
ment of vegetation in terms of its floristic elements. Proponents of vari-
ous “‘geofloral” concepts have unfortunately confused flora and vegeta-
tion, which inherently leads to a confusion of floristic and vegetational
history. It is extremely improbable from the genetic and physiological
viewpoint that many lineages could have remained in association through-
out the Tertiary; that is, that a given vegetational type remained florist-
ically unchanged (Mason, 1947; MacGinitie, 1962; Wolfe, 1964). Re-
cent work has indeed shown that many “‘Madro-Tertiary” elements in
Nevada represent lineages that were present earlier in the mesic Miocene
vegetation of the Pacific Northwest (Wolfe, 1964). Work in Alaska has
also shown that the concept of an ‘“‘Arcto-Tertiary Geoflora” is invalid
(Wolfe, et al., 1966; Wolfe, 1966; 1969; Wolfe and Leopold, 1967;

- Wahrhafting, et al., 1969; Hopkins, et al., in press). The discussion of

the Neogene of the Pacific Northwest involves an understanding of the
history not only of that area but of much of northwestern North America.

I wish to thank H. D. MacGinitie and H. E. Schorn, University of Cali-

fornia, Berkeley, and E. B. Leopold, U.S. Geological Survey, for their

helpful discussions of the subjects covered in this report and for their

“critical reading of the manuscript. K. M. Piel, Union Oil Company of

California, kindly provided his unpublished pollen count for the late Mio-
| cene Quesnel diatomite of British Columbia.

1]
it

Publication of this paper has been authorized by the Director, U.S.

Geological Survey.

84. MADRONO

; vA 3;
“0 Pliocene yi oe
A late - j
ihe ‘i
/ x middle > Miocene if
140" mearly ) ~/ /

Lo /

/ )
/
/
/
L /
—— 2 ee
35°" 1252
Oo 100 200 300 400 500 MILES
ess) Ea ee ee ee ed
100 O 100 200 300 400 500 KILOMETERS

Ee ee a

Fic. 1. Location of some Neogene and early Pleistocene assemblages in northwestern
North America. Numbers correspond to assemblages as follows:

Early Miocene

1. Capps Glacier 2. Collawash 3. Eagle Creek
4. Maupin 5. Seldovia Point 6. Upper Cedarville
7. Upper Healy Creek

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 85

METHODS OF DETERMINING VEGETATION

The paleobotanist, after completing identification of the fossils from a
given assemblage or series of assemblages, has at least a partial list of the
flora. The significance of such a list in terms of vegetation is not agreed
on by all paleobotanists. Most poleobotanists working on Tertiary as-
semblages of western North America have used a two-fold approach to
determine vegetation: 1, a strict application of uniformitarianism to the
tolerances of a given lineage, genus, or family (i.e., tolerances have not
changed through time), and 2, counts of the megafossils.

That lineages have had different tolerances in the past than at present
should be obvious from theoretical considerations. Assuming that the
dicotyledons, for example, at one time evolved in a limited area that had
a limited number of habitats and climates, during the spread outward
from such an area the organisms in any lineage would have met new en-
vironmental conditions; the same reasoning can be applied to the lineages
comprising any genus or subgenus. The fact that the dicotyledons today
occupy an extremely wide range of habitats is an indication that lineages
are capable of adapting to new environmental conditions. The fossil rec-
ord in fact supports such a concept of changing tolerances: one lineage of
Pterocarya, for example, first appears in North America during the Eocene
in subtropical and tropical forests (Wolfe, 1968). During the Oligocene,
the lineage is represented in tropical forest during the early part of that
epoch, but by the late Oligocene it was present in warm temperate vege-

Middle Miocene

59. Troutdale

60. Type Clamgulchian

8. Cache Creek _ 9. Cape Blanco 10. Fingerrock
11. Fish Creek 12. Frederika 13. Grand Coulee
14. Houston 15. Latah 16. Mascall
17. Middlegate 18. Monument 19. Rockville
20. San Antonio 21. Suntrana 22. Unga Island
23. Wishkaw River
Late Miocene
— 24. Aldrich Station 25. Blue Mountains 26. Brock Road
27. Chalk Hills 28. Chloropagus 29. Chuitna River
30. Fallon 31. Faraday 32. Grubstake
33. Hidden Lake 34. Hog Creek 35. Lower Ellensburg
_ 36. Marble Point 37. Mashel 38. Neroly
| 39. Pit River 40. Pribilof Canyon 41. Quesnell
| 42. Skonun 43. Skunk Creek 44. Stewart Spring
. 45. Stinking Water 46. Table Mountain 47. Thorn Creek
_ 48. Trapper Creek 49. Trout Creek 50. Type Homerian
51. Weyerhauser 52. Wilkes
Pliocene and early Pleistocene
53. Bering Platform 54. Dalles 55. Deschutes
_ 56. Elk River 57. Middle Ellensburg 58. Rattlesnake

86 MADRONO [Vol. 20

tation (Wolfe, 1959). During the early and middle Miocene, the lineage
participated in the Mixed Mesophytic forest, and during the late Miocene
the lineage is represented only in conifer forest. Pterocarya is today rep-
resented only in broad-leaved deciduous forests. In most instances, of
course, tolerances of the Neogene representatives of a given genus or
lineage probably more closely approximate the present tolerances of the
genus or lineage than do the Paleogene represntativs. Note, however,
that Chamaecyparis nootkatensis participated in vegetation that repre-
sents a conifer-live oak association during the late Miocene in southwest-
ern Nevada (Wolfe, 1964); clearly C. nootkatensis included during the
late Miocene physiological races that are no longer extant. Uniformi-
tarianism should be applied cautiously to vegetational reconstructions.

If, of course, the ‘‘associational method” of determining the flora is
utilized, such anomalous associations as that of Chamaecyparis nootka-
tensis and Quercus chrysolepsis will not be known. The “‘holotype” of the
associational method of determination (Cain, 1944, p. 43) is in fact an
excellent demonstration of the weakness of the method. Knowlton (1902)
originally described Cinnamomum bendirei from the Bridge Creek assem-
blage of Oregon. Chaney (1927), however, interpreted this assemblage as
a redwood forest and thus considered Cinnamomum an incongruous ele-
ment. He therefore transferred Knowlton’s species to Philadelphus, which
would be expected in a redwood forest. Brown (1940) pointed out that
the leaves morphologically could not be Philadelphus but had the diag-
nostic characters of Lauraceae; he thus transferred the species to Sassa-
fras. This transfer was ignored by Chaney (194b, p. 350), who still con-
sidered that Philadelphus was a better choice for a redwood forest. After
the discovery of the living Metasequoia and the realization that the
Bridge Creek assemblage contained not Sequoza but rather Metasequoza,
Chaney (1952) accepted Brown’s transfer of the species to Sassafras. The
leaves, however, have the small areoles lacking branched, freely ending
veinlets and the continuous marginal vein of Cinnamomum, Lindera, and
Neolitsea,; Sassafras has large areoles intruded by branching veinlets and
lacks a marginal vein (Wolfe, 1960). Knowlton’s determination was more
valid than either Chaney’s or Brown’s.

The second widely used method of reconstructing past vegetation is
through the use of leaf counts. The basic assumption of this method is
that the representation of leaves in a fossil bed is proportional to the rep-
resentation of the plants that bore them in the ancient forets. Chaney
(1959) has discussed the variables affecting such an assumption. and was
forced to the conclusion (p. 46) that the assumption was perhaps valid
only for generalities. Without any statistical basis, Chaney (1959, p. 46)
stated: “I believe that any species that has provided as much as one-
fifteenth [about 7 percent] of the record of foliage and fruit in a fossil
flora must have been numerous enough to be considered abundant in the
forest .. .”’” Whether Chaney’s belief is valid is, of course, unknown.

It is extremely doubtful that the relative representation of plant mega-

1969] WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 87

fossils is significant in reconstructing ancient vegetation. Megafossil as-
semblages contain an overrepresentation of fluviatile and lacustrine spe-
cies that grew at or near the site of deposition (MacGinitie, 1953, p. 46;
Fegri and Iversen, 1964, p. 39). As yet, no one has demonstrated any
sizable correlation between the relative representation of organisms and
the relative representation of their larger organs either on the forest floor
or at the sites of deposition. One attempt to do so is that of Chaney (1924).
The correlation coefficients he obtained were indeed sufficiently high to
indicate that leaf counts might be valuable in reconstructions of vegeta-
tion. The only species for which Chaney gave the raw data is Alnus rubra
[= A. oregona|, the coefficient computed was 0.49. Note, however, that
negative matches, situations in which neither organisms or remains of this
species were found, were included in the computations. Such an inclusion
hardly seems justifiable; the coefficient would be even higher if a worker
included negative matches from Nevada, where the species does not grow.
Recomputation of the correlation coefficient after elimination of all nega-
tive matches yields a coefficient of 0.35, 1.e., very close to a universe in
which no correlation exists. I conclude that although there is a slight cor-
relation, it is insufficient for leaf counts to be considered useful in the
reconstruction of vegetation.

Fossil leaves are, however, not entirely useless in determining vegeta-
tion. The physiognomic features of foliage are largely independent of tax-
onomy and appear to be highly sensitive to the environment (Richards,
1952, p. 154). The correlation between vegetational types and the leaf
margin—whether entire or nonentire—is striking (Bailey and Sinnott,
1915, 1916). Several paleobotanists have applied this correlation to vege-
tational interpretations of fossil assemblages. In general, in a mesic cli-
mate, the percentages of species that have entire margined leaves can be
correlated to vegetational type as follows:

76+. Tropical Rain forest
57-75 Paratropical Rain forest (extratropical rain forest of Wang,
1961; Subtropical Rain forest of Richards, 1952)
40-56 Subtropical forest
10-35 Temperate forest

Summer dryness and extreme winter low temperatures increase the per-
centage for the temperate forest; this increase is probably related to the
physiological aridity of both far northern and truly arid climates. In
Alaska, for example, the leaf margin percentage is 42, but a large number
of the woody dicot species that have entire margins are thick- and small-
leaved Ericaceae. The leaves of species of mid latitude, arid environments
show similar adaptations. A consideration of leaf size should, therefore,
be considered along with the type of leaf margin.

For Neogene assemblages, the microfossil assemblages are probably an
excellent source of data for reconstructions of vegetation. Microfossil as-

88 MADRONO [Vol. 20

semblages do not, insofar as the pollen of woody plants is concerned, rep-
resent local vegetation only; the pollen rain of a given region appears to
be rather uniform and reflects the predominant vegetation type in the
region (Davis and Goodlett, 1960). The variables affecting the interpre-
tation of a pollen diagram have been and are being investigated; correc-
tive factors that take into account many of the variables have been pro-
posed (see for example, Fegri and Iversen, 1964, p. 99-123). Although
some of the variables, for example, amount of pollen production, may
have changed for various genera, the comparison of two approximately
isochronous spectra or diagrams should yield a reasonable idea of the dis-
tribution of vegetational types during that interval of time. It should be
emphasized, however, that interpretation of a single spectrum can be
highly misleading; in a region of active volcanism, for example, one major
eruption could drastically change the regional vegetation and hence pollen
rain. Reconstruction of even a short diagram through several feet of sec-
tion should help eliminate the effect of such short term changes in
vegetation.

EFFECT OF ALTITUDE

Some workers have argued that previous interpretations of vegetational
and floristic history in the Pacific Northwest have failed to take into ac-
count the varying altitudes at which the fossil assemblages lived. Thus
Axelrod (1964) considered that a particular group of assemblages from
the Pacific Northwest is isochronous and of middle Miocene age; these
assemblages purportedly show a strong floristic and vegetational zonation
according to altitude. The only method of testing the validity of Axelrod’s
conclusion is to demonstrate the contemporaneity of the assemblages.
Various data—mammalian, radiometric, diatom, freshwater molluscan,
and stratigraphic—all of which are high reliable for determining relative
ages in the Neogene rocks of the Pacific Northwest, are available for sev-
eral of these assemblages:

Basis for Probable
Assemblage age assignment Altitude® “absolute” age
Trapper Creek diatoms, mollusks, 3,000ft. 11-12 m.y.
stratigraphy!
Trout Creek radiometric” 2,300 13.1 m.y.
Mascall radiometric” 1,500 15.4 m.y.
Upper Cedarville radiometric” 1,100 19.8 m.y.
Rockville (“Succor Creek,” in part) radiometric? 600 16.7 m.y.
Grand Coulee radiometric* 250 15.7-16.8 m.y.
Latah radiometric?:4 250 15.5-20.6 m.y.
1 Mapel and Hail, 1959. 4 Gray and Kittleman, 1967.
* Evernden and James, 1964. 5° Axelrod, 1964.

3 Obradovich, unpub. data.

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 89

These data illustrate that age, not altitude, is the important factor. The
early Miocene Upper Cedarville flora represents a cool time interval as
compared with the warmer (“lower altitude”) middle Miocene assem-
blages such as the Grand Coulee, Latah, Mascall, and Rockville (Wolfe
and Hopkins, 1967). The age of the Upper Cedarville has been a matter
of considerable discussion. I (1964), for example, have considered this as-
semblage to be of late Miocene age. The problem arises from the fact that
LaMotte (1936) included in this flora material from ’49 Camp, where the
early Miocene radiometric age was obtained, and from the Pit River area
many miles to the east. The Pit River assemblage contains Platanus dis-
secta Lesq., which is a typically late Miocene species. The Upper Cedar-
ville (‘49 Camp) and Pit River are here considered as distinct floras. The
Rockville-Succor Creek problem is analogous to that of the Upper Cedar-
ville-Pit River. The Succor Creek (late Miocene) and Rockville (middle
Miocene) assemblages are here considered as distinct.

The late Miocene assemblages likewise represent a cooler time interval
than do the middle Miocene assemblages. These middle and late Miocene
assemblages do show a gradation from lower to higher, but this gradation
is stratigraphic and not altitudinal.

EARLY MIOCENE

The concept of early Miocene varies from one paleontological discipline
to another. The mammalian workers use the term Arikareean for this in-
terval; radiometric dates indicate a duration for the Arikareean from
about 21 to 25 or 26 million years ago (Evernden, et al., 1964). The
foraminiferal workers, on the other hand, consider their Saucesian Stage
to be basal early Miocene, and the base of this stage has a radiometric
age of 22.5 million years (Turner, 1968). The moluscan paleontologists
consider their Vaqueros “‘stage”’ to be of earliest middle Miocene age and
this is thought to be equivalent in the foraminiferal sequence to the
Saucesian stage. The base of the Miocene in western North America thus
can be placed from about 23 to 28 million years, depending on which
biostratigraphic framework is accepted. The only plant assemblages in
this time interval (early Miocene) that have radiometric dates are the
small Maupin assemblage, the upper member of the John Day Formation,
which is younger than 23 million years (Evernden, et al., 1964) and the
Upper Cedarville with an age of 19.8 million years (Evernden and James,
1964). These assemblages are, on paleobotanical grounds, correlative to
the Miocene zone 2 of Wolfe (1962) and to the lower part of the Seldo-
vian Stage of the paleobotanical geochronology. The lower Seldovian as-
semblages which are the oldest under consideration here may thus not be
basal Miocene and may represent only the later part, 19 to 23 or 24 mil-
lion years, of the early Miocene.

The known lower Seldovian assemblages are well represented in Alaska
and in conterminous United States. In Alaska, they include the Seldovia
Point, Capps Glacier, and Upper Healy Creek assemblages (Wolfe, et al.,

90 MADRONO [ Vol. 20

1966; Wolfe, 1966; Wahrhaftig, et al., 1969). A total of about 70 mega-
fossil entities is known, and the pollen floras have also been extensively
studied. In conterminous United States, aside from the Maupin (Wolfe,
unpublished data) and Upper Cedarville (LaMotte, 1936), lower Seldo-
vian assemblages include the Collawash (Wolfe, unpublished data) and
Eagle Creek (Chaney, 1920). Approximately 160 megafossil entitles are
known, but, except for the Collawash, the microfossil assemblages appar-
ently have not been studied.

The Alaskan assemblages are preserved in rocks that, for the most part,
were deposited in large coal basins at low altitudes. The Upper Healy
Creek beds were deposited at least 300 miles from the coast, and the
Capps Glacier assemblage may have been only about 100 to 150 miles
from the coast. The Seldovia Point assemblage was nearer to the coast
than the other two assemblages but the beds containing their assemblage
were deposited in a valley in an area of moderate relief (Wolfe, et al.,
1966).

The topographic setting of the assemblages in Oregon is known with
reasonable certainty. The basalts of the Columbia River Group buried
and thus preserved some of the early Miocene topography in northwest-
ern Oregon. At the time, the Cascades were probably more than 1,500 feet
in height (Peck, et al., 1964, p. 28); the Collawash beds are at about the
altitude at which the basalts thinned out against the range. To the north,
in the area of Eagle Creek deposition, the basalts attain a thickness of
about 2,500 feet. The distribution and thickness of the basalts thus indi-
cates that the Eagle Creek assemblage probably was in a broad valley not
far above sea level, whereas the Collawash assemblage was at about 2,500
feet elevation on the northern margin of the Cascade Range in Oregon.
The original elevation of the Maupin assemblage is somewhat uncertain,
but the small size of the assemblage makes its vegetational and floristic
significance impossible to evaluate at this time.

The original elevation of the Upper Cedarville beds is extremely un-
certain; certainly northwestern Nevada had a considerable elevation even
in the early Miocene, but an altitudinal interpretation based on geologic
data would be highly conjectural.

FiLora. The lower Seldovian flora is known to contain at least 180
megafossil species, and, combined with the microfossil floras, over 200
species are represented. The flora is particularly rich in species of Taxo-
diaceae, Salicaceae, Juglandaceae, Betulaceae, Fagaceae, Rosaceae, and
Aceraceae; also typically present are Ulmas, Zelkova, Cocculus, Liquid-
ambar, and Platanus.

The sources of many lineages in the lower Seldovian floras are known
with reasonable certainty. One source is earlier vegetation of the Pacific
Northwest; the lineages occurred in vegetation that represented Para-
tropical Rain forests, i.e., similar to the vegetation of lowland Taiwan
and Hong Kong, or Subtropical forests. These lineages have displayed a

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 91

considerable adaptability to changes in climate. Another source is the
Paleogene temperate to marginally subtropical upland assemblages along
the Rocky Mountains (Wolfe, 1960); some of these lineages had already
appeared in the Pacific Northwest by the time that temperate vegetation
occupied this area in the later Oligocene. A third, although minor, source
is represented by lineages that first appear in the later Oligocene tem-
perate vegetation of Alaska.

The lineages of Fagaceae for the most part are of unknown descent.
One of the particularly striking features of Miocene as compared with
older floras is the presence of “lobed” oaks, particularly the black oaks.
The black “lobed” oaks are apparently restricted to North America, al-
though Chelebaeva (1968) has referred a fragmentary specimen from
the middle Miocene of Kamchatka to this group. The predecessors of the
“lobed” black oaks must almost certainly be within the other members
of the subgenus Erythrobalanus, and possibly forms such as Quercus
peritula Cocker. from the marginally subtropical Florissant assemblage
of the early Oligocene of the Rocky Mountains or Q. pregrahami MacG.
from the subtropical Weaverville assemblage of the late Oligocene of
California may be ancestral. The origin of the “lobed” white oaks is even
more problematic. MacGinitie (1953) considers the subgeneric assign-
ment of the Florissant Q. lyratiformis Cocker. to be dubious, and this is
the only pre-Miocene material from North America known to me that
has been referred to Leucobalanus. The close relationship between the
extant east American and the Miocene west American members of Leuco-
balanus indicates the probability that there was a common source in the
Oligocene of the Rocky Mountains.

Some regional floristic differentiation is apparent in the early Miocene
of northwestern North America. Most species of Salicaceae, for example,
are distinct between Alaska and the Pacific Northwest. Rosaceae are
more diverse in the Pacific Northwest and Lauraceae are unknown in
Alaska. The Alaskan flora has a definite Asian element that did not reach
the Pacific Northwest: Acer fatisiifolia, A. ezoanum, Kalopanax, Ulmus
longifolia, Populus reniformis. More than half the known Alaskan lower
Seldovian species are, however, also known in the Pacific Northwest, thus
indicating that the two areas should be considered as parts of the same
floristic province during the early Miocene.

Altitudinal zonation of the flora in the Pacific Northwest does not ap-
pear to have been pronounced during the early Miocene. Most species
known from the low altitude Eagle Creek flora are also known in the up-
land Collawash flora. Considering the little latitudinal floristic zonation
during the early Miocene, it should be expected that altitudinal floristic
zonation would also be slight.

VEGETATION. The vegetation of the early Seldovian of northwestern
North America was broad-leved deciduous. An apparently continuous
deciduous forest extended from Japan and northern China north into

92 MADRONO [Vol. 20

Alaska and south into the Pacific Northwest. Evergreens were an impor-
tain part of this forest; coniferous evergreens became increasingly im-
portant in the north and broad-leaved evergreens in the south.

The Alaskan pollen assemblanges from the Cook Inlet region indicate
that locally the conifers such as Picea were present in the lowland for-
est—presumably because of the cool summers (Wolfe and Leopold,
1967). In the interior of Alaska, however, only deciduous conifers of
Taxodiaceae appeared in significant numbers with the broad-leaved decid-
uous plants. Broad-leaved evergreens were apparently rare in Alaska.
The leaf margin percentage for the Alaskan lower Seldovian flora is 15,
i.e., similar to that for the temperate forest of the Mid-Atlantic Staes
and the northern border of the Mixed Mesophytic forest in China today.

In the Pacific Northwest, broad-leaved evergreens were more diverse
and include: Quercus, Magnolia, Cinnamomum, Litsea, Persea, Umbrel-
lularia, Exbucklandia, Cercocarpus, Lyonothamnus, Garrya, and Ar-
butus. The leaf margin percentage for a large assemblage such as the
Collawash is 25, i.e., similar to that for the region occupied by the Mixed
Mesophytic forest in central China. The pollen assemblages from the
Collawash beds contain only minor amounts of Picea, indicating that
coniferous forest was not present even at 2,500 feet altitude. In eastern
Asia today the Mixed Mesophytic forets has a latitudinal range of about
14°, but in western North America this vegetational type spanned at
least 25° of latitude; even in the early Miocene latitudinal zonation of
vegetation was much less pronounced than today.

MIDDLE MIOCENE

In this report, the term middle Miocene denotes an interval from
about 14 or 15 to about 19 million years age. This interval is equivalent
to the Hemingfordian and early Barstovian ages of the mammalian
paleontologists. Assemblages of this age in Alaska include the Unga
Island (locs. P9978, P9993 of Burk, 1965), Suntrana (Wahrhaftig,
et al., 1969), two assemblages in the Kenai formation, the Houston and
Cache Creek (Wolfe, et al., 1966) and the Frederika (Wolfe, unpub-
lished data). In the Pacific Northwest, assemblages of this age include
the Fish Creek, Cape Blanco, Wishkaw River, and Monument (Wolfe,
unpublished data), as well as published assemblages such as the Latah
(see Chaney and Axelrod, 1959, for list of references), Grand Coullee
(Berry, 1931), Mascall (Chaney and Axelrod, 1959), and Rockville
(Graham, 1965), which is, in part, the Succor Creek flora of some au-
thors. In Nevada, only two middle Miocene assemblages have been de-
scribed, the Middlegate (Axelrod, 1956) and the Fingerrock (Wolfe,
1964); an additional assemblage is known from the San Antonio Range
(Wolfe, unpublished data) and three other assemblages are currently
under study by Axelrod. In central California, no assemblages of middle
Miocene age have been thoroughly studied.

The depositional setting of the assemblages from Alaska is, with one

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 93

exception, lowland. The exception is the Frederika, which is preserved
in beds that were deposited in an area of considerable relief. These beds
were formed in part because of the damming of the drainage by the first
extrusion of the Wrangell lavas (E. M. MacKevett, pers. comm., March
1969).

In the Pacific Northwest, both the Wishkaw River and Cape Blanco
assemblages occur in intertonguing marine-nonmarine rocks, and hence
represent coastal lowland vegetation. The Fish Creek assemblage occurs
in beds that overlie the basalts of the Columbia River Group; presum-
ably the Fish Creek assemblage lived at about the same elevation as the
early Miocene Collawash assemblage, 1.e., about 2,500 feet. In eastern
Oregon, the Monument assemblage, which occurs in beds that are pre-
Columbia River Group but post-John Day Formation, probably grew in
a region of little relief. The Mascall assemblage probably grew in an up-
land basin surrounded by considerable relief, to judge from the thick
volcanic sequence that grades laterally into the Mascall Formation
(Thayer and Brown, 1966). The Latah assemblage occurs in lake beds
that were formed by the damming of the drainage by the lavas of the
Columbia River Group (Pardee and Bryan, 1926). Pardee and Bryan
suggest that the lavas, which are much thicker to the east and south
than it is now, i.e., higher than 1,500 to 2,000 feet (A. B. Griggs, pers.
the Latah region by a chain of hills; the lavas finally attained a thick-
ness sufficient to flow over the hills and ended deposition of the Latah.
Several miles to the east of the Latah basin, the lavas lap onto the old
granitic highland. These data indicate that the Latah Formation was
deposited in an upland basin; if anything, the Latah basin was higher
than it is now, i.e., highe rthan 1,500 to 2,000 feet (A. B. Griggs, pers.
comm., March 1969). Geologic data bearing on the original altitude of
the Rockville and the Middle Miocene assemblages from Nevada is lack-
ing, although it is presumed that these regions were uplands of at least
moderate elevation.

Friora. The flora of the middle Miocene (upper Seldovian) in Alaska
and the Pacific Northwest differs little from that of the early Miocene.
An almost complete generic list for the early and middle Miocene is
given in Table 1. Almost all lineages in the late Seldovian were also
represented in the early Seldovian. Some groups, e.g., Salicaceae, were
more diverse in Alaska during the late Seldovian than during the early
Seldovian, and possibly this represents diversification of the family.
Pinaceae are better represented in the megafossil floras than previously,
particularly late in the middle Miocene. A fir related to the extant 4 dies
bracteata has an earlier record in the upland conifer forests of the
Oligo-Miocene of the Rocky Mountains and makes its first appearance
in the Pacific Northwest during the middle Miocene.

In Alaska one of the few upland assemblages of Miocene age, the
assemblages from the middle Miocene Frederika Formation, contains a

94 MADRONO [Vol. 20

diversity of Pinaceae in contrast to the lowland Alaskan assemblages.
Included in the Frederika assemblage are Abies, Picea, Pinus, and Tsuga,
which are accompanied by Pterocarya, Fagus, Ulmus, and Acer.

Over 230 species and 110 genera of presumed ligneous plants are now
known in the early and middle Miocene of northwestern North America.
Considering the incompleteness of the fossil record, the richness is im-
pressive. Despite extensive search, about 20 species with highly distinc-
tive leaves from the Collawash assemblage have yet to be identified;
it is conceivable that some of the leaves belong to extinct genera analog-
ous to the epibiotic and/or monotypic genera of the extant Mixed Meso-
phytic forest of eastern Asia.

TABLE 1. COMPOSITION OF THE EARLY AND MIDDLE MIOCENE FLORA OF
NORTHWESTERN NortH AMERICA.

‘“‘y”? denotes a record based largely or entirely on pollen. PNW = Pacific North-
west.

species Alaska PNW aane Alaska PNW
Ginkgo 1 1 1 Celtis 1 0 1
Cephalotaxus 1 0 1 Ulmus 4 2 3
Abies 4 2p 3 Zelkova 1 1 1
aff. Cedrus 1 p 1 Schoepfia/Anacolosa 1p 0 1p
Keteleeria 1 0) 1 Aristolochia 1 0 1
Picea 2 1 2 Cercidiphyllum 1 i 1
Pinus 3 2 3 Clematis 2 0 2
Pseudotsuga 1 p 1 Mahonia 3 0 3
Tsuga 1 3p 0) Cocculus 1 1 1
Chamaecy paris 1 1 ©) Liriodendron 1 0 1
Fokienia 1 p 1 Magnolia 1 0 1
Calocedrus 1 0 1 “Laurophyllum” 6 0) 6
Thuja 1 1 1 Cinnamomophyllum 1 0) 1
Cunning hamia 1 0 1 Sassafras 1 0 1
Gly ptostrobus 2 1 1 Hydrangea 1 1 1
Metasequota 1 1 1 Itea aly p p
Sequoia 1 1 1 Exbucklandia 1 0 1
Taxodium 1 1 1 Fothergilla 1 1 1
Populus 12 7 6 Liquidambar 1 1 1
Salix 13 8 6 Platanus 1 1 1
Comptonia 1 1 0 Amelanchier 2 0 y
Carya 4 3 4 Cercocarpus 1 0) 1
Juglans 2 1 1 Crataegus 3 1 3
Pterocarya 4 3 4 Holodiscus 1 0 1
Alnut 7 5 7 Lyonothamnus 1 0 1
Betula $ 2 2 Prunus 3 1 8)
Ostrya 3 2 1 aff. Peraphyllum 1 0) 1
Ostryo psis 1 1 ) Pyrus 1 @) if
Castanea 1 0 1 Rosa 1 0 1
Castano psis 1h 0) 1 Rubus 1 0 1
Fagus 7 2 5 Sorbus 2 1 1
Quercus 11 3 10 Spiraea 1 1 0

|
'

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 95

No. No.
species Alaska PNW species Alaska PNW
Albizzia 1 0 1 X ylonagra 1 0 1p
Cercis 1 0 1 Gordonia 1 0) 1
Cladrastis 1 1 1 Idesia il 0 1
Gymnocladus 1 0 1 Shepherdia 1 1 0
Sophora 1 0 1 Cornus 2 0) 2
Ptelea 1 ) 1 Nyssa 1 1 1
Ailanthus 1 @) 1 Oreopanax 1 0 1
Cedrela 1 0) 1 Alangium 1 1 0
Rhus 1 0 1 Clethra 1 0 1
T oxicodendron 1 0 1 Arbutus 2 0 2
Pistacia 1 ) 1 Leucothoe 1 0 1
Tlex 4 p 4 Rhododendron 2 1 1
Acer ie, 7 8 Diospyros 1 0 1
Aesculus Oye. 1 i Kalopanax 1 1 0
Allophylus 1 0 1 Halesia 1) 0 1
Ceanothus i 0 1 Fraxinus 3 1 3
Colubrina 1 0) 1 Catalpa 1 0) 1
Karwinskia iL 0) 1 Diervilla 1p 1p 0
Sageretia 1 0) 1 Sym phoricar pos 1 1 0
Zizyphus i! 0 1 Sambucus 1 0 1
Vitis 4 1 3 Viburnum 1s p 1
Tilia 3 1 2 Clerodendrum 1 0 1

Noteworthy in this flora is a so-called Madro-Tertiary element. In-
cluded in this category are:

Quercus (part), Juglans (Rhysocaryon), Mahonia, Cercocarpus,
Lyonothamnus, aff. Peraphyllum, Ceanothus, Colubrina, Karwinskia,
Garrya, X ylonagra, and Arbutus.

Note that most of these genera formed an important part in the up-
land forest of the late Miocene of Nevada; although they have been
termed ‘Madro-Tertiary” by some workers, i.e., the lineages supposedly
migrated northward into Nevada during the Neogene, the history of the
lineages indicate that they were derived from a mesic forest of the Pa-
cific Northwest during the middle to late Miocene interval.

The sources of the Mixed Mesophytic forest of the Miocene of west-
ern North America have been briefly touched on in this report (see also
discussion by Wolfe, 1969). One significant element that contains both
evergreen and deciduous broad-leaved plants was derived at various
times during the later Paleogene from the evergreen broad-leaved for-
ests. Several lineages representing genera such as Quercus (species that
have “unlobed” leaves), Alnus, Carya, Pterocarya, Magnolia, Cocculus,
Cinnamomophyllum, Persea, and other genera of the “Laurophyllum”
type; Liquidambar ; and Platanus, can be traced from the Paratropical
Rain or Subtropical forests of the Eocene and earlier Oligocene into
the temperate vegetation of the Miocene. A second and major element

96 MADRONO [Vol. 20

that primarily contains deciduous broad-leaved plants was derived dur-
ing the Eocene and earlier Oligocene from the temperate to marginally
subtropical vegetation that lived in the uplands, especially the Rocky
Mountains. Lineages that display such a distribution represent, for
example, Populus, Salix, Sassafras, Amelanchier, Cercocar pus, Crataegus,
Sorbus, and Acer. A third but minor element was derived from the tem-
perate vegetation of the later Oligocene of Alaska; most of these line-
ages represent Salicaceae or Betulaceae. Many of the lineages from these
various sources diversified after entering the Mixed Mesophytic forest;
members of Salicaceae, Judlandaceae, and Rosaceae, exemplify such a
pattern.

VEGETATION. As in the early Miocene, the vegetation of northwestern
North America was largely broad-leaved deciduous forest. Some zona-
tion of vegetation is evident because broad-leaved evergreens, except for
Ericaceae, were absent from Alaska but formed a significant element in
the Pacific Northwest. Upland vegetation in the Northwest had only a
minor element in Pinaceae, but in Alaska conifers of this family were
dominant in the uplands.

The middle Miocene was warmer than the early Miocene. This is
indicated by comparing the leaf margin percentages of the upland Colla-
wash assemblage, 25, with those of the upland Fish Creek, 31, and
Latah, 32, assemblages. Although the percentages for the Collawash and
Latah are not strikingly different, the large size of both assemblages
indicates that statistically the percentages are highly reliable. The
warming could have brought subtropical vegetation farther north along
the coast, and indeed the Cape Blanco assemblage has a leaf margin
percentage of 35, which is closely approaching subtropical.

In reference to probable altitudes of the assemblages from central and
southwestern Nevada, note that the leaf margin percentage for the San
Antonio, Middlegate, and Fingerrock assemblages combined is 19. This
is considerably less than the percentages for upland Oregon assemblages
such as the Mascall or Fish Creek. Broad-leaved evergreens such as
Exbucklandia, Magnolia, and Lauraceae are lacking in Nevada. I think
it highly probable that the Nevada assemblages must have been signifi-
cantly higher than the known middle Miocene assemblages from the
Pacific Northwest, some of which grew at altitudes of at least 2,500 feet,
i.e., the Nevada assemblages probably lived at altitudes of 4,000 to
5,000 feet or more. This suggestion takes into account the fact that not
only do the Nevada assemblages appear to represent a cooler vegetation
than that of the Pacific Northwest, but also that the Nevada assem-
blages are farther south than those in the Northwest.

LATE MIOCENE

The term late Miocene as used in this report represents the later part
of the Barstovian and all the early Clarendonian mammalian ages. In

1969] WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 97

terms of available radiometric ages, this interval lasted from about 14
to about 10 million years ago. Assemblages for which independent ages
are available are, in Alaska, the Grubstake (Wahrhaftig and others,
1969), Pribilof Canyon (Hopkins, et al., in press), and Marble Point
(locs. P5182, P9990 of Burk, 1965). The largest assemblages, however,
are those from the Kenai Formation—the Chuitna River and those from
the type section of the Homerian Stage (Wolfe, 1966). In British Co-
lumbia, late Miocene assemblages includes the Skonun (Martin and
Rouse, 1966) and the Quesnel (Peil, unpublished data). In the Pacific
Northwest this interval is represented by the Mashel, Wilkes, Faraday,
Weyerhauser, Skunk Creek, Hidden Lake, and Brock Road assemblages
(Wolfe, unpublished data), and by many previously published assem-
blages: Lower Ellensburg (Smiley, 1963), Stinking Water (Chaney
and Axelrod, 1959), Blue Mountains (Chaney and Axelrod, 1959),
Trout Creek (Graham, 1965), Thorn Creek (Smith, 1941), Trapper
Creek (Axelrod, 1964), Hog Creek (Dorf, 1936), and Pit River (La-
Motte, 1936). Not included in this discussion is the small assemblage
from the Payette Formation. In Nevada, late Miocene assemblages in-
clude the Cloropagus, Fallon, and Aldrich Station (Axelrod, 1956),
Chalk Hills (Axelrod, 1962), and Stewart Spring (Wolfe, 1964). In
central California, only two late Miocene assemblages have been studied:
the Table Mountain (Condit, 1944) and the Neroly (Condit, 1938).

In all instances, the Alaskan late Miocene assemblages represent low-
land vegetation, as interpreted from the geologic data. Some were
coastal (Pribilof Canyon, Marble Point), some were slightly interior
(Chuitna River and type Homerian), and one was several hundred miles
in the interior (Grubstake). The Skonun assemblage from the Queen
Charlotte Islands was obtained from predominantly marine beds, and
thus can be considered coastal lowland. The original altitude of the
Quesnel assemblage is unknown, except that this area of British Colum-
bia probably had at least moderate elevation during the late Miocene.

The Pacific Northwest assemblages represent both lowland and up-
land. The Mashel, Wilkes, Faraday, and Weyerhauser assemblages come
from the Puget-Willametie lowland, and geologic evidence does not indi-
cate that during the late Miocene this area was at a different altitude
than today. During the late Miocene, the High Cascade Range had not
yet developed, the Western Cascades being the only significant upland
between the eastern and western Pacific Northwest. The fact that a few
thousand feet of middle to late Miocene basalt did not override the Cas-
cade Range in southern Washington is an indication that the Western
Cascade Range there had at least moderate elevation, 3,000 feet, at the
time of extrusion of the basalts. A chain of volcanoes formed the crest
of this range in the central and northern parts of the Cascades in Oregon
(Peck, et al., 1964, p. 31). Note that the Brock Road assemblage was
situated near the crest of the range, whereas the Skunk Creek and Hid-

98 MADRONO [Vol. 20

den Lake assemblages were east of the crest. All are now west of the
crest of the present Cascades.

The topographic setting of the assemblages from eastern Oregon and
Idaho cannot be precisely determined. The Hog Creek assemblage was
probably in a broad valley of the ancestral Snake River. All the other
assemblages, however, were in a broad sense upland. Geologic data are
likewise imprecise in interpreting the original altitude of the Nevada
assemblages, although they too were upland.

In California, the Table Mountain assemblage, which is now in the
foothills of the Sierra Nevada, probably was lower than the present
altitude, ca. 2,000 feet. The Neroly assemblage occurs in an intertongu-
ing maine-nonmarine section, and was thus coastal.

Frora. After the middle Miocene, many genera and species of the
Mixed Mesophytic forest became extinct in northwestern North Amer-
ica, and are not found even in the lowland regions west of the Cascade
Range; the Faraday, Weyerhauser, Wilkes, and Mashel assemblages
represent the flora of the Puget-Willamette lowland. Notably lacking in
these assemblages are genera such as Castanea, Schoepfia/Anacolosa,
Cercidiphyllum, Cocculus, Magnolia, Cinnamomophyllum, Laurophyl-
lum, Exbucklandia, and many others that were present even in the up-
land early and middle Miocene. Specific diversity within genera such as
Carya, Fagus, and Quercus was also less in the late than in the early to
middle Miocene.

Floristic provincialism was more pronounced during the late Miocene
than earlier in the Neogene. A similarity matrix was constructed (table
2) based on comparison of the specific composition of the megafossil
assemblages. Five groupings are apparent: Alaska, Columbia Plateau-
Cascade Range, Nevada, Puget-Willamette, and California. The first
three floristically intergrade. Alaska and Nevada had floras that were
more closely related than were the floras of Nevada and California or
Nevada and the Puget-Willamette area.

Late Miocene assemblages, particularly those in Alaska, display a
greater diversity of Salix and Ericaceae than earlier assemblages. From
Alaska south to the Columbia Plateau and Nevada conifers were better
represented than before. The ‘“‘Madro-Tertiary” elements were largely
restricted to Nevada.

The group that suffered most extinction is composed of lineages that
were derived from the paratropical and subtropical vegetation during
the later Neogene. The Mixed Mesophytic elements that were derived
from the upland assemblages of the Paleogene Rocky Mountains were
proportionately better represented in the late than in the early or middle
Miocene. Notable exceptions, however, are some lineages of Juglanda-
ceae, Betulaceae, Liquidambar, and Platanus, all of which were of para-
tropical or subtropical extraction. These long-ranging lineages were also —
widely distributed in the Mixed Mesophytic forest and thus probably —
had broader tolerances than many of their associates. |

1969 |

WOLFE: FLORISTIC AND VEGETATIONAL HISTORY

TABLE 2. SIMILARITY MATRIX OF SOME LATE MIOCENE MEGAFOSSIL ASSEMBLAGES

IN NORTHWESTERN NortTH AMERICA.

Coefficient of similarity (or association) is that of Dice and Sgrensen as given in
Sokal and Sneath (1963, p. 129):

Sp = 2nsx/(2Nsx + U)

where S, = coefficient of similarity, n;; = number of positive matches, and u =
number of negative matches.

Gruusteke

“Te
Wxe, pt beaten)

Vorwu }avser

el = 1,00.

|
lil <==

020 = 0629

0—20 = 0629

0.10 = 0.19

y
=
Minh

aoa |i
ae

b=)
() 3 Y
ae Poe 4 56
: i, er ii TEE:
ae = fu a ma Shay 2&8 as Cates ask Y PA

Chalk
Hills
Table
Mountain

The modern aspect of the late Miocene flora is clear. Not only were
many exant lineages represented, but in some instances the extant spe-
cies was represented. Note, however, that many late Miocene lineages
have not survived to the present day. One particularly significant ex-
ample is the lineage represented by aff. Cedus; this lineage is represented
by cone scales in which the seeds were born entirely on the cone scale
as in Cedrus but that had a long subtending bract as in Abies. At the
four localities at which these cone scales have been found, pollen samples
typically display an abundance of Cedrus-like pollen and presumably
the two organs represent the same genus.

100 MADRONO [Vol. 20

It is noteworthy that in the late Miocene some related lineages had
attained certain distribution patterns which have been largely main-
tained to this day, and some lineages had attained a restricted distribu-
tion either west or east of the Cascade Range. Thus the lineages of
Alnus incana, Betula papyrifera, and B. occidentalis are known only
from assemblages that lived to the east of the late Miocene crest of the
Cascade Range; the lineages to which A. oregona and A. rhombifolia
belong were, on the other hand, represented only west of the Cascades.
In reference to north-south distribution, the probable ancestor of Holo-
discus dumosus occurs in assemblages that were east and south of the
assemblages in which the probable ancestor of H. discolor is represented.
The lineage of Betula papyrifera was largely northern and the lineage
of B. occidentalis was largely southern. However, some lineages, e.g.,
that to which Quercus kelloggi belongs, were represented both east and
west of the Cascades, whereas these lineages are today entirely western.

VEGETATION. The leaf margin percentages for both the Alaska and
Nevada assemblages are approximately 30, whereas those for the Puget-
Willamette and Columbia Plateau assemblages range from 14 to 24,
typically about 17 to 21. The high percentage for Alaska, combined
with the small size of the leaves, is interpreted as representing an ap-
proach to cold, and hence physiologically arid, conditions; the present
percentage for Alaska is 42. In Nevada, however, a summer dry climate
had probably set in, which is of course physiologically arid (Wolfe and
Hopkins, 1967). Percentages similar to those for Alaska have been ob-
tained for the modern vegetation of the Pacific Northwest. The Puget-
Willamette assemblages were probably analogous to the vegetation of
the Middle Atlantic states, i.e., temperate broad-leaved deciduous forest.

The extensive amount of pollen data allows an excellent understand-
ing of vegetational regions during the late Miocene (fig. 2). The Alaskan
assemblages represent a cool conifer forest, which was floristically more
diverse than the present boreal forest. Broad-leaved deciduous plants,
other than Betulaceae and Salicaceae, were represented, although sparse-
ly. The Skonun assemblage shows some similarity to the Alaskan as-
semblages in the high representation of Betulaceae, but the moderate
representation of Juglandaceae and Fagaceae indicate a mixed conifer
broad-leaved deciduous forest. The Skonun-Mashel-Weyerhauser-Fara-
day spectra intergrade from the mixed forest in the northern lowlands to
a broad-leaved deciduous forest in the southern lowlands. Juglandaceae,
Fagaceae, and Liquidambar appear to have been the major constituents
of the more southern forest. This could be designated a hickory-oak-
beech assemblage.

The vegetational differences between the Willamette lowland and
the higher elevations in the Cascade Range can be readily determined
from the pollen spectra (fig. 3). All upland spectra have large amounts
of Pinaceae, particularly aff. Cedrus and Picea, whereas, except for

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 101

sis Bara

feet aff. Cedrus

:
|

i755
|

|
'

‘i

Pseudotsuga

Ill ines

Se Fa ad Sy /

Ye ee Wi ee a
Jf es Taxodiaceac/ Cupressaceae:

os ; oN x 7 Wa 2:1: Broad+leaved exotics
| \ }

YP cg, Ae Aivos_and Betuls

ae e | Ht, eS ay i = a | i
ae | iii eee ell | |
: ' [ i

Fic. 2. Pollen spectra of some late Miocene assemblages in northwestern North
America. The following assemblages are included (number preceding corresponds to
figure 1; numbers in parentheses indicate number of samples and total grains
counted): 25. Blue Mountains (2:600); 33. Hidden Lake (7:2112); 37. Mashel
(1:300) ; 39. Pit River (1:300) ; 40. Pribilof Canyon (2:600) ; 41. Quesnel (1:430) ;
42. Skonun (10:1575); 44. Stewart Spring (1:300; 47. Thorn Creek (1:300); 48.
Trapper Creek (1:300); 50. Type Homerian (6:1027); 51. Weyerhauser (3:964) ;
Data for Quesnel furnished by K. M. Piel and data for Skonun from Martin and
Rouse (1966). Some conifers are too sparsely represented in some spectra to be in-
cluded in this illustration. Most of the solid black segment in Mashel spectrum rep-
resents Keteleerza. In the Pribilof Canyon spectrum Cy = Cyperaceae and Grami-
neae and in the Type Homerian spectrum Er = Ericales.

[ Vol. 20

MADRONO

102

OooT

4eguep mb17 :

sn2s2ney nn.

SS SS el ta ee eee!

SD)!wW OT S i)

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 103

Pinus, Pinaceae are a minor element in the lowalnd spectra. Broad-
leaved deciduous trees, notably members of Juglandaceae, Fagaceae,
and Liquidambar, are dominant in the Weyerhauser spectrum and rare
in the upland spectra. The Faraday spectrum is in some respects
transitional between the Weyerhauser and the upland spectra. This
transitional character could be expected because of the proximity to
the site of deposition of the Faraday beds of a volcano that was active
during the late Miocene (Peck, et al., 1964). The pollen profile for the
Faraday beds displays a marked decrease in pollen of Pinaceae immedi-
ately above an ash layer; presumably the eruption of the ash from the
local volcano resulted in the death of many Pinaceae that grew on the
slopes of the volcano. Note that Tsuga, primarily T. heterophylla type,
attains its highest relative abundance in the Faraday spectrum; Tsuga
may have been a prominent member of the forest at altitudes inter-
mediate between the Faraday and Hidden Lake sites of deposition.

The upland conifer forest, which can be characterized as a spruce-
cedar forest, occupied a broad area from central British Columbia south
to northeastern California and east to the mountains of Idaho. The
Trapper Creek assemblages appears to be intermediate between the
spruce-cedar association and the Nevada vegetation, as indicated in
the Trapper Creek by the paucity of aff. Cedrus and the presence of
Ephedra, Sarcobatus, and Artemesia. Axelrod (1964, p. 65) has sug-
gested that the presence of Ephedra and Sarcobatus reported earlier
from Trapper Creek (Leopold, 72 Mapel and Hail, 1959) was due to
modern contamination. An additional sample, with which great care was
taken to eliminate and prevent contamination, has also yielded these
pollen types. There is no reasonable basis for the exclusion of these
genera from the Trapper Creek flora.

The megafossil assemblages from Nevada have been characterized as
representing a spruce-live oak-cedar (Chamaecyparis, not aff. Cedrus)
association (Wolfe, 1964). The pollen assemblages from Stewart Spring
that are now under study by H. E. Schorn confirms such an interpretation.

Judging both mega- and microfossil data, the following associations
appear to have occupied northwestern North America during the late
Miocene:

Alaska. Conifer forest of Abies grandis, Picea sitchensis, P. glauca,
Pinus monticola, and Tsuga heterophylla, accompanied by Betula papy-
rifera, i.e., a birch-pine forest. Streamside vegetation largely Salicaceae
and Alnus. Undergrowth largely Rosaceae and Ericaceae.

Fic. 3. Pollen spectra of some late Miocene assemblages in the Willamette Valley
and Cascade Range of Oregon. Profile of Western Cascade Range is hypothetical but
based on available geologic data (Peck, et al., 1965; D. L. Peck, pers. comm.). As-
semblages are, from left to right (numbers in parentheses indicate number of samples
and total grains counted): Weyerhauser (3:964), Faraday 28:7757), Skunk Creek

(1:300) a,ynd Hidden Lake (7:2112). Small amounts of Keteleeria are included in
the Abies totals.

104 MADRONO [ Vol. 20

Columbia Plateau-Cascade Range. Conifer forest ofAbzes concolor,
Abies magnifica, aff. Cedrus, Keteleeria, Picea breweriana, P. magna,
Pinus monticola, P. ponderosa, Tsuga heterophylla, Thuja plicata, and
Sequoia sempervirens, accompanied by Betula papyrifera, Quercus
chrysole pis, Ulmus spp., Acer spp., and Arbutus idahoensis, i.e., a spruce-
cedar forest. Streamside vegetation largely Salicaceae, Alnus, and Plata-
nus. Undergrowth mostly Rosaceae.

Nevada. Conifer forest of Abies concolor, Picea breweriana, Picea
magna, Chamaecyparis, nootkatensis, Tsuga heterophylla, accompanied
by Quercus chrysolepsis, i.e., a spruce-cedar-live oak forest. Streamside
vegetation largely Salicaceae. Undergrowth mostly Rosaceae.

Puget-Willamette lowland. Broad-leaved deciduous forest of Carya
bendirei, Fagus, Quercus deflexiloba, Liquidambar, and Arbutus, ac-
companied by Thuja plicata and Sequoia sempervirens, i.e., a hickory-
oak-beech forest. Streamside vegetation largely Salicaceae and Betula-
ceae. Undergrowth mostly Rosaceae and Ericaceae. Locally, a Taxodium-
Nyssa association persisted.

Central California—mixed broad-leaved evergreen and deciduous wood-
land of Quercus chrysolepis, Castanopsis, Carya, Persea, and Arbutus,
accompanied by Pinus (closed-cone), i.e., a live oak-madrone woodland.
Streamside vegetation mostly Salicaceae. Undergrowth mostly Rosaceae
and Ericaceae. Locally, a Taxodium-N vssa association persisted.

PLIOCENE AND EARLY PLEISTOCENE

From about 10 to 2 or 3 million years ago, the fossil record in north-
western North America is notably poor. Pliocene assemblages are known
in Alaska: the Type Clamgulchian, represented by many large collections
from the Kenai Formation, and several assemblages from the Bering plat-
form area (Hopkins, et al., 1960; Wolfe, unpublished data) that are
probably of late Pliocene and/or early Pleistocene age. In the Pacific
Northwest, Pliocene assemblages are represented by the Troutdale
(Chaney, 1944b), Dalles (Chaney, 1944a), Middle Ellensburg (Smiley,
1963), and Deschutes (Chaney, 1938). One early Pleistocene assem-
blage is known from the Cape Blanco area (Wolfe unpublished data).
Pliocene assemblages from regions south of the Pacific Northwest appear
to have little to-contribute to an understanding of the vegetation or flora
of the Northwest.

The topographic setting of all the known Alaskan Pliocene and early
Pleistocene assemblages is coastal lowland. The Troutdale assemblage
in Oregon is the only Pliocene assemblage known from the Puget-Wil-
lamette lowland. The assemblages from eastern Washington and Oregon
probably were at moderate elevations. The assemblage from the Elk
River beds at Cape Blanco was obtained from rocks that are predomi-
nately marine.

FLorA. Compared to the Miocene, the Pliocene flora was depauperate.
In Alaska, nearly all the broad-leaved deciduous trees had become ex-

1]
1]
|

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 105

tinct; note, however, that Glyptostrobus, which today is the most tropical
member of Taxodiaceae, persisted into the Type Clamgulchian assem-
blage. Salicaceae and two species in the Betulaceae were the primary
constituents of the Alaskan ligneous dicotyledonous flora, and Pinaceae
are well represented by Picea and Pinus in the microfossil assemblages.

In the Northwest, the flora still contained some broad-leaved decidu-
ous elements now exotic to the region, such as Pterocarya, Ulmus, Plata-
nus, and Aesculus. Some of these exotics, e.g., Ulmus, are known in the
Deschutes assemblage (Wolfe, unpublished data), which is only five to
six million years old (Evernden and James, 1964). By the early Pleisto-
cene, however, broad-leaved exotics were apparently almost extinct in
the Northwest. The Elk River assemblage contains some Platanus, but
otherwise the flora has a modern aspect.

The known megafossil species in the Northwestern Pliocene and
Pleistocene assemblages are primarily members of lineages that were
present in this region during the Miocene. No significant amount of mi-
gration of woody lineages into the region appears to have taken place
during the Pliocene, although the fossil record is sufficiently poor to
emphasize “‘appears.”” As noted in the discussion of the late Miocene
flora, the flora of the Pacific Northwest was modern in aspect by the
late Miocene.

VEGETATION. The character of the vegetation in the Northwest during
the Pliocene is not clear, largely because of the lack of pollen assemblages
and the small size of the megafossil assemblages. The leaf margin per-
centages for both the Troutdale and Middle Ellensburg show a definite
increase compared to the Faraday and Lower Ellensburg. This change
was not, therefore, the result of orogenic activity, because the Faraday-
Troutdale sequence is west of the Cascade Range. The increase in leaf
margin percentages is prcbably due to the onset of a definite summer dry
climate in the Pacific Northwest (Wolfe and Hopkins, 1967). This
change would account for the extinction of many lineages, e.g., those of
Carya, Fagus, Liquidambar, and some of Acer. The persistence of Ulmus
east of the Cascade Range should not be surprising, because some Asian
species of the genus have proved to be highly successful in cultivation on
the Columbia Plateau.

The Troutdale assemblage does not indicate a cooler climate than that
of the late Miocene, despite the probable change to summer dry condi-
tions. The extinction of various lineages by the end of the late Miocene
can be readily explained in terms of the changed precipitation regime.
Insofar as known, the Troutdale assemblage could represent broad-
leaved deciduous forest derived from the late Miocene vegetation. None
of the Troutdale localities have yielded records of Pinaceae more diverse
than the Pinaceae of, for example, the Weyerhauser assemblage.

The Ellensburg assemblages (Smiley, 1963) may offer a clue as to
the direction of temperature changes. The late Miocene Lower Ellensburg

106 MADRONO [Vol. 20

assemblage lacks Lauraceae except for Sassafras, whereas the early Plio-
cene Upper Ellensburg assemblage contains Persea. Similarly, the Middle
Ellensburg assemblage contains ligneous legumes lacking in the Lower
Ellensburg. Although the evidence is not conclusive, the early Pliocene
may have been somewhat warmer than the late Miocene in the Pacific
Northwest.

The Deschutes assemblage indicates that by the later Pliocene, conifer
forest was probably not present near the sites of deposition; the late
Miocene rocks, however, consistently contain some megafossils of Pina-
ceae. This absence of Pinaceae in the Pliocene megafossil assemblages
may reflect the increasing aridity from the rain shadow created by the
accretion of the High Cascades. The conifer forest would thus be re-
stricted to higher elevations away from the sites of deposition. Clearly,
however, much more paleobotanical, especially palynologic, data are
needed for an understanding of the vegetation of the Columbia Plateau
during the Pliocene.

ORIGINS OF THE MODERN FLORA AND VEGETATION

From the preceding discussion, it is clear that many phylads and line-
ages now extant in the Pacific Northwest were present in this region
prior to the Pleistocene and were in fact present in the Miocene. Several
of the lineages were at one time members of the summer wet Mixed Meso-
phytic forest and have since adapted to summer dry conditions. Extant
species that have such a history in the Pacific Northwest include: Popu-
lus tremuloides, P. trichocarpa, Salix commutata, S. fluviatilis, S. lasi-
andra, S. lasiolepis, Quercus garryana, Q. kelloggi, Alnus rhombifolia,
A. rubra, Betula occidentalis, Castanopsis chrysophylla, Celtis douglassi,
Clematis columbiana, Mahonta aquifolium, M. repens, M. nervosa, Ame-
lanchier alnifolia, Cercocarpus montanus, Crataegus douglassi, Holodis-
cus discolor, Osmaronia cerasiformis, Peraphyllum ramoisissimum, Prunus
demissa, Sorbus scopulina, Spiraea, densiflora, Toxicodendron radicans,
Acer grandidentatum, A. glabrum, A. macrophyllum, A. negundo, Ceano-
thus velutinus, Arbutus menziesi, Leucothoe davisae, Rhododendron
occidentalis, Cornus nuttalli, Garrya elliptica, Shepherdia utilis, and
Sambucus glauca.

Some of the extant species, however, appear to have entered the Pacific
Northwest after the extinction of the Mixed Mesophytic forest, i.e., after
the middle Miocene. Some appear to have entered from the north, be-
cause their first fossil records are in the Miocene or Pliocene of Alaska;
note that this does not mean that the lineages are of northern origin but
may have evolved in Eurasia and migrated through Alaska. Included are:
Salix barclayi, S. scouleriana, S. hookeriana, S. glauca, S. monticola, S.
pipert, Myrica californica, Alnus, incana, A. sinuata, Ribes triste, Prunus
subcordata, Rubus idaeus, Acer circinnatum, Gaultheria shallon, Vac-
cinium alaskaense, and Symphoricar pos albus.

The majority of these species live today in the mountains of the

1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 107

Pacific Northwest, although some, particularly Salix hookeriana and its
allies, are lowland species. These willows apparently attained a coastal
distribution from Cook Inlet south to western Washington during the
later Miocene. Some lineages may have entered the Pacific Northwest
from the south. Peraphyllum, for example, was a member of the Mixed
Mesophytic forest in the Northwest, but the only late Miocene record of
the genus is in Nevada; when summer dry climates prevailed in the
Northwest at the close of the Miocene, lineages such as Peraphyllum may
have reentered from the south. Most ‘“‘Madro-Tertiary” elements in the
modern flora of the Northwest may have followed such a pattern.

The known Tertiary vegetation of the Pacific Northwest does not
closely match the modern vegetation floristically. The deciduous broad-
leaved forest of the late Miocene western lowlands has survived in a
highly modified form in the Willamette and Rogue River valleys. At
least 50 percent of the late Miocene species of this vegetation, however,
no longer participate in this forest type. Most have become regionally
extinct, but others, for example, Leucothoe aff. L. davisae, are today
montane species.

The late Miocene conifer forest of the Columbia Plateau-Cascade
region has also been highly modified since 10 million years ago. The
area perhaps floristically most closely related to this vegetation is the
present Siskiyou Mountains and adjacent areas. In this region, most
of the late Miocene conifer species have survived, including some of the
associated ligneous dicotyledons. Again, however, at least 50 percent
of the late Miocene species associated with the conifer forest no longer
participate in this extant forest type. Although extinction accounts for
much of the floristic change, some of the lineages, e.g., those to which
Quercus chrysolepis sensu stricto (excluding Q. vaccinifolia) and Plan-
tanus racemosa belong, are no longer part of the conifer forest. The
primary difference, however, between the late Miocene and modern
conifer forest of the Northwest is that Pseudotsuga did not play as sig-
nificant a role in the vegetation as it does today. Hansen’s (1949) data
indicate that Pseudotsuga pollen in the pollen rain occurs in about the
same percentage as the percentage of Pseudotsuga in the surrounding
forest as measured in basal area. Pollen of Pseudoteuga is, however,
not abundant in any Neogene pollen assemblage known to me in the
Northwest. Even the early Pleistocene assemblages at Cape Blanco
have low amounts of Pseudotsuga (Wolfe, unpublished data), whereas
interglacial deposits of the Puget lowland (Leopold, unpublished data)
contain large quantities of the genus. It appears, therefore, that the
dominance of Pseudotsuga in the present conifer forest of the Northwest
was attained during the middle or late Pleistocene.

Note also that, since the late Miocene, the conifer forest has received
some immigrants from the north as well as incorporating some elements
from the late Miocene broad-leaved forest. To the latter group probably

108 MADRONO [Vol. 20

belong: Arbutus menziesi, Leucothoe davisae, and Rhododendron macro-
phyllum.

The conifer forest of the late Miocene of Alaska has, of course, suf-
fered from extensive extinction. Vegetationally, this conifer forest is
probably most like that of southern British Columbia, and there are also
strong floristic similarities. The apparent absence of Acer and Rhodo-
dendron (Rhododendron) coupled with the presence of Pterocarya and
Ulmus in the Alaskan late Miocene, however, indicate that as a vegeta-
tional and flloristic unit the Alaskan late Miocene forest did not migrate
southward.

The floristic changes that have occurred in the vegetation of the
Pacific Northwest during the Neogene indicate strongly that the present
associations are also most probably transitory. As Mason (1947) has
noted, the association must be continually redefined at each point in
time because of the coincidental nature of associations.

U.S. Geological Survey, Menlo Park, California

LITERATURE CITED

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Publ. Geol’ Scie 33:

. 1962. A Pliocene Sequoiadendron forest from western Nevada. Univ.
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BarLey, I. W., and E. W. Sinnott. 1915. A botanical index of Cretaceous and Ter-
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Berry, E. W. 1931. A Miocene flora from Grand Coulee, Washington. U.S. Geol.
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Brown, R. W. 1940. New species and changes of name in some American fossil
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Burk, C. A. 1965. Geology of the Alaska Peninsula—Island arc and continental
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Cain, S. A. 1944. Foundations of plant geography. Harper, New York.

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. 1924. Quantitative studies of the Bridge Creek flora. Amer. J. Sci. 8:127-

144.
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476:187-216.
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interpretation. Publ. Carnegie Inst. Wash. 617:1-134.
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1969 | WOLFE: FLORISTIC AND VEGETATIONAL HISTORY 109

CuecesBaEva, A. I. 1968. The Neogene flora of the River Pirozhnikovaya in Kam-
chatka. [In Russian.] Bot. Zurn. (Moscow & Leiningrad) 53:737-748.

Conpit, C. 1938. The San Pablo flora of west-central California. Publ. Carnegie Inst.
Wash. 476:217-268.

. 1944. The Table Mountain flora. Publ. Carnegie Inst. Wash. 553:57-90.

Davis, M. B., and J. C. GoopretrT. 1960. Comparison of the present vegetation with
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41:346-357.

Dorr, E. 1936. A late Tertiary flora from southwestern Idaho. Publ. Carnegie Inst.
Wash. 476:73-124.

EVERNDEN, J. F., and G. T. JAMEs. 1964. Potassium-argon dates and the Tertiary
floras of North America. Amer. J. Sci. 262:945-974.

.. D. E. Savace, G. H. Curtis, and G. T. JAMEs. 1964. Potassium-argon
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262:145-198.

Fecri, K., and J. IversEN. 1964. Textbook of pollen analysis. Hafner, New York.

GranHaM, A. 1965. The Sucker Creek and Trout Creek Miocene floras of southeastern
Oregon. Kent State Univ. Bull. 53.

Gray, J., and L. R. Kitrteman. 1967. Geochronometry of the Columbia River
basal and associated flora of eastern Washington and western Idaho. Amer. J.
Sci. 265:257-291.

Hansen, H. P. 1949. Pollen content of moss polsters in relation to forest compo-
sition. Amer. Midl. Naturalist 42:473-479.

Hopkins, D. M., F. S. MAcNEIL, and E. B. Leoporp. 1960. The coastal plain at
Nome, Alaska: a late Cenozoic type section for the Bering Strait region. Rep.
21st Int. Geol. Congr. pt. 4:46-57.

——., D. W. ScuHo it, W. O. Appicott, R. L. Pierce, P. J. Smiru, J. A. WOLFE,
D. GrersHANOVICH, B. Kotronev, K. E. LoHMAN, and J. OBrapovicn. In press.
Cretaceous, Tertiary, and early Pleistocene rocks from the continental margin
in the Bering Sea. Bull. Geol. Soc. Amer.

KNOWLTON, F. H. 1902. Fossil flora of the John Day basin, Oregon. U.S. Geol. Surv.
Bull. 204.

LaMortTe, R. S. 1936. The Upper Cedarville flora of northwestern Nevada and
adjacent California. Publ. Carnegie Inst. Wash. 455:57-142.

MacGinitte, H. D. 1953. Fossil plants of the Florissant beds, Colorado. Publ. Car-
negie Inst. Wash. 599.

Mason, H. L. 1947. Evolution of certain floristic associations in western North
America. Ecol. Monogr. 17:201-—210.

Maret, W. J., and W. J. Har. 1959. Tertiary geology of the Goose Creek district,
Cassia County, Idaho, Box Elder County, Utah, and Elko County, Nevada.
U.S. Geol. Surv. Bull. 1055-H.

Martin, H. A., and G. E. Rouse. 1966. Palynology of late Tertiary sediments from
the Queen Charlotte Islands, British Columbia. Canad. J. Bot. 44:171-208.

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Prof. Paper 140-A: 1-16.

Peck, D. L., A. B. Griccs, H. G. ScuHiickEr, F. G. WELLs, and H. M. Dore. 1964.
Geology of the central and northern parts of the Western Cascade Range in
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Ricwarps, P. W. 1952. The Tropical Rain Forest. Cambridge Univ. Press.

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35:159-276.

SmiTH, H. V. 1941. A Miocene flora from Thorn Creek, Idaho. Amer. Mid]. Natural-
Ist 25:473-522.

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110 MADRONO [Vol. 20

Tuayver, T. P., and C. E. Brown. 1966. Geologic map of the Aldrich Mountain
quadrangle, Grant County, Oregon. U.S. Geol. Surv. Map GQ-438.

TurNeER, D. L. 1968. Potassium argon dates concerning the Tertiary foraminiferal
time scale and San Andreas fault displacement. Ph.D. thesis. Univ. Calif.,
Berkeley.

WaHrRuHArFTIG, C., J. A. WoLre, E. B. LEopotp, and M. A. LANPHERE. 1969. The coal-
bearing group in the Nenana coal field, Alaska. U.S. Geol. Surv. Bull. 1274-D.

Wanc, Cui-Wvu. 1961. The forests of China. Pub. Maria Moors Cabot Found. Bot.
Rés. 5.

Wotre, J. A. Tertiary Juglandaceae of western North America. M.A. thesis. Univ.
Calif., Berkeley.

. 1960. Early Miocene floras of northwest Oregon. Ph.D. thesis. Univ. Calif.,

Berkeley.

. 1962. A Miocene pollen sequence from the Cascade Range of northern

Oregon. U.S. Geol. Surv. Prof. Paper 450-C:C81-C84.

. 1964. Miocene floras from Fingerrock Wash, southwestern Nevada. US.

Geol. Surv. Prof. Paper 454-N.

. 1966. Tertiary plants from the Cook Inlet region, Alaska. U.S. Geol. Surv.
Prof. Paper 398-B.

. 1968. Paleogene biostratigraphy of nonmarine rocks in King County,

Washington. U.S. Geol. Surv. Prof. Paper 571.

. 1969. Paleogene floras from the Gulf of Alaska region. U.S. Geol. Surv.
open-file report.

—., and D. M. Hopkins. 1967. Climatic changes recorded by Tertiary land

floras in northwestern North America. Jn K. Hatai, Tertiary correlation and

climatic changes in the Pacific. Sasaki, Sendai, Japan.

.. D. M. Hopxtns, and E. B. LEoporp. 1966. Tertiary stratigraphy and bio-
stratigraphy of the Cook Inlet region, Alaska. U.S. Geol. Surv. Prof. Paper
398-A.

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northwestern North America and northeastern Asia. In D. M. Hopkins, ed., The

Bering Land Bridge. Stanford Univ. Press, Stanford.

ECOLOGIC PLANT GEOGRAPHY OF THE
PACIFIC NORTHWEST

R. DAUBEN MIRE

The purpose of this phytogeographic sketch is to provide an introduc-
tion to the natural vegetation of the Pacific Northwest for the use of
botanists from other areas who will be attending the XI International
Botanical Congress in Seattle in 1969. Attention will be centered on the
State of Washington, with secondary emphasis on the adjoining areas.
If my friends in British Columbia feel that I have slighted their Prov-
ince, this is more a consequence of maps terminating at the international
border than of any intent of mine to confine attention to the ‘Pacific
Northwest,”—a nationalistic and ambiguous though useful term, which
I shall not try to define!

The visitor, like some of us who reside here, may be appalled by the
scarcity of natural vegetation in a region which was opened up by white
explorers as late as 1805 (the Lewis and Clark expedition). Neverthe-
less the account is centered on remnants of natural vegetation with the
intent of helping the visitor recognize some of the common types and
see how they fit into a regional pattern.

THE LAND SURFACE

The Rocky Mountains (only a limited fringe of them shows at the
right edge and upper half of fig. 1) are the oldest of our major moun-
tain systems, the uplifting and folding that first gave them definition
having taken place well back in the Cenozoic Era. Their rocks are partly
granitic and partly sedimentary, and show a wide variation in the de-
gree of metamorphism, but neither chemically nor physically do they
present extreme conditions that h ave been acclaimed of much phyto-
geographic importance.

Later in the Cenozoic another major geologic event of interest to us
here was the building up of a great basaltic plateau of some 26,000,000
ha? in extent that covered the non-mountainous parts of eastern Wash-
ington, eastern Oregon, and southern Idaho. Layers of the columnar
basalt that comprise this plateau are conspicuous features of canyon
walls throughout the plateau region.

In Pliocene time the Cascades were uplifted, these mountains reaching
considerable height and thereby stimulating so much rainfall that their
basaltic veneer was in large part soon eroded away exposing the intrusive
core. Since basalt gives rise to much more fertile soil than acid igneous
rocks, there are some significant botanical differences to be observed in
going from basaltic to granitic areas. For example, from Mt. Rainier
southward basalt is the prevailing rock still mantling the Cascades,
whereas northward acid igneous rocks outcrop. The ubiquity of Purshia

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114 MADRONO [Vol. 20

tridentata in the igneous foothills, in contrast with its restriction to a few
special habitats in the basalt area to the south is correlaed with this geo-
logic discontinuity.

In Fig. 1 a chain of low mountains is shown following the Pacific
shoreline. These are the Coast Ranges. They are well developed in south-
western Oregon where they merge with the Cascades, but otherwise they
are quite low in Oregon and Washington with the outstanding exception
of the Olympic Mountains. To the north in British Columbia they again
become imposing features of the land.

Continental glaciation terminated in Washington and northern Idaho
(fig. 1), leaving the area to the north mantled with glacial till. In places
melt waters flowing outward from the ice cut great chasms in the basalt
flows, and elsewhere washed the surface clean in broad drinageways, de-
positing large sheets of gravel and sand farther downstream.

Vulcanism during the late Cenozoic raised a string of spectacular vol-
canic peaks far above the general level of the Cascade crest from British
Columbia to California, but spectacular as these peaks are, their major
significance for ecology lies in the ejecta spread over the landscape when
two of them erupted.

In approximately 12,000 B.c. Glacier Peak, situated a little to the
northwest of the center of Washington in Fig. 1, erupted and winds at
that time spread fine white ash over considerable area to its leeward.
Then in 6600 B.c. the stupendous explosion of Mt. Mazama, in the south-
west quarter of Oregon in Fig. 1, gave rise to a blanket of materials so
deep and extensive as to dwarf the Glacier Peak event. (Crater Lake
now occupies the vent where Mt. Mazama exploded.) Fine white ash
fanned out eastward and northward over most of the Pacific Northwest,
extending well into British Columbia and Montana, to provide a valuable
marker in sediments that has been generally useful in geochronology,
archaeology and the correlation of fossil sequences in bogs. In the more
immediate vicinity of Mt. Mazama coarse pumice was deposited to
great depths, smothering pre-existing vegetation and providing a sub-
strate that supports highly distinctive vegetation even today. For ex-
ample, Pinus contorta, nearly everywhere a seral tree in the mountains,
is the only tree that can grow over a large portion of the pumice area,
and here it maintains reproduction as a climax dominant.

In some period prior to the last (Wisconsin) glaciation there were
several episodes of loess accumulation on the basaltic plateau between

Fic. 1. Physiography of Washington and Oregon, with adjoining parts of British
Columbia, Idaho, Nevada and California. These state and province names have been
indicated only by centrally placed letters, respectively: W, O, BC, I, N and C. The
crnamented line crossing the map in the approximate latitude of central Washington
indicates the southern limits of the last (Wisconsin) continental glaciation. Areas
subjected to mountain glaciation are not indicated. Map reproduced through the
courtesy of Edwin Raisz and the publishers of W. W. Atwood’s “Physiographic
provinces of North America.”

1969 | DAUBENMIRE: PLANT ECOLOGY nals)

the Cascades and the Rockies. The volcanic ash which fell later became
incorporated with this loess, or in places was washed into hollows. On
the forest-covered mountain slopes the ash was trapped and fixed where
it fell, giving the forest of northern Idaho in particular a relatively com-
plete cover of fine-textured fertile material (usually very fine sandy
loam) so that the occurrence of thin stony soil is not so important a
factor in forest ecology there as in most mountain areas of western North
America. Down on the basal plain in Washington, the loess mantle with
its incorporated ash was in places stripped away in the path of extensive
Wisconsin glacial floods, so that desert-like areas of bare basalt became
exposed. These areas which support sparse vegetation mainly of dwarf
shrubs and the tiny caespitose grass Poa secunda, are labelled ‘‘chan-
nelled scablands”’ in Fig. 1.

Other botanically significant geologic features which are more localized
are the outcroppings of serpentine (southwestern Oregon and the east
slope of the Cascades in central Washington), recent outpourings of lava
(locally in the Cascades of Washington and Oregon, and especially in
Craters of the Moon National Monument in southern Idaho), and cones
of coarse ejecta that dot the landscape in central Oregon, and form Mt.
St. Helens in Washington.

CLIMATE

Far greater climatic complexity is encountered in the surroudings of
these meetings than has characterized the areas where most of the pre-
ceding congresses have convened. All of those areas had diversity result-
ing from the latitudinal climatic gradient, and some had a significant
superimposed gradient, usually extending more or less at right angles,
that involved degrees of oceanicity or continentality. But it would be
difficult to find a match for the intensification of the north-south cli-
matic gradient that results from the strong influence of the Westerlies
that are centered approximately on the Canada-United States border, or
a match for the steepness of the oceanic-continental gradient that results
from massive mountains rising to height of several thousand meters
within a few km of tidewater.

When the topographic map (fig. 1) is compared with the precipitation
map (fig. 2), the “approach effect” is clearly shown by the high precip-
itation on lowlands as the Westerlies approach major mountain masses
from the southwest. A sizable area in the lowlands to the southwest of
the Olympic Mountains has precipitation in excess of 2,540 mm a year.
Quinault Ranger Station has an average annual value of 3,414 mm. It is
in this general area that cryptogamic epiphytes are so spectacularly de-
veloped, and a trip up the Hoh or Quinault Valleys is eminently worth-
while to see the phenomenon. The approach effect is again evident along
the west base of the Cascades and even along the west base of the
Rockies across the arid intermountain trough.

The effect of mountains on precipitation also has its negative aspect,

116 MADRONO [Vol. 20

the creation of a relatively dry rain shadow along the leeward base of
each of the major mountain masses. To the northeast of the Olympic
Mountains a weak but well-defined rain shadow provides many more
sunny days at places like Sequim than in surrounding areas, with the
influence clearly discernible even in Seattle and on the nearby islands
in Puget Sound. The most pronounced rainshadow is along the eastern
base of the Cascades in central Washington. In the Yakima Valley the
mean annual precipitation drops as low as 175 mm within easy sight of
heavy forests of mesophytic conifers that encircle the glacier-clad slopes
of Mount Rainier where precipitation is at least 2,972 mm. Both the
rainshadow and the approach effect are well illustrated (from British
Columbia to Oregon) by both weather statistics and the resultant veg-
etation patterns.

Not only does our precipitation vary greatly from one place to another
but its unequal distribution throughout the year is botanically important.
Each summer the belt of the Westerlies recedes northward, and failing
to drag heavy supplies of moisture inland at this season, the summers
are left remarkably short of precipitation. Where the annual sum is low,
the shortage becomes critical, and those who visit the rainshadow at the
east flank of the Cascades will experience weather that for a few months
in summer remind one of true desert conditions. Only the coolness and
shortness of summers west of the Cascades (the spring and autumn sea-
sons are quite long) keep drouth from reaching serious proportions there.

Winter rains are gentle and almost mistlike, often accompanied by
heavy cloud and fog that bedevil air transportation. Coastal areas get
only half the possible hours of sunshine. In summer convectional showers
are so rare that storm-induced erosion is not severe. Strangers seeing the
wheatlands of southeastern Washington (the “Palouse” in fig. 1) for the
first time are usually amazed at the sight of cultivated slopes so steep
that self-levelling harvesting machines are used, and the trucks that ser-
vice them frequently overturn. Such slopes are not cultivatable under
rainfall conditions that prevail toward the center of the continent.

Along the coast the frost-free season is approximately 250 days long,
and snow is rather ephemeral, whereas in the mountains frost is likely any
day in summer and snowbanks may persist throughout the summer even
in the subalpine forest belt.

In winter a high pressure system forces Arctic air masses southward
down the plain east of the Rockies and often this air spills over the
mountains westward to temporarily overwhelm the oceanic character of
the climate between the Rockies and the Cascades. These sudden cold
spells are very damaging to fruit trees and ornamentals (all aliens), and
sometimes even the native plants are damaged. The Columbia Gorge is
large enough to allow oceanic influences to penetrate inland readily, and
often cold air influences extend farthest west down this gorge. At its
western extremity many trees are flag-shaped in consequence of ice
storms in winter when cold air pours westward down the valley, and at

1969 | DAUBENMIRE: PLANT ECOLOGY 117

the eastern extremity they are flag-shaped in the opposite direction by
the pressure of winds blowing eastward while branches are actively
growing.
SOILS

In discussing ‘‘the land surface” the general character of the parent
materials for soils has been indicated. In Fig. 3 the types of profiles that
might be expected on deep loams of undulating topography are shown.
These zonal soils are closely related to the pattern of rainfall, but since
there are large areas of steep topography in which surface materials are
moved by gravity from time to time, the large areas of parent material
of geologically recent origin (sand and gravel, bare rock, volcanic ejecta,
desiccated lake beds, etc.), the map units are highly complex mixtures of
azonal, intrazonal, and zonal soils.

ALPINE VEGETATION

Where the mountains rise so high that heat is no longer adequate to
allow the development of the tree life-form, the vegetation of the tree-
less area is called alpine tundra (figs. 4, 5). Near the coast the elevation
of this ecologically critical level is about 900 m in west central British
Columbia, rising to about 2,700 m at the southern end of the Cascade
Mountains in California. Eastward in the more continental climates of
the Rocky Mountains the critical level is about 800 m higher at equiva-
lent latitudes. Although this timberline is relatively stable, one can fre-
quently see evidence of trees having invaded contiguous areas of herba-
ceous vegetation, with most of the invasion having taken place during
the very dry period in the early 1930’s.

In the highest part of the alpine region glaciers and lichen-covered
rock debris dominate the landscapes, whereas down next to timberline
lush meadows dominated by forbs (Lupinus, Castilleja, Valeriana, Pedic-
ularis, Anemone) mixed with graminoids, or by low shrubs (Empetrum
nigrum, Luetkea pectinata, Phyllodoce, Cassiope) prevail. At interme-
diate elevations there are stony fell fields where the plant cover is in-
complete and cushion plants exemplified by the holarctic Silene acaulis
are best developed. Mid- to late July is approximately the height of the
flowering season for alpine plants in west central North America. The
communities they form have been well studied only in Montana.

SUBALPINE FORESTS

The forests just below upper timberline differ considerably on either
side of a line drawn a little east of the divide of the Cascades (figs. 4, 5).
East of this line Abies lasiocarpa and Picea engelmannii are the conspic-
uous species in old-growth stands. Since they are geologically recent de-
rivatives of Abies balsamea and Picea glauca, which characterize the
transcontinental taiga belt, the term montana taiga has been applied to
subalpine forests of the Rockies. Climatic similarity, i.e., heat budgets
marginal for tree development, is another point of similarity that is syne-
cologically significant.

118

MADRONO

[Vol. 20

1969 | DAUBENMIRE: PLANT ECOLOGY 119

Undergrowth plants in this forest likewise bespeak strong affinity with
the far North, e.g., Cornus canadensis, Linnaea borealis, Pyrola ssp., and
Sorbus scopulina. In the vicinity of the Canada-United States border
Menziesia ferruginea and Xerophyllum tenax usually dominate over all
other undergrowth species. Elsewhere a widespread type has the dwarf
Vaccinium scoparium as the chief ground cover.

Following fire or logging the first generation forests are typically
composed of Pinus contorta, P. albicaulis, P. monticola, and Larix occi-
dentalis in varying proportions. Abies lasiocarpa and Picea engelmanni
sometimes invade simultaneously but usually they are late-comers.

Nearly everywhere the A dies slowly increases at the expense of all others,
ultimately dominating the climax, except in the few localities where
Tsuga mertensiana occurs. There the T'suga proves the best competitor.
In the krummholz at upper timberline, where the severe climate keeps
the tree cover so open that differences in shade-tolerance are not critical,
Abies lasiocarpa and Pinus albicaulis or P. flevilis are the main species.

Subalpine forests of the Cascade crest and the few high mountains
still closer to the ocean might be bracketed under the term montane taiga
only by virtue of their heat budget, since floristic affinities with the
transcontinental taiga are essentially nil. Although Abies lastocarpa
makes a limited penetration into the strongly oceanic climates, reaching
even to the Olympic Mountains, Picea engelmanni does not follow it,
and instead the major climax dominants there are Abies amabilis, A.
magnifica, Tsuga mertensiana, and Chamaecyparis nootkatensis. Beneath
these trees the common plants include Menziesia ferruginea, Rhododen-
dron albiflorum, Xerophyllum tenax, Vaccinium ovalifolium, V. mem-
branaceum, Pachistima myrsinites, and Pyrola spp.

Conspicuous trees in the initial regeneration that follows deforestation
are Pseudotsuga menziesii, Larix occidentalis, Abies procera, A. lasio-
carpa, Pinus contorta, P. monticola, and P. albicaulis. The last- men-
tioned becomes especially prominent with Tsuga mertensiana in the
krummbholz at upper timberline.

MONTANE AND LOWLAND FORESTS CHARACTERISTIC
OF WET OCEANIC CLIMATES

In the region about Seattle natural forest succession usually trends
toward the dominance of Tsuga heterophylla on well-drained mineral
soil. Acer circinata and Cornus nuttallii are common understory trees.
Beneath these trees the two most significant dominants, both evergreen,
are Polystichum munitum on relatively moist sites or Gaultheria shallon
on soils having a slight tendency toward dryness. Other common species,
most conspicuous under intermediate moisture conditions, are Berberis
nervosa, Clintonia uniflora, Cornus canadensis, Phegopteris dryopteris,
Ribes sanguineum, and Rubus spectabilis.

Fic. 2. Mean annual precipitation in inches. Areas with more than 2,540 mm and
less than 254 mm have been shaded for emphasis (after Kincer, 1922).

120 MADRONO [Vol. 20

GREAT SOIL GROUPS, MAINLY:

- SIEROZEM

- BROWN

- CHESTNUT

- CHERNOZEM & PRAIRIE

- NON-CALCIC BROWN

- BROWN PODZOLIC, GRAY BROWN
PODZOLIC, GRAY WOODED, WESTERN
BROWN FOREST & PODZOL

- REDDISH BROWN LATERITIC
ALPINE TURF

- GRUMUSOL

SOLONCHAK & SOLONETZ
ALLUVIAL & HUMIC GLEY

- REGOSOL

- LITHOSOL

1969 | DAUBENMIRE: PLANT ECOLOGY 121

In somewhat swampy situtions Thuja plicata maintains dominance in-
definitely, and the conspicuous undergrowth plants include Oplopanax
horridum, Athyrium filix-foemina, and Lysichiton, americanum. In val-
leys too dry for Tsuga or Thuja, Abies grandis is the climax dominant.

The vegetation as a whole has a lush aspect, with many of the broad-
leaved trees, shrubs, and herbs evergreen, as are nearly all the coniferous
trees.

Owing to the high incidence of fires in the past, the uplands are most
usually dominated by coastal ecotypes of Psuedotsuga menziesu, which
is the most valuable forest tree of the region and the mainstay of its im-
portant forest industry. Old virgin stands of this tree, already established
when white man first arrived but no longer common outside national
parks, contained individuals up to 127 m tall, with butt diameters up to
7.6 m. Alnus rubra is another seral species that is highly aggressive on
disturbed land. This is the largest species in its genus and is extensively
used for furniture. On recently cut-over lands Pteridium aquilinum or the
alien Cytisus scoparius commonly determine the physiognomy until the
first seral forest gets established. In southwestern Oregon Ulex europea
plays this role.

On moist but not swampy terraces Acer macrophyllum is quite com-
mon, this tree being an especially good “host” for Selaginella oregana
which so copiously festoons its branches on the windward side of the
Olympic Mountains (especially in the Hoh and Quinault Valleys).
Populus trichocarpa, Rhamnus purshiana, and Fraxinus oregana are
other broad-leaved trees common on stream terraces.

Picea sitchensis is mainly confined to a very narrow strip along the
open ocean where it plays a climax role under moderate salt spray influ-
ence, but becomes seral to 7'suga on sandy soils slightly farther back
from the ocean. Other trees that are found only close to the ocean are a
salt-tolerant ecotype of Pinus contorta that is most common on dunes
or bogs, and Chamaecyparis lawsoniana which is seral to Tsuga hetero-
phylla in the Coast Ranges of southern Oregon. Forest along the coast
is frequently interrupted by dunes supporting such characteristic species
as Elymus mollis, Carex macrocephala, Lupinus littoralis, Abronia lati-
folia, Poa macrantha, and Lathyrus littoralis, or by tidewater marshes
in which Distichlis spicata and Salicornia pacifica are the dominants.

Coniferous forests quite similar to those described as characteristic of
uplands and swamps back from the influence of sand and salt spray con-
tinue from near sea level up the mountains to the lower limits of the
subalpine belt, and from British Columbia to southern Oregon, with a
disjunct area reappearing on the seaward slope of the major mountain
ranges in northern Idaho, northern Montana and southeastern British

Fic. 3. Principal soil regions. Only the characteristic zonal soil present in each map
unit has been indicated (after Various Authors, 1964).

122 MADRONO [Vol. 20

Columbia (‘‘cool moist forests” in figs. 4 and 5). In this inland exten-
sion the most notable differences are (1) the substitution of Pinus mon-
ticola and Larix occidentalis as the prevailing seral trees, and (2) the
absence of Gaultheria shallon and scarcity of Polystichum munitum in
the undergrowth. In place of these one finds mainly Clintonia uniflora,
Tiarella unifoliata, Phegopteris dryopteris, Viola orbiculata, and Vac-
cinium membranaceum, all of which occur in the coastal segment, but
are overshadowed there by the more conspicuous Gaultheria, Poly-
stichum, Berberis nervosa, etc.

The major climax dominants of upland forests in the interior are seg-
regated, with Tsuga heterophyla in the most moist situations. Thuja
plicata in less moist places (but a seral tree on Tsuga sites), and Abies
grandis on soils tending to be drouthy.

Immediately after a fire destroys forests in this group a rich variety
of shrubs invades the landscape, and until a new forest canopy develops,
this shrubbery provides an abundance of winter browse for deer and
wapiti which abound in the area.

Along the coast a related but evidently distinct forest area is one in
which Sequoia sempervirens dominates, this area extending as a narrow
strip from the southwest extremity of Oregon to a little south of San
Francisco Bay (see its delimitation in fig. 4). One living Sequoia meas-
sured at 117 m tall is thought to be the tallest tree left standing in North
America. Pure stands of this species characterize floodplains but on con-
tiguous hillsides Lithocarpus densiflora, Abies grandis, and Pseudotsuga
menziesi form a lower tree stratum. The most abundant plants of the for-
est floor include Oxalis rubra, Polystichum munitum, Berberis nervosa,
Gaultheria shallon, and Vaccinium spp. Thuja and Tsuga heterophylla
are sparingly represented in the Sequoza area.

The southern and inland limits of the Sequoia sempervirens forest are
associated with the limits of frequent summer fog which results from
warm breezes blowing landward across very cold water along the coast.

FORESTS CHARACTERISTIC OF MODERATELY DRY TO DRY CLIMATES

Soils of the subalpine forests remain moist throughout summer, ex-
cept perhaps in the top 20 cm or so, and the same is true of those forests
below in which Tsuga, Thuja or Abies grandis are climax dominants. But
in passing from the Abies grandis areas into land where the soils regu-
larly dry several to many decimeters deep there is a clear ecologic dis-
continuity. Here Pseudotsuga menziesii (or ecologic equivalents) remains
free from competition from more shade-tolerant species and so can per-
sist as a major climax dominant, if not the only one. It is desirable to
consider separately three geographic subdivisions of this Pseudotsuga belt.

In the Willamette Valley of central Oregon (fig. 4) the Douglas fir
forest is distinctive for the abundance of Quercus garryana, a low decid-
uous tree which is usually seral to Pseudotsuga, but may form pure

1969 | DAUBENMIRE: PLANT ECOLOGY 123

climax stands in certain relatively dry sites. Small areas in which the
Quercus plays much the same role also occur on the gravelly outwash
plain in Washington southward of Olympia and Tacoma. The deciduous
Holodiscus discolor is a characteristic shrub of the Douglas fir forest.

Another distinctive segment of what is here referred to as the Pseudo-
tsuga belt occurs in the mountains of south central Oregon, extending
from there southward in an inverted V-shaped area (delimited in fig. 4)
on both sides of the interior valley of California. In this area Pseudotsuga
is variously associated with Libocedrus decurrens and Abies concolor.
Pinus ponderosa, P. jeffreyi, P. lambertiana, and Sequoiadendron gigan-
teum are all seral trees here. The last mentioned is famous for its mas-
sive trung, which grows to a diameter of 11 m and a height of 100 m,
thus dwarfing the climax trees of shorter stature and life-span that grow
around it. Seqguoiadendron trunks may have more than 3500 xylem layers.

Another distinctive feature of this southern Oregon-California sector
of Douglas fir belt is the abundance of evergreen sclerophyllous shrubs
in the undergrowth. These increase conspicuously in early stage of re
generation cycles following deforestation.

A third subdivision of the Douglas fir belt has more distinctly conti-
nental climates, occurring on the east-facing slopes of the Cascades,
thence eastward in the foothills across the breadth of the Rockies. Here
Pseudotsuga is the sole climax dominant on zonal soils, with seral trees
including Pinus ponderosa, P. contorta, Larix occidentalis, and Populus
tremuloides. In the undergrowth one of the following usually dominates
the physiognomy: Calamagrostis rubescens, Physocarpus malvaceus,
Holodiscus discolor or Sym phoricar pos albus. On the east flank of the Cas-
cades, Pinus contorta is the only conifer that can form forests on large
areas of coarse pumice in what appears to be the equivalent climatic belt.

Progressing down the moisture gradient into areas drier than Pseudo-
tsuga can tolerate, one sometimes comes abruptly onto steppe on the
basal plain, but elsewhere one or two distinctive forest belts may inter-
vene.

If the Douglas fir belt gives way to another forest belt, the later is
typically dominated by Pinus ponderosa growing in pure stands. These
climax pine stands contain a wide variety of undergrowth types depend-
ing on variation in climate, aspect or soil, all of which still allow the pine,
and only this pine, to form the tree stratum. Some of the undergrowth
plants that are locally conspicuous are Purshia tridentata, Physocar pus
malvaceus, Symphoricarpos albus, Festuca idahoensis, F. scabrella, Ag-
ropyron spicatum, Stipa comata, and Aristida longiseta. In certain of
these forest types the pine grows slowly and is subject to attack and de-
formation by the parasitic Arceuthobium campylopodum, but elsewhere
it grows rapidly and provides one of our most valuable timbers. The
best growth of the tree, however, is in higher zones where Pseudotsuga
and Abies grandis are climax.

124 MADRONO

[ Vol. 20

Te
it

eS )
2 Gi) Gam yr he eres
SEL |S a acer “a
EC | A i KR
ot A Ve

SN SS

\ AS eee) eae
(2 Gaaeaney Nestieta oe
— c Nace ce

\ 5

ALPINE VEGETATION & SNOWFIELDS

COLD SUBALPINE FORESTS OF STRONGLY
OCEANIC CLIMATES

COLD SUBALPINE FORESTS OF QUASI-OCEANIC
OR CONTINENTAL CLIMATES

COOL MOIST FORESTS WITH TSUGA HETERO-
PHYLLA, THUJA OR SEQUOIA

DRY FORESTS WITH PSEUDOTSUGA, PINUS
PONDEROSA, PINUS CEMBROIDES OR JUNIPERUS
STEPPE

CHAPARRAL

HALOPHYTIC VEGETATION

EIGN) ee

Fic. 4. Vegetation of northwestern U.S.A. (after Shantz & Zon, 1924).

1969] DAUBENMIRE: PLANT ECOLOGY 125

P| ALPINE VEGETATION & SNOWFIELDS

COLD SUBALPINE FORESTS OF STRONGLY
OCEANIC CLIMATES

COLD SUBALPINE FORESTS OF QUASI-OCEANIC
OR CONTINENTAL CLIMATES

SUBARCTIC (PICEA GLAUCA) FOREST

COOL MOIST FORESTS WITH TSUGA HETERO-
PHYLLA OR THUJA

DRY FORESTS WITH PSEUDOTSUGA OR

PINUS PONDEROSA

et Cnc © fete) ie) (04 ial eee: eprenel eh one
jes

> Gap. Sef! 18 or ISVisy Crea ei Le Bel lei Neel eile

spies te eile! eter 3, 10

ef e* ae

Sonewies Te
eT yee’ Nal "cr ter eglan otis

tn,
[ "h,

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wh yas
Se

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See,

Fic. 5. Vegetation of British Columbia (after Rowe, 1959).

126 MADRONO [Vol. 20

Again as one reaches the lower and drier limit of the Pinus ponderosa
belt, he may come out onto an unforested basal plain, or he may have
one more forest belt to pass through. The last is perhaps better charac-
terized as woodland savanna, for the species of Juniperus (J. scopulorum,
J. occidentalis, J. osteospermum) and Pinus flexilis which comprise this
type are only a few meters tall and rarely form a closed canopy. This
lowest conifer belt is fairly well developed in western Montana, and
more locally represented in southern Idaho, and in central Oregon (where
the pine does not occur).

On the west side of the Rockies in Washington and British Columbia,
groves of Populus tremuloides become a conspicuous feature of land-
scapes in the margin of the steppe, where they are confined to locally
moist places.

CHAPARRAL VEGETATION

Chaparral vegetation in North America (corresponding to maquis,
garigue, fynbos and mallee in other parts of the world) is rather easily
divided into three physiognomic categories: scrub (up to 2 m tall),
woodland and savanna. All three types are well represented about the
margin of the great valley in central California (fig. 4), and the first two
extend in fragmentary fashion into southern Oregon.

Chaparral scrub in which Ceanothus cuneatus, Arctostaphylos viscida,
A. canescens, Cercocarpus betuloides, Eriodictyon californicum, Garrya
fremonti, Rhamnus californica, and Rhus triloba are major species is lo-
cally well represented east of the Coast Ranges in the upper Rogue River
Valley of Oregon, with a trace still farther north in the Umpqua River
Valley. This scrub occupies a belt marginal to what was originally a
small area of steppe, (all too small to show in fig. 4), with Quercus gar-
ryana, Pinus ponderosa and Pseudotsuga menziesi in progressively wet-
ter situations.

Relatively dry ridge summits in southwestern Oregon are in places
dominated by Lithocarpus densiflora and Arbutus menziesii growing as
low forest trees with Rhododendron macrophyllum forming a tall shrub
layer beneath. Stands of this character may be interpreted as the north-
erly limits of chaparral woodland, which is floristically much more com-
plex southward.

STEPPE VEGETATION

The low-lying unforested areas between the Cascades and the Rockies
are often referred to as “desert” by the local populace. But the few
months of intensive summer heat and drouth, combined with the oc-
currence of cacti, rattlesnakes, scorpions, and tarantulas that perhaps
suggest this classification to the layman, do not impress the plant geog-
rapher as strongly as does the moderately heavy to heavy stand of per-
ennial grass that grew nearly everywhere before the era of domestic live-
stock. The term steppe is more appropriate to the botanist.

1969] DAUBENMIRE: PLANT ECOLOGY 127

A good grass cover is possible on practically all soils throughout the
arid parts of the Pacific Northwest, for despite the low rainfall, nearly
all of it falls in winter when evapotranspiration is feeble and much of the
water can percolate deep enough to get below the reach of direct evapo-
ration into the air, thus providing water to sustain a vigorous spurt of
vegetative and flowering activity as temperature rises in spring. Summer
is mostly a period of aestivation with more plants inactive at this season
than in mid-winter. East of the divide of the Rockies most of the rain
falls in summer, and the steppe grasses there delay their activity
accordingly.

On zonal soils in the steppes of the Pacific Northwest the most char-
acteristic dominants are Agropyron spictaum, Festuca idahoensis, F.
scabrella (chiefly in British Columbia), and Poa secunda. On sandy soils
these species are replaced by Stipa comata, Oryzopsis hymenoides,
Agropyron dasystachyum, and Sporobolus cry ptandrus.

The driest parts of the intermountain trough from Central British
Columbia to central Oregon, together with practically all the intermoun-
tain area farther south, originally supported conspicuous but low-grow-
ing gray-leaved shrubs that formed a layer well above the grasses, making
the term shrub-steppe applicable to this area. Over most of the shrub-
steppe Artemisia tridentata was the outstanding representative of this
shrub layer, with the monotony somewhat relieved locally by other
shrubs such as Chrysothamnus nauseosus, C. viscidiflorus, Tetradymia
canescens, Gravia spinosa, Purshia tridentata, and Gutierrezia sarothrae.
All these shrubs are deep-rooted, and except for Grayia they retain their
leaves through the dry summers, with Artemisia holding its leaves
throughout the year. Herbaceous plants, for the most part the same
species as in contiguous steppe lacking shrubs, go into aestivation in
midsummer as moisture in the upper meter or more of the profile is
used up.

Perennial forbs (Arnica, Balsamorhiza, Castilleja, Crepis, Geranium,
Lupinus, Potentilla, etc.) are conspicuous in both steppe and shrub-
steppe in inverse ratio to the degree of aridity, i.e., they achieve their
maximum importance in communities just below lower timberline.

Three cacti occur in the Pacific Northwest—Opuntia polyvacantha
(most widespread), O. fragilis, and Pediocactus simpsonit.

Over most of the intermountain steppe area the most aggressive plant
to assume dominance as grazing eliminates native perennials is the alien
Bromus tectorum. In the less arid eastern fringe of the steppe it is lo-
cally moist enough for Poa pratensis to take over this role. In the Great
Basin area the noxious alien Halogeton glomeratus is relatively more
conspicuous in this role.

In eastern Washington and northern Oregon thin stony soils overlying
basalt outcroppings are abundant and these support a wide variety of
special communities in nearly all of which the tiny perennial Poa secunda
is the chief grass, with one or more species of dwarf shrubs conspicuous.

128 MADRONO [Vol. 20

Artemisia rigida is the most widespread of these dwarf shrubs, but sev-
eral species of Eviogonum are also common.

Other special soil types in the steppe region support communities in
which one of the following is the characteristic dominant: Eurotia lanata,
Artemisia arbuscula, A. nova, and Atriplex nuttallit.

Saline basins in the Great Basin of Utah and Nevada support a rich
assortment of halophytic shrubs, forbs, and grasses. The major halo-
phytes here, roughly in order in increasing tolerance of wetness and
salinity, are: Salicornia rubra, Allenrolfea occidentalis, Suaeda depressa,
S. intermedia, Distichlis stricta, Atriplex patula, A. argentea, Sporobolus
airoides, Puccinellia airoides, Triglochin maritima, Bassia hyssopifolia,
Sarcobatus vermiculatus, Atriplex confertifolia, A. nuttallii, and Kochia
americana. Northward only a few of this group play the role of major
dominants. Here in the north Distichlis stricta is ubiquitous. In some
places it is associated with Elymus cinereus, a coarse caespitose grass
growing about 2 m tall, and elsewhere with Sarcobatus vermiculatus, a
succulent-leaved chenopodiaceous shrub. Marshes in which Typha lati-
folia and Scirpus spp. cover large areas are conspicuous in south central
Oregon.

Department of Botany, Washington State University, Pullman

GENERAL REFERENCES

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FRANKLIN, J. F., and N. E. WEst. 1965. Plant communities of Oregon: A bibliog-
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FRYXELL, R. 1965. Mazama and Glacier Peak volcanic ash layers: relative ages.

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Haseck, J. R., and E. Hartrey. 1965. The vegetation of Montana—A bibliography.
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Hansen, H. P. 1947. Postglacial forest succession, climate, and chronology in the
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SOIL DIVERSITY AND THE DISTRIBUTION OF PLANTS,
WITH EXAMPLES FROM WESTERN NORTH AMERICA

A. R. KRUCKEBERG

Discontinuity of pattern and form is an ubiquitous feature of living
things. All along the scale of organizational complexity, from cell to eco-
system, some degree of environmental hiatus separates the elements of a
system. Mitochondria are discreet entities separated from one another
by the microenvironments of dissimilar subcellular phases. At a higher
level, individuals of the same population are not confluent; time, space,
chance and inhospitable habitats contrive to maintain temporary or par-
tial discontinuities. But it is at the level of the species that isolation is
most apparent and evolutionarily significant. The modern Darwinist sees
the immense diversity in the organic world as resulting from the inter-
actions over geological time of the variant heredities of organisms, the
natural selection of adapted variants, and the manifold factors which
promote the isolation of those adapted variants. Evolution of our present
diverse biota without discontinuity is unthinkable.

The discontinuities in the physical environment which isolate popula-
tions and species, though richly unlimited in degree and permuting inter-
action, can be reduced to but two broad groups: climatological and geo-
logical. In other words, the limits to distribution of kinds of organisms
are largely imposed by differences in climate or geology, or both. That
climate is primary in controlling the broad distribution of organisms is
undisputed. Tolerance spans of terrestrial organisms are chiefly limita-
tions in reaction to stressful levels of moisture and temperature. Within
areas of broad climatic similarity, though, geological variability provides
the major source of regional biotic diversity. The contribution of geolog-
ical phenomena to vegetational discontinuity takes a number of forms.
Variations in topography, in mineral content and physical properties of
parent rock account for most of the regional differentiation brought about
by geological processes. When microclimatic and biotic features act in
concert with geology, the mosaic of habitats is greatly enriched.

The soils derived from parent rocks owe their distinct qualities to a
set of interacting factors. Hans Jenny (1941) characterizes the soil for-
mation process as a set of variables in a functional array. In Jenny’s
formula, s = f (cl, 0, r, p, t), there are five independent variables that
define the soil system: climate (cl), organisms (0), topography (r), rock
type or parent material (p), and time (t). If all but one factor, say p
(parent material), remain constant, then variations in the end product
(soil) are due to differences in parent material.

It is this one variable, parent material (p), that will be the central
theme of this review of soil diversity and plant distribution. The Jenny

129

130 MADRONO [ Vol. 20

formulation, however, reminds us that the biological properties of soils
cannot be analyzed one factor at a time. When we abstract just parent
material from the total soil ecosystem, we achieve manageable simplicity
only at the expense of setting aside the interacting whole system. Our
primary concern will be to examine the possible effects of chemical vari-
ations of parent material and soil on plant distribution. Such chemical
diversity will condition the kind and amount of mineral nutrients avail-
able to the vegetation. To paraphrase Jenny’s factorial approach, we can
ask: Other factors being equal, what effects do differences in soil mineral
content have on the distribution of plants?

Soils can control the distribution of plants in other ways, however,
and we will look briefly at some of these. The physical properties of soil
and parent material, weathering processes, soil microclimate, and those
properties uniquely induced by the biota are also the domain of the
edaphic factor complex; they will be touched upon wherever appropriate.

Though we may be operating at a microcosmal level where other en-
vironmental influences might vastly overshadow the effects of soil chem-
ical differences, such differences can be dramatically effective in altering
plant distribution. Examples to follow will support this contention. I
will offer evidence to support two general hypotheses that bear on the
probable interactions between mineralogical composition of parent mate-
rial and discontinuities of plant distribution.

First Hypothesis: Given a regional climatic framework, much of the
plant species diversity and discontinuity in the region is governed by
variations in soil chemistry, and thus by specific variations in the min-
eralogy of rock substrates.

Second Hypothesis: Speciation within a regionally contiguous genus is
largely a response to environmental discontinuity within the confluent
area. Sharp discontinuities in soil chemistry can serve as isolating phe-
nomena to bring about species diversification.

HISTORICAL BACKGROUND

We can scarcely doubt that discerning humans through the ages have
been aware of sharp vegetational discontinuities arising from differences
in soil. Geological and vegetational diversity go hand in hand in regions
around the Mediterranean and the near east, the scene of Man’s agricul-
tural beginnings. A much later record from the Age of New World Ex-
ploration specifically ties vegetation to soil. Columbus is said to have
capitalized on a specific soil-plant association when he had to replace a
mast on a ship of his first fleet (Buck, 1949). The story goes that he was
counselled to choose a log o f pine growing on red soil in nearby Cuba;
the red limonitic soils of Cuba are known to be high in iron and to have
furnished durable timbers.

Correlations between substrate and vegetation really became a part of
botanical science much later—in the 19th cenutry. It was the young and
alert Austrian botanist, Franz Unger, who first emphasized the signif-

1969 | KRUCKEBERG: SOIL DIVERSITY 131

icance of geological formation for plant distribution. From the pen of the
master botanist-naturalist, Anton Kerner von Marilaun, we get an in-
triguing accunt of the patterning of vegetation that set Unger to develop
his concept of the chemical concept of plant distribution. I quote from
the English version of Kerner’s Natural History of Plants, (Kerner and
Oliver, 1902): “‘The little town of Kitzbuhel, in the Northeast Tyrol, has
a very remarkable position. On the north rises the Wilde or Vorder
Kaiser, a limestone chain of mountains with steep, pale, furrowed sides,
and on the south the Rettenstein group, a chain of dark slate mountains
whose slopes are clothed far up with a green covering. The contrast pre-
sented by the landscape in its main features is also to be seen in the vege-
tation of these two mountain chains. On the limestone may be seen
patches of turf composed of low stiff Sedges, Saxifrages whose formal
rosettes and cushions overgrow the ledges and steps of the rugged lime-
stone, the yellow-flowered Auricula, the Rock-rose-flowered Rhododen-
dron, and white-flowered Cinquefoil adorning the gullies, dark groups of
Mountain Pines bordered with bushes of Alpine Rose; and opposed to
these on the slate mountains are carpets of thick turf composed of the
Mat-grasses sprinkled with Bell-flowers, Arnica montana and other Com-
posites, groups of Alpine Alder and bushes of the rust-colored Alpine
Rose—these are the contrasts in the plant-covering which would strike
even a cursory observer, and would lead a naturalist to ask what could
have been the cause. No wonder that the enthusiastic Botanist, Franz
Unger, was fascinated by this remarkable phenomenon in the vegetable
world. In his thirtieth year, furnished with a comprehensive scientific
training, he came as a doctor to Kitzbuhel, and with youthful ardour he
used every hour of leisure from his professional duties in the investiga-
tion of the geological, climatic and botanical conditions of his new lo-
cality, devoting his fullest attention to the relations between the plants
and the rocks forming their substratum. The result of his study was his
work, published in 1836, on the Influence of Soil on the Distribution of
Plants as shown in the Vegetation of the North-east, Tyrol, which
marked an epoch in questions of this sort. The terminology introduced
in the book found rapid entrance into the botanical works of the time.
Unger divided the plants of the district accordingly to their occurrence on
one or other of the substratums—in which lime and silica respectively
predominated—into (1) those which grow and flourish on limestone
only; (2) those which prefer limestone, but which will grow on other
soils; (3) those which grow and flourish on silica only; and (4) those
which, whilst preferring silica, will grow on other soils.”’

Until the advent of modern soil science, arguments pro and con for
Franz Unger’s chemical theory of plant-soil relationships persisted with-
out the full understanding of the nature of plant mineral nutrition. The
essence of Unger’s view—that mineral content of soils is the primary
edaphic influence on plant distribution—is vindicated by contemporary
soil chemistry. Qualitative and quantitative differences in elemental

132 MADRONO [Vol. 20

(ionic) content of both the exchange complex and the soil solution
do cause selective responses in the composition of vegetation cover as I
will relate shortly.

Parent material of whatever sort, igneous, metamorphic, sedimentary
rocks, and organic materials, become soil by weathering. Through the
action of temperature changes, wind abrasion, water and other chemical
agents, as well as biological influences, rocks weather to those textural
and particle size classes of materials that constitute mineral soil. Mineral
soil, then, is a mixture of particles ranging downward in size from rock
fragments through gravel and sand to silt and clay. The most reactive
phase for plants is the colloidal clay fraction. Ionic exchange between
root systems and the soil is mediated by clay colloids. Major and minor
elements required for plant growth are adsorbed on clay colloid surfaces.
Since weathering frees primary minerals to generate secondary clay min-
erals and to participate in ion exchange, it is to be expected that the
mineralogical composition of the weathering parent material will deter-
mine the quality of the reactive mineral content of soils.

The diversity of the geologic parent materials available at the earth’s
surface for soil formation is vast and rich. The range of rock types is
derived from variations in both mineralogical content and mode of origin.
Thus, at one end of the spectrum are the acid rocks, rich in feldspars
(silicates of K, Na, Al and Ca). Acid rocks exist as granites (batholithic
or intrusive igneous), rhyolites (volcanic surface flows), or as schists and
gneisses (metamorphics), or as consolidated sediments (sandstones,
etc.). Omitting the broad range of transitional members along the scale
from acid to basic rocks, we come to the other end of the spectrum. Here
are the ultrabasic rocks, chiefly of iron-magnesium silicates, plentiful and
worldwide in distribution. Both igneous and metamorphic types occur;
common examples of ultrabasics are peridotite and serpentinite.

PLANT RESPONSES TO DIFFERENCES IN CHEMICAL COMPOSITION OF
SoIL AND UNDERLYING PARENT MATERIAL

How sensitive is the plant to variations in chemical content of soils?
For cultivated plants, man’s agricultural experience is rich and his skill-
ful manipulations of crops and soils have had bountiful returns. Applica-
tion of macro- and micronutrient fertilizers is a cornerstone of good farm
practice. But what of natural vegetation and its response to chemical
variations in soil? Positive evidence is clear for peculiar vegetational and
floristic displays on a number of truly abnormal soils. The term “abnor-
mal” signifies 1, the abundant occurrence in soils of one or more elements
rarely found in such excessive amounts in agricultural soils, or 2, the
absence of one or more of the essential plant nutrients usually available
in cultivated soils, or yet 3, some combination of these exceptional ele-
mental constitutions. However, I would repeat my earlier ‘“disclaimer”’.
Soil as a part of the living ecosystem is the product of many inter-
actions, both biotic and environmental. To say that a soil is abnormal

1969 | KRUCKEBERG: SOIL DIVERSITY 133

and gives rise to exceptional vegetational responses due to mineral com-
position is valid to the extent that of all the soil forming influences, the
quality of parent material is primary in its effect on plant growth.

I will present examples largely from the “abnormal” group of soils.
Not only are the vegetational responses so striking, but their careful
study may reveal guidelines for determining the chemical effects that
may exist for plants on more normal, yet chemically variable, soils.

‘““ABNORMAL” (AZONAL) SOILS

Soils and Vegetation of Limestone and Dolomite

It was the stark contrasts in vegetation between limestone and slate
slopes in the Tyrolean Alps that led Franz Unger (1836) to his chemical
theory of edaphic restriction. Striking differences in physiognomy, species
composition, and plant morphology are associated with rocks rich in cal-
cium carbonate. Examples of contrasts between calciphile and calciphobe
elements of a regional flora abound in the early European literature.
Plants favoring limestone soils are calciphiles or calcicoles ; plants avoid-
ing limestone soils are calciphobes or calcifuges. The occurrence of vicari-
ism (selective replacement of closely related species or varieties on con-
trasting soils) is frequent. It will suffice to give one or two examples of
contrasts in floristic composition to reveal the nature of the vegetational
discontinuity caused by limestone.

The remarkable flora on vast outcrops of limestone and chalk in Great
Britain has fascinated botanists and naturalists for decades. This sus-
tained interest is delightfully recounted in two modern books, “Wild
Flowers of the Chalk and Limestone,” by J. E. Louseley (1950), and
“Downs and Dunes, Their Plant Life and its Environment,” by E. J.
Salisbury (1952). Only a fragmentary account of the rich chalk and lime
floras can be given here. The gamut in degree of constancy of species to
soils derived from rocks rich in calcium carbonate begins on the side of
the rare and obligate lime inhabitants such as the two orchids, Orchis
simia, monkey orchid, and O. muilitaris, military orchid, and Helianthe-
mum polifolium, white rockrose. Franz Unger (1836, p. 168) would have
called such exacting plants “‘bodenstet” (or “‘soil-fast”’). The other ex-
treme, plants common not only on limestone, but on other soils, he re-
ferred to as “bodenvag”’ (or “‘soil-wanderer”’) species. In addition, lime-
stones and chalks in Britain have their share of calcifuges—plants that
avoid the calcareous substrates. Foxglove, Digitalis purpurea, and broom,
Sarothamnus scoparius, though widespread, are conspicuously absent
from these soils. Louseley says of the two species, “. . . (they) are such
excellent soil indicators that on train journeys it is often easy to tell im-
mediately when the railroad line leaves chalk or limestone by their pres-
ence on the railway banks.” Ericaceous species, long known to gardeners
for their aversion to limestone, are equally discriminating members of
the natural vegetation. Only where the chalks are surface-leached and an

134 MADRONO [Vol. 20

acid humus has developed can species of the heath family get a local toe-
hold on lime.

Limestone vegetation in other parts of the world is no less remarkable
for possessing a high proportion of indicator species. For example, the
geology of Japan and Taiwan is especially rich in calcareous deposits. The
botanical composition of 63 limestone outcrops (Shimizu, 1962), were
categorized in grades of fidelity, the degree to which a species is re-
stricted to a particular community type. There are 75 species in Fidelity
Class 5 (‘‘exclusives”—high restriction to limestone) ; this class contains
a large number of ferns, shrubs, and herbaceous perennials, but few trees.
The next Fidelity classes, 4 and 3, (‘‘selectives” and “preferents’”’) with
48 and 112 species respectively, add considerably to the total floristic
richness of the calcicolous floras.

There is no question, then, that limestone parent materials have ex-
erted a profound selection on regional floras, resulting in unique vegeta-
tional composition, physiognomy, and soil formation. It remains now
to look at possible physiological explanations of accommodation to
limestone.

Physiological explanations of preference for, or avoidance of, lime-
stone soils are not wholly satisfying. Obscuring the search for answers
are a number of complicating ecological and soil chemistry factors. Do
calcicoles require a medium high in calcium or are they merely able to
tolerate high calcium in exchange for a release from greater competition
stress on non-calcareous soils? Is the limestone effect one of pH pref-
erence or more fundamentally a nutritional problem? The high pH values
generated by some limestone soils no doubt exert strong side effects on
the availability of other elements, e.g., iron, aluminum, manganese, phos-
phate. A physiological approach has been fruitful in the case of differ-
ences in calcium preference of grasses in the genus Agrostis (Clarkson,
1965). Of four species of Agrostis grown in controlled solution cultures
of various calcium regimes, the well-marked calcifuge species, A. setacea,
has a significantly different capacity for calcium uptake. The results for
the three calcicole and one calcifuge species appear to be related to dif-
ferences in the capacities of their active transport system—the metabol-
ically controlled mechanism for moving ions from soil to root interior.
The calcifuge species, A. setacea, seems to have a calcium transport sys-
tem of lower capacity than the other three. The ecological assessment of
these results would appear to be that a calcifuge species may have an
inherently lower threshold to calcium uptake.

A now classic study of a calcicole-calcifuge species pair by A. C. Tans-
ley, pioneer British ecologist, has a timely and contemporary message
for anyone studying the effects of a single soil variable on plant distribu-
tion. Tansley (1917) demonstrated that species interactions greatly al-
tered the effect of soil type on plant growth. The two bedstraws, Galium
saxatile and G. sylvestre, were grown in pure and mixed stands on acid
peat and calcareous soils. ‘“Both species can establish and maintain

1969 | KRUCKEBERG: SOIL DIVERSITY L395

themselves—at least for some years—on either soil,” but ‘the calcicole
species is handicapped as a result of growing on acid peat and therefore
is reduced to subordinate position in competition with its calcifuge rival,
which is less handicapped,” and “. . . the calcifuge species (Galium sax-
atile ) is heavily handicapped especially in the seedling stage, as a direct
effect of growing on calcareous soil, and is thus unable to compete effec-
tively with its calcicole congener, Galium sylvestre.” Tansley’s work em-
phasizes the obvious but often overlooked danger of reading too much
ecological significance into results obtained from plants studied in arti-
ficial isolation. The current research on plant competition by John
Harper (1967) and his associates in Britain underscores the importance
of variability in plant response as influenced by biotic interaction.

Lists of species pairs, calcicole versus calcifuge, imply that taxonom-
ically recognizable kinds of plants have different tolerances and that the
members of a given pair may be closely related. Though there may be
some question as to the advisability of giving such vicariads taxonomic
recognition, there is no doubt that there are interpopulational differences
to an edaphic factor such as limestone. In pursuit of this possibility, it
has now been amply demonstrated that species occupying a diverse array
of edaphic habitats have responded genetically to variant selective agents
of the soil; i.e., such species exhibit ecotypic differentiation. Working
with Trifolium repens, a species with wide edaphic range, genecologists
in Britain have found intraspecific variation in tolerance to calcareous
and acid soils. Snaydon (1962) concludes that, “. . . the wide edaphic
range of T. repens is due, at least in part, to the presence within the
species of specifically adapted physiological types.”” When such intraspe-
cific but interpopulational differences include morphological characters,
and when the contrasting edaphic factors act as isolating barriers, both
taxonomic separation and microevolutionary divergence are demonstrable.

It is too simple to hope for a one-to-one correspondence between the
chemical nature of a calcareous substrate and a selective action on the
potential flora the rock may support. Species which may be highly re-
stricted to limestone in one area may be indifferent to such substrates
elsewhere along their range. This is likely the case for the peculiar floris-
tic composition of the Convict Creek basin in the Sierra Nevada of Cali-
fornia. Major and Bamberg (1963) describe a remarkable aggregation of
geographically disjunct species in the basin. Several taxa, otherwise found
only far to the east or north, occur on a narrow band of marble in the
basin; this highly distinctive calcareous substrate intrudes locally at
Convict Creek, the monotonous granodiorite of the Sierras. It appears
that the locally arid outcrops and the contrasting moist seeps provide
habitats not otherwise available on the high eastern slope of the massif.
The high calcium content of the marble is thus only secondary or even
irrelevant to the local occurrence of the disjunct species.

The end result of interactions between substrate and floristic composi-
tion can be clearcut: the patterning of vegetation we see can then be di-

136 MADRONO [Vol. 20

Fic. 1. Mosaic of vegetation types in the White Mountains, California. Sage-
brush on sandstone and bristlecone pines on dolomite. Photograph taken by Albert
Hill and furnished by Harold Mooney.

rectly related to the distribution of the parent materials. But when one
is led to sort out the properties of the environment which yield the flor-
istic end product, the story becomes complex. Take a recent case, that of
the clearly substrate-oriented distribution of bristlecone pine, Pinus aris-
tata, of the White Mountains in eastern California. The pines, now ac-
claimed the real patriarchs of the plant world, occur chiefly on dolomitic
limestone, whereas sagebrush is dominant in adjacent granitic and sand-
stone soils (fig. 1). Physiological ecologists (Wright and Mooney, 1965)
find that it is the interaction of physical, nutritional and biotic factors
that lead to the complementary distribution of the pine and sagebrush
(Artemisia tridentata) dominants. The light-colored dolomitic soils are
moist and cool, and yet are highly deficient in phosphates. Sagebrush is
excluded from the dolomite by the phosphorus deficiency, and recip-
rocally the pines prefer the cooler, moister dolomites, while tolerating the
low phosphorus status. The temperature-moisture difference and the
phosphorous deficiency thereby effect a competitive relationship which
results in the visible substrate-oriented patterning of vegetation.
Vegetation on Acid Soils

The story of vegetation on limestone tells us that the plant response
runs the gamut from narrow calcicolous restriction through broad toler-

—— eee ———

1969] KRUCKEBERG: SOIL DIVERSITY 137

ance or indifference to clear avoidance of the substrate. Such an array of
responses largely repeats itself whenever exceptional chemically limiting
edaphic responses occur. At the risk of being too inclusive, I want to ex-
amine this range of plant response for other chemically unique substrates.
For most of them only limited discussion is possible. The mere catalog
of other unusual substrates is intriguing in itself. Thus, to use low pH
as a crude basis for compilation, there are the highly acid soils induced
by a variety of exceptional parent materials: Aluminum-rich bauxites or
terra rossa soils of the tropics, silica-rich soils (sands, diatomaceous
earths, slates, laterites, etc.), hydrothermally altered volcanics (rich in
sulfates), and the soils of lead mine tailings and zinc deposits.

Studies prior to 1957 on the effects of aluminum on plant life led to
the generalizations, 1, that soils with high Al may restrict dicot weed
competition in grass pastures, 2, that on Al-rich soils there are three
levels of Al uptake: a, plants requiring aluminum ions in their metab-
olism, b, plants known as “aluminum accumulators,” which concentrate
Al ions in plant tissues with visible but non-lethal effects, and c, plants
which are tolerant of Al but collect little or only small percentages of Al
in their tissues, and 3, that certain plant families or genera either re-
quire Al for normal growth, e.g., Ericaceae, Moraceae, Ferns, and Lyco-
podiaceae, or are accumulators of Al (Carpinus, Rubiaceae, and Melasto-
maceae). Several of these generalizations have now been put to the test of
careful field observation by Howard and Proctor (1957). A major por-
tion of the lowland land surface of Jamaica contains aluminum-rich baux-
ite deposits. Although agriculturally poor, the bauxitic areas do support
a mixture of cultivated crops and a native vegetation in varying stages
of secondary succession. Since undisturbed vegetation on bauxite is in
remote areas and as yet unsampled, studies on the effects of Al on vege-
tation were restricted to disturbed sites. The authors concluded: “it ap-
pears that the bauxite flora of Jamaica consists of plants which are un-
affected by aluminum and tolerant of its presence . . . to the present we
have found no species characteristic of bauxite soils, nor have we dem-
onstrated that the vegetation of adjacent areas currently not found on
the bauxite deposits will not grow on the bauxite soils. To the contrary,
the invasion of plants from adjacent areas on barren, mined-out pits and
the plantations established in these pits indicate that factors other than
the concentration of aluminum will control the success or failure of these
species on bauxite.”’ At this point we would have to conclude the effects
of Al on floristic pattern and vegetation are unresolved.

Highly acid and infertile soils underlain by sands and other siliceous
substrates often support unique plant assemblages. The pine barrens of
New Jersey, the shale barrens of the Appalachians, and possibly the
coastal sands of the Carolinas are eastern representatives of the type.
Notable in the West are the Mendocino barrens and the laterites-seri-
citis schists in the Sierra Nevada foothills of California, while in the

138 MADRONO [Vol. 20

Great Basin’s desert and mountain country the hydrothermally altered
volcanics create local vegetational discontinuities.

Just back of the Pacific coastline in Mendocino Co., California, is a
dissected sandstone plateau which supports that most remarkable vege-
tation, the “pygmy forest” (Jenny, et al., 1969). In its most extreme rep-
resentation a dense growth of cane-like dwarfed individuals of Cupressus
pygmaea and Pinus bolanderi, not over eight feet tall, cover the ashy
gray podsolic soils. Notable associates of the pygmy conifers are several
ericaceous shrubs. Of the latter, Arctostaphylos nummularia is endemic
to the pygmy forests and is one of the three rare acid-soil endemics in the
section Schizococcus. Since other species of conifers occur on exceptional
soils, the possibility exists that there are features in common among sev-
eral atypical (non-zonal) soil types in California that yield unusual
floras. The general conclusions from thorough field study, greenhouse
culture work (mineral nutrition) and laboratory analysis of soils and
plant material (McMillan, 1956) merit our attention. Two problems
needed explanation: 1, the restricted distribution of Cupressus species
on a variety of exceptional soil types including the Mendocino acid bar-
rens, and 2, the anomaly of good growth of cypress seedlings on fertile
and infertile soils in greenhouse culture. McMillan suggests 1, that
edaphic restriction of native plants is not tied to a particular nutritional
requirement provided solely by the unique substrate; 2, some common
physiological tolerance, e.g., to low calcium availability, may be the
basis of generic differentiation in Cupressus but not so for other genera
of similar edaphic predisposition, and 3, that the pine barren plant com-
munity is an array of species that results from “‘the overlapping of differ-
ent tolerance ranges of the component individuals for environmental
conditions presented by a particular habitat.”

In the Great Basin region of the West, broad expanses of sagebrush,
juniper, and saltbrush desert are occasionally interrupted by isolated
stands of yellow pine and other disjunct subordinate species. These re-
markable floristic islands are usually found to be growing on local non-
zonal soils of exceptional nutrient characteristics. In both Nevada and
Utah, such restricted isolates of vegetation occur on highly acid soils de-
rived from hydrothermally altered lavas and volcanics. Billings (1950)
found that the altered andesites northwest of Reno were ‘‘very deficient
in exchangeable bases, phosphorous, and nitrogen as compared to” adja-
cent zonal soils supporting pinyon-juniper and sagebrush. Billings con-
cluded that ‘“‘the pine stands are relicts which have remained because of
the inability of sagebrush zone dominants to invade these mineral-de-
ficient soils.”’

The vegetation of desert “islands” of altered volcanics in Utah sub-
stantiate Billings’ views on the casual nature of the floristic isolation.
Salisbury (1964) adds to the total picture by suggesting that succession
to zonal soils typical of the regional climate can occur under the influence

1969] KRUCKEBERG: SOIL DIVERSITY 139

of the vegetation itself especially through humus accumulation and even-
tual plant succession . . . even on these altered volcanics of low pH. The
soil profiles from extreme (non-zonal) to zonal sites show an amelioration
of the pH and nutrient status of the soil. Leaching of the undesirable
elements and the biological addition of essential nutrients appears to
achieve the successional change. We would predict, however, that succes-
sion to zonal status could be achieved only under ideal conditions of
topography, moisture and vegetation cover. Steep slopes of altered ande-
site with high runoff would undoubtedly persist as non-zonal, sterile soils.
In fact, we could generalize to say that severe topography coupled with
exceptional parent material will permanently arrest soil formation at the
azonal or skeletal state; the biological consequence would be the per-
sistence of a pioneer, edaphically specialized endemic flora.

The last example of non-zonal acid soil and its influence on plant dis-
tribution is not only fascinating in its own right, but fosters some far-
reaching generalizations. The remarkable restrictions of Arctostaphylos
myrtifolia in almost pure stands to Eocene laterite and to sericitic schists
in the Sierra Nevada foothills of California has been thoroughly studied
by Gankin and Major (1964). Near Ione and San Andreas a non-zonal
acid heath association abruptly interposes itself within the regional cli-
max vegetation. The Ione manzanita occurs often in dense heath-like
stands on substrates of low base status, low fertility, of exceptionally low
pH values, 2.0 to 3.95, and of high soluble aluminum values. It is con-
tended that it is the high soil acidity and high aluminum content which
exert such a strong selective inclusion-exclusion effect on the regional
flora. After citing a number of other examples of edaphically controlled
endemism and disjunct distributions, the authors seek a common cause
(p. 803): “The above examples could be expanded, evidently indef-
initely. Once this principle of disjunct and endemic plant occurrence on
non-zonal sites is accepted, examples become almost too numerous. In all
these cases,, explanations of why the rare plants occur where they do in
terms of plant physiological reactions are completely lacking. Judging
from the cases cited, they would have to be conflicting. The only explan-
ation which fits the diversity of facts—that is, plants occurring at higher
or lower altitudes than normal, in wetter habitats or drier, with less cal-
cium or more—is in terms of plant competition. All the cases fit the con-
clusion that rare or disjunct (non-zonal) plants can occur in a given
area where competition is decreased by some kind of extraordinary soil
parent material or other continuously effective disturbance of climax
vegetation development.” With that conclusion I would concur, but
would at the same time suggest that the competition hypothesis opens up
still another “‘Pandora’s Box” of complex biotic interactions. Competi-
tion, like endemism, soil infertility, and pH, is as yet a rather vague
concept, at least in contemporary plant ecology. Attempts to analyze
“competition” and to test its complex nature are only recently gaining
fruitful momentum.

140 MADRONO [Vol. 20

Soils and Vegetation on Serpentine and other Ultramafic Rocks

The last and certainly most spectacular “abnormal” (azonal) soil to
be discussed is that derived from serpentine and other ferromagnesian
rocks. The plant life on such soils has held particular fascination for gen-
erations of botanists. On nearly every major land mass of the world,
ferromagnesian (ultramafic) outcrops weather to soils that exert a pro-
found selective influence on the regional flora. Stark contrasts between
the barrenness of ultramafic and the comparative luxuriance of adjacent
non-ultramafic sites, as well as the pronounced differences in species
composition are familiar and striking features of this discontinuity in
vegetation dominated by geology. Although the most celebrated manifes-
tations of ultramafic vegetation are in Europe (the “dead” Alps, the Bal-
kan Peninsula and northern Sweden) and North America (central Califor-
nia to Oregon and Washington, and the Gaspé Peninsula) other areas,
both tropical and temperate, show tell-tale vegetational responses to
these soils. Cuba, New Caledonia, New Guinea, New Zealand, and Japan
also have notable areas of serpentine and related rocks which in turn
support unique floras (Krause, 1958; Whittaker, e¢ al., 1954).

Before turning to the floristics and ecology of serpentines, we should
set the scene. I propose to use the word ‘‘serpentine” broadly to encom-
pass all ultramafic rocks and soils weathering from them. The term
“ultramafic” (or “ultrabasic’’?) embraces those rock types in which the
mineralogical composition is largely in the form of silicates of iron and
magnesium, as exemplified by the mineral, olivine. The commonest ultra-
mafics are the igneous rocks, peridotite and dunite, and their metamor-
phic derivative, serpentine. Soils weathering from such rocks are high in
magnesium and low in calcium; because of other minerals, pyroxene,
amphibole, chromite, etc., in additional to the crucial olivine, the soils may
also contain unusually high amounts of nickel and chromium. A secon-
dary biological effect during soil genesis is the common deficiency in
nitrogen and phosphorus. Serpentine soils are both unfit for most agri-
culture and highly selective for native plant species. The calcium-mag-
nesium ratio of much less than 1.0 is considered to be a crucial selective
soil factor for the distribution of plant species. Serpentine usually has
both a physiognomic as well as a taxonomic effect on plant life. Serpen-
tine vegetation is sparse, with much intervening barren ground; dwarfing
and xerophytism are common. Species composition is both depauperized
and often unique; endemism and range disjunction are frequently the
most outstanding floristic attributes.

Both because they are spectacular samples of serpentine vegetation
and are reasonably representative of temperate zone ultramafics, I will
confin my discussion to the serpentines of western North America. For
convenience, we can distinguish three physiographic regions in which
serpentines abundantly occur: 1, the Central California Coast Ranges—
Sierra Nevada foothills area, 2, the Klamath-Siskiyou area, and 3, the
Northern Cascades—Wenatchee Mountain areas. We will look first at

1969] KRUCKEBERG: SOIL DIVERSITY 141

Fic. 2. Outcrop of serpentine overlooking meadow of mixed alluvium, three miles
northeast of Middletown, California.

the vegetational and floristic responses, then at the genotypic reactions
of populations to serpentine, and finally develop hypotheses to account
for the evolution and adaptation to the serpentine habitat. All along the
north-to-south transect, especially from Douglas Co. in Oregon, to San
Luis Obispo Co. in California, abundant and often extensive ultramafic
outcrops serve to further complicate the already intricate environmental
mosaic. The North Bay counties (Napa, Lake, Marin and Sonoma) of
central California afford an ideal locale in which to sample the central
Californian version of the vegetational discontinuities associated with
serpentine outcrops (fig. 2). The serpentines here stand in sharp contrast
to the adjacent non-serpentine sites which support largely wide-ranging
woody dominants of either the oak woodland, mixed conifer, or chaparral
type. Such sclerophyllous shrubs as Quercus durata, Ceanothus jepson,
Garrya congdonii, and even the small coniferous trees, Cupressus sar-
gentu, and C. macnabiana, are unmistakable ‘indicator’ species because
of their typical restriction to and numerical dominance on serpentine
soils. It is not these dominant woody species, however, which have made

142 MADRONO * OLVol. 20

Californian serpentines celebrated as a source of rare and endeimc plants.
The transient spring flora of the dry serpentine hills still continues to be
a source of “new or otherwise noteworthy” additions to the California
flora. From the time of E. L. Greene and W. L. Jepson to recent collect-
ing by Freed Hoffman, John Thomas Howell, John Morrison, Helen
Sharsmith and others, the list of herbaceous rarities endemic to serpen-
tine has grown and continues to grow. A genus of crucifers, Streptanthus,
is particularly rich in serpentine forms and well serves as an example of
wholesale evolutionary diversification on this selective substrate. At least
12 species in California and southern Oregon occur on serpentine: for
example, S. niger, Tiburon Peninsula; S. batrachopus, Mount Tamalpais;
S. insignis, San Benito Co.; S. polygaloides, Sierran foothills; S. hesper-
idis and S. brachiatus, Lake Co.; and S. morrisonit, upper Austin Creek,
Sonoma Co. A few are just as obligate on serpentine but of wider range:
S. howellit. Siskivou Mts.; and S. barbatus, S. brewert, S. barbiger, and
S. drepanoides, Napa to Trinity counties. Still others have a broader eda-
phic tolerance and occur both on and off serpentine: S. glandulosus, San
Luis Obispo Co. to southern Oregon; and S. tortuosus, Sierra Nevada—
Coast Range-Siskiyou triangle, though some named intraspecific taxa ap-
pear to be local serpentine endemics. Populations of S. glandulosus when
grown on test serpentine soil proved to be most instructive in the quest
for an explanation of serpentine restriction (Kruckeberg, 1951). Collec-
tions from non-serpentine sites were clearly intolerant of serpentine soil,
while morphologically indistinguishable serpentine samples grew vigor-
ously on the same test soil. Infraspecific variation in physiological tol-
erance is clearly demonstrated here and expands the idea of ecotypic
differentiation of species beyond climatic response to that on soil differ-
ences. Streptanthus glandulosus is therefore interpreted as a species orig-
inally possessing several edaphic biotypes and that through time non-
serpentine biotypes have been gradually eliminated. Ultimately its fate
may be that of its obligate serpentine endemic relatives, restricted to ser-
pentine. I would concur with Gankin and Major (1964) that it is the
‘pressure’ of competition—though its action unspecified as yet—that re-
duces biotype diversity and forces ultimate confinement to serpentine. In
this sense then, some of the narrow endemics of Streptanthus appear to
be “depleted” species. Biotype depletion need not be the prelude to ex-
tinction, however. Having found refuge as edaphic specialists on serpen-
tine, diversification within the serpentine environment may ensue. This
seems to have ben the speciational history in at least three subsections,
Insignes, Pulchelli, and Hesperides, of the genus.

When we move from the xeric chaparral-covered serpentines of Cali-
fornia to the more mesic serpentine habitats in the Siskiyou Mountains
to the North, we find a definite shift in composition of vegetation.
Though the contrast between serpentine and non-serpentine plant asso-
ciations is often as striking as those to the South, both species and life-
form composition are clearly different. Annuals and chaparral shrub spe-

1969] KRUCKEBERG: SOIL DIVERSITY 143

cies no longer dominate the ultrabasic landscape. Rather, it is the par-
ticular blend of widely spaced conifers and intervening broadleaved shrub
and herbaceous layers that characterize the mountainous serpentines
the Siskiyous. The forest-shrub complex on serpentine (Whittaker, 1960)
gives way abruptly to climax (?) montane mixed coniferous forests on
neighboring non-serpentine soils. Repeating the character of the highly
acid non-zonal soils discussed earlier is the occurrence of open mixed
stands of conifers, often stunted, composed in part of species not common
on adjacent ‘‘normal”’ soils. The occurrence of certain conifers on ser-
pentine appears to be the result of altitudinal and/or geographical exten-
sions of more typical ranges of the species. Pinus jeffreyi, Jeffrey pine,
and P. attenuata, knobcone pine, fit this category of disjunct distribu-
tions, repeating here in the Siskiyous what is notable about their distri-
butions elsewhere in the far West.

Most remarkable is the shrub cover of Siskiyou serpentines. Nearly
every taxon of the sclerophyllous shrub layer is a varietal xeromorph of
a species more typically of mesophytic and arborescent habit. Whittaker
has observed example after example of these ‘‘trees-turned-shrubs,” or
mesic-turned-xeric shrub. “Quercus chrysolepis is represented on serpen-
tine by var. vaccinifolia, the most abundant single shrub species there;
Lithocarpus densiflora is represented by var. echinoides, Umbellularia
californica by an unnamed shrubby variant, and Castanopsis chrysophylla
by var. minor (uncommon in the study area). Quercus garryvana occurs
on serpentine as the shrubby var. breweri. Among other trees and shrubs
a series of congeneric pairs appear in non-serpentine and serpentine
floras with the serpentine species in each case of smaller stature: Amel-
anchier florida and A. gracilis, Garrya fremonti and G. buxifolia, Rham-
nus purshiana and R. california occidentalis, Holodiscus discolor and H.
dumosus, Ceanothus integerrimus and C. pumilus, and Berberis nervosa
and B. pumila.”

Local moist seeps at the base of serpentine-peridotite slopes are havens
for some of the most spectacular of the Siskiyou endemics: Cypripedium
californicum, Rudbeckia californica, Darlingtonia californica, Trillium
rivale, Lilum bolanderi, and L. occidentale. There is no doubt that the
rich endemism of the Siskiyou Mountains can be correlated in large part
to the “insular” occurrence of ultrabasic rocks.

Contrasts between serpentine plant life of the Siskiyous and that of
western and central Washington are muffled by the presence in both of
a coniferous forest cover. Yet species differences between the two areas
are truly sharp. The most telling contrast is in the reduction in species
diversity on the Washington ultrabasics. Though there are remarkable
species discontinuities and edaphic restrictions in the state, the serpen-
tine flora is markedly depauperate compared to that on the Siskiyou and
Californian serpentines. But before we deal in specifics, let me set the
physiographic scene for display of plants on ultrabasics in Washington.

Ultramafics in the state occur in two major settings (Kruckeberg,

144 MADRONO [Vol. 20

Fic. 3. Barren serpentine slopes within coniferous forest type, headwaters of
Boulder Creek in Cle Elum River drainage, Wenatchee Mountains, Washington.

1969). The largest exposure is in montane portions of Kittitas and ad-
jacent Chelan counties. The sites are all in the Wenatchee Mountains
which form an easterly extending spur of the Cascade Range. The out-
crops occur either as peridotite, dunite, or serpentine; exposures of the
rock may be massive, of many square miles in extent, or very local (fig.
3). Old altered volcanics (greenstones), sedimentary rocks, gneisses and
schists, as well as acid igneous granodiorite border or even interfinger
with the ultramafics. The region is thus lithologically rich and complex.
The terrain is rugged, with steep slopes and high ridges that culminate
in the ultramafic peaks, Earl, Navaho, and Ingalls, from 5000 to 7000 feet
altitude. The clearest and most spectacular contact between ultramafic
and non-ferromagnesian rock types is along upper Ingalls Creek where
the east boundary of peridotite at the creek abruptly gives way to the
massive granodiorite of the Stuart Range.

All of the Wenatchee Mountains ultramafics occur in areas of conif-
erous forest. At altitudes from 2400 to 4000 feet, the forest consists of
open stands of Douglas fir, yellow pine, and western white pine; this
forest type grades insensibly upward into a mixture of subalpine fir,
mountain hemlock and whitebark pine. The stand are invariably open,
the barren slopes between the scattered trees lightly populated with

1969 } KRUCKEBERG: SOIL DIVERSITY 145

grasses and forbs, some of which are highly characteristic of ultramafic
soils.

The next largest series of ultramafic occurrences in Washington is in the
northwestern counties of Snohomish, Skagit, San Juan, and Whatcom.
The most outstanding of these is Twin Sisters Mountain, a westerly out-
lier of the northern Cascades; it is pure dunite, an igneous ultramafic
composed primarily of the mineral olivine. Rock of similar origin occurs
locally at low elevations to the west; Fidalgo Island and Cypress Island
have the most extensive of this series of ultramafic outcrops.

The vegetation on the Twin Sisters dunite contrasts strikingly with
that on the adjacent non-ferromagnesian parent materials. The luxuri-
ance of the Humid Transition forest abruptly gives way to stunted
Douglas fir, lodgepole pine, western white pine and shrubby Juniperus
communis. The insular ultramafics also support conifers, largely Douglas
fir, Pinus contorta, and J. scopulorum.

The coniferous forest on ferromagnesian substrates is by no means
dense and continuous. The trees are largely stunted and widely spaced;
often on steep, stony serpentinized outcrops there are no trees present.
On such barren, continuously eroding slopes, as well as on talus, in rock
fissures and on sparsely forested slopes, one is almost sure to find a rep-
resentation of species restricted to the ferromagnesian soils and rock.

The flora of the Wenatchee Mountains has received the lion’s share of
my attention. The serpentines of this rugged range support a depauper-
ate flora, a shifting, variable mosaic of both indicator-endemics and wide-
ranging edaphically indifferent (“bodenvag”) species. The ultramafic
rocks at the same time exert a pronounced exclusion effect on much of
the regional flora on adjacent non-serpentine habitats. From field records
of observations on 36 serpentine and 30 non-serpentine sites a picture of
partial floristic discontinuity has emerged and yields fruitful generaliza-
tions on the effects on the flora of these northern ultramafic soils. 1. Ex-
treme barren serpentine habitats are depauperized in species number,
especially in species of the tree and shrub life form. Such serpentine bar-
rens within the coniferous forest biome appear as though the alpine and
timberline zones have been eccentrically displaced downward in altitude.
2. There are indeed endemic and indicator species on Washington ser-
pentines. These are: Polystichum mohrioides lemmoniu, Cheilanthes sili-
quosa, Poa curtifolia, Eriogonum pyrolaecfolium coryphaeum, Arenaria
obtusiloba, Claytonia megarhiza nivalis, Anemone drummondiu, Thlas pi
alpestre, Ivesia tweedyi, Lomatium cuspidatum, Douglasia dentata ni-
valis, and Chaenactis thompsonii. These twelve species are strong indi-
cators of serpentine; all are herbaceous (two ferns, a grass and nine dicot
forbs). 3. Conifers for the most part do not show any marked edaphic pref-
erences. I have observed certain low to mid-montane coniferous species to
occur at higher altitudes on serpentine: Pinus contorta latifolia, lodge-
pole pine; P. ponderosa, yellow pine; P. monticola, western white pine;
and Taxus brevifolia, western yew. On massive dunite of the more west-

146 MADRONO [Vol. 20

erly Twin Sisters Mountain, lodgepole pine is the dominant timberline
tree; it is unknown in this role elsewhere in the Pacific Northwest. A
reverse displacement occurs for the three other conifers. Pinus albicaults,
whitebark pine; Abies lasiocarpa, subalpine fir; and the shrubby Junip-
erus communis occur at lower than normal elevations on serpentine.

With respect to the flora on nearby non-serpentine soils, it is clear that
a large number of species (35) avoid serpentine. This ‘‘serpentinophobia”’
is most evident where serpentine rocks contact other rock types such as
granite, greenstone, sandstone, etc. For some plants, avoidance of ser-
pentine becomes a family or generic matter. Though rich in species in
the Pacific Northwest, the genus Penstemon fails to occur on serpentine.
The Ranunculaceae, Saxifragaceae, Leguminosae, Rosaeae, and Erica-
ceae are conspicuous by their scarcity or absence on serpentine.

In Washington as in other parts of the world, some elements of the re-
gional flora appear to act indifferently to serpentine. The soil-wanderers
(bodenvag species) in Washington form a conspicuous element of the
flora, especially since most are conifers. There is no doubt though that
species of this category often are responding genetically to the serpen-
tine habitat. Ecotypic differentiation into serpentine tolerant strains has
been demonstrated for Washington serpentine flora just as clearly as for
the Californian examples (Kruckeberg, 1967). Nine of the 18 bodenvag
species tested clearly showed differences in serpentine tolerance. Six
showed signs of the same phenomenon, while only three species failed to
show ecotypic variation. The clearest responses were with herbaceous
perennials, e.g., Achillea lanulosa, Fragaria virginiana, Prunella vulgaris,
and Rumex acetosella. The two latter species are especially noteworthy
inasmuch as they are introduced species on serpentine. They have re-
spondd adaptively to selection for serpentine tolerance probably within
the last 50—75 years. At first it appeared that coniferous bodenvag species
were not ecotypically differentiated. Only after a long period of growth
(2 years) has it been possible to detect ecotypic response in lodgepole
pine, P. contorta latifolia.

The most faithful indicators of Washington serpentines are two ferns,
Cheilanthes siliquosa, rock brake, and Polystichum mohrioides var. lem-
moni, (Kruckeberg, 1964). The Cheilanthes rarely fails to appear on
even the most isolated and smallest ultrabasic outcrop, and at all alti-
tudes from sea level to timberline. Though restricted to higher altitudes
the Polystichum is just as reliable an indicator. One is led to assume that
spores of these serpentinophytes are widely dispersed or at least in a
regionally chain mail fashion, but only do they establish populations fol-
lowing germination on soils of ultramafic origin. The distribution of C.
siliquosa spans the North American continent from the Gaspé of Quebec
to British Columbia and thence to central California and nearly always
its discontinuous range coincides with the outcropping of ultramatfics.

“Normal” (Zonal) Soils
Admittedly arbitrary is the antithesis—normal versus abnormal soils.

1969 } KRUCKEBERG: SOIL DIVERSITY 147

Yet to the extent that climate or parent material are primary in deter-
mining the quality of a soil, the distinction is justified. We would, there-
fore, expect that normal soils will be characterized by properties derived
from other parent materials. In other words, given a range of unexcep-
tional parent materials in a region of similar climate, all normal soils
would be nearly alike, converging on common properties due to the over-
riding effects of the regional climate. The vegetational response to a
single soil type on differing parent materials should be homogeneity,
other factors being the same.

Does this in fact ever occur? A test of this progression to sameness of
soil from different parent materials could only be made under ideal con-
ditions. For example, chemically similar and “‘normal”’ parent materials
of varied origin, e.g., volcanic, intrusive, metamorphic and sedimentary
rocks would be expected to have weathered over the same periods of time,
would have to be subjected to the same succession of biota and would
have to be compared under similar topographic sites. The concurrence of
all of these seems unlikely. Even where the properties of the differing
parent materials are not extreme, physically or chemically, soil and vege-
tation differences are likely to exist. Two of the examples cited earlier can
be brought to bear on this point. According to Wright and Mooney
(1965), it is the dolomite which is the preferred substrate for the White
Mountain bristlecone pine. The sandstone and granite both support the
sagebrush dominant and much of the subordinate vegetation. Yet the
species composition on the two latter substrates does differ: between the
two more normal rock types there are substrate preferences by the flora.

Whittaker’s (1960) study of soil preferences by the flora of the Sis-
kiyou Mountain included comparisons between two rock types, diorite
and gabbro, less extreme than serpentine-peridotite. Soils derived from
diorite and gabbro are much alike chemically. Assuming uniform soil
forming factors other than parent materials, the two parent materials
should support rather similar vegetation. On the contrary, differences in
vegetational composition still do occur and are substrate-dependent.
Even dominant tree species show differential responses; species on the
gabbro occur with greater frequency on more mesic sites. Whittaker gen-
eralizes this ‘‘shift toward the mesic” as a common trend associated with
change toward substrates higher in ferromagnesian minerals.

It is therefore likely that whenever geological diversity exists in moun-
tainous regions there will be corresponding floristic diversity. The more
extreme the lithological differences, the greater the differences in flora.
Substrate dependence will more likely be minimal in areas of minimal
topographic relief and/or on alluvial substrates. The latter condition
appears to hold for the Pacific Northwest where valley alluvium from
volcanics, sedimentaries, or granodiorites supports the same climax conif-
erous forest—including much the same species in the subordinate veg-
etation. Still another possibility exists: several vegetation types on a
single and local substrate. Wells (1962) finds this to be true in the cen-

148 MADRONO [Vol. 20

tral Californian coastal vegetation. He attributes this kind of vegeta-
tional heterogeneity to a history of fire, grazing and other disturbances.

Other Soil Properties Affecting Plant Distribution

Once formed from parent material by the processes of weathering, soil
comes into it own as a substrate which can develop unique properties in-
dependent of its mineralogical origin. The interplay between organism
and soil introduces new dimensions and creates new properties. Two
significant attributes of soil that affect plant distribution in remarkable
ways are currently under vigorous and fruitful scrutiny. The first of
these, soil microtopography, is usually intrinsic to soil and can vary
apart from the activity of associated biota. The other influential soil
property is the presence in soils of substances of biological origin that
promote or inhibit growth. Studies of these two properties are providing
unexpected insights into the basic ecological problem of competition (in-
terference) and evoke possible mechanisms for such ecological phenom-
ena as a succession, spatial pattern, endemism, etc.

Soil Microtopography

The fate of seed, once shed from the parent plant, is largely a matter
of chance. Where a seed is deposited will be a primary determinant of
germination. Even dispersal to a suitable soil or organic substrate is not
enough to ensure success. From the “ant’s eyeview” the surface of the
substrate can be mountainously irregular; and to the seed, major differ-
ences in soil microtopography may spell the difference between a favor-
able microenvironment for germination and failure. Harper, et al. (1965)
has studied the effect of microtopography on germination and has em-
phasized germination on ‘‘safe” versus “unsafe” germination sites as a
potent control of plant populations. Their earlier experiments (Harper,
1961) were done with seed of annual grasses, Bromus species, sown on
two contrasting soil surfaces. On a uniformly rough surface, irregular
clumps of soil about ™% inch in diameter, there was a linear increase in
germination with increase in sowing density. But on a soil of regular
surface, checked by drying into smooth sectors bounded by cracks, ger-
mination failed to increase beyond a rather low density. Clearly the rough
surface provided ‘‘many more potential germination sites” than did the
smooth one. Only those seed that landed in the crevices germinated and
there the number of “safe” sites was limited! Harper, e¢ al. (1965)
greatly elaborated on this theme by using a variety of species of differing
seed size and increasing the variety of soil surfaces. There is no question
but that the physical heterogeneity of soil surfaces provides a range of
microhabitats both suitable and unsuitable for germination. The probable
effect of ‘‘safe’’ microsite may be to provide suitable moisture and tem-
perature conditions for germination. The effectiveness of soil pathogens
on reducing seedling survival must also depend on the quality of the par-
ticular microsite.

1969] KRUCKEBERG: SOIL DIVERSITY 149

Variations in soil microtopography are the product of soil-forming fac-
tors. This idea can be formalized in terms of Jenny’s soil-forming factor
equation (s = (cl, 0, r, p, t)) where “‘s” now is “soil microtopography.”’
We can draw from Harper’s work some rather far-reaching inferences
which bear upon competition, succession and plant distribution. 1. Dif-
ferent species will respond uniquely to different configurations of soil
microsites and thereby create local differences in species distribution. 2.
Microsite variations decrease the pressure of interspecific interference, if
the species have distinct safe site preferences. 3. Changes in microsite
through time from bare mineral soil through various successional stages
will result in the selection for different species at each stage of the seral
sequence. The concept of “‘safe” site thus is pregnant with experimental
strategems for studying a species niche, plant distribution, and the dy-
namics of vegetation.

Soil Inhibitors

The effect of a metabolite of one species on the survival and/or com-
petitive ability of another is well known to the protistan (microbial) and
aquatic animal ecologist. That an equivalent interaction mediated by
soil intake-output of metabolites can occur in higher plants thus appears
most reasonable. Though repeatedly suggested ever since the time of
Liebig, the possibility of promotion or inhibition of growth by metab-
olites which pass from plant to plant via the soil has only occasionally
been given serious attention. Paradoxically the botanist who now finds
good evidence for the phenomenon was in the position of having to deny
the ecological effectiveness of the first modern case of allelopathy, the
effect of plants on each other through their metabolites. Muller (1953)
could not substantiate under field conditions the inhibitory effect of me-
tabolites of Encelia, a desert shrub, on other plants which had been
found in laboratory tests of leachates. The inconclusive nature of anti-
biotic effects of plant-on-plant is attributed to the differences between
natural conditions in the field, dilution, microbial decomposition, soil
sequestering of leachates, etc., and the more concentrated doses com-
bined with ideal though artificial conditions in laboratory experiments.
More recently Muller and his associates (1964; 1965), have developed
incontrovertible evidence for the inhibition of vegetation by volatile
terpenes given off by species of Salvia and Artemisia in the California
coastal sage community. The causal basis, toxicity of terpenes, for swaths
or perimeters of sterile ground around the sage species has been con-
firmed in laboratory studies. The suppression is greatest against annual
grassland species. Moreover the toxic effect of the terpenes can be re-
tained by the soil for several months. Natural inhibitors can have a far-
reaching effect on floristic composition. A substance produced by the
shrub by chamise, Adenostema fasciculatum, excludes species of grass,
which in turn would otherwise exclude Dodecatheon clevelandii, a shrub-
tolerant herb. Thus this two-step biochemical exclusion creates an in-

150 MADRONO [ Vol. 20

hibitor-induced association of the chamise with the Dodecatheon.

The deposition in soil and recycling of organic metabolites is now well
established. A wide range of secondary metabolic products of plants,
carbohydrates, amino acids, organic acids, volatiles, alkaloids, etc., can
be recovered in the tissue of plants grown on substrates, soils or culture
solutions, that contain the substances (Grummer, 1961; Tukey, 1962;
Winter, 1961). Evaluation of the ecological role of such exogenous sub-
stances confronts the same problem of complex factor interaction that
persistently vexes the ecologist. A laboratory test of toxicity may not be
complemented by positive evidence of toxicity in the field. Differences
in concentration, unavailability of exudate-leachate due to adsorption or
microbial activity, rainfall-temperature effects, all may lessen or negate
the influence of the metabolite. Despite these reservations, it is becoming
increasingly clear that interference phenomena, and in turn, pattern and
distribution of species in communities, even the effective niche of a plant
species, can be importantly influenced by organic compounds cycled
through the plant-soil-plant system.

Evolutionary Consequences of Variations in Soils

Plants are ever responsive to differences in their environment that oc-
cur over spans of time and space. The effective level of response is the
population. The outcome of organism-environment interaction is either
adaptive response through natural selection or failure to colonize the
habitat, and hence, exclusion or extinction. Variations in soils, then, as
significant parts of the fabric in the environmental mosaic, operate as
agents of natural selection. Discontinuity of the soil features will further
act to isolate adaptive variants. The result of such interactions can be
discerned at various hierarchical levels and will be expressed in a variety
of modes. Microevolutionary reaction to soil differences will take the
form of degrees of ecotypic differentiation, the development of broad
genotypic tolerance, or ecological exclusion. The origin of species re-
stricted to edaphically unique habitats is a higher order of evolutionary
divergence. Raven (1964) has invoked the concept of catastrophic selec-
tion to account for diversification of edaphic specialists. Rapid selection
of exceptional genotypes under the stringent environment of azonal soils
is presumed to lead to fixation of unique, incipient populations. When
soil and biological discontinuities become congruent, isolation and spe-
cies formation then are promoted. Examples of evolutionary change up to
the level of species as occasioned by the selective action of soil differences
have been presented above in the section on “‘abnormal”’ soils.

Would we not expect some degree of edaphic preference to be expressed
in still higher levels of the taxonomic system? Couid not sections of
genera, entire genera, or even families show in substantial degree singular
edaphic restrictions? Such expression of specificity can be induced by
climate and is a major ingredient of speciation in the direction of adap-
tive radiation or extinction. The high incidence of the genus Sireptan-

1969] KRUCKEBERG: SOIL DIVERSITY BS.

thus to serpentine, many caryophylls to ultrabasics, Ericaceae to acid
soils, Cyperaceae to water-logged soils, genera of the Chenopodiaceae
and Amaranthaceae to nitrogenous or saline soils are all suggestive of
edaphic specialization. We would contend then, that soil, as is climate, is
a potent selective agency in securing evolutionary change.

Plants as Indicators of Mineral Deposits

The non-random distribution and abundance of plant populations in a
circumscribed habitat is the expression of one or more of a set of environ-
mental controls. It is as though the unique composition of a flora is tell-
ing the observer that some factor is having an overriding effect on the
composition of the plant cover. Plants which act as assay organisms for
some environmental component are called indicator species. The recog-
nition of plant indicators has been a traditional approach to the study
of environmental restraints on plant distribution. Ecologists, agricul-
turalists, foresters and range managers all use the sensitivity of plants to
environment in attempting to control or manage vegetation.

Plant indicators have been exploited in yet another way—prospecting
for mineral deposits. Deposits of a variety of minerals have been located
by searches in the field for the tell-tale displays of eccentric patterns of
plant occurrences or equally startling absences of occurrences. It is when
the indicator plants are found to contain unusual quantities of some min-
eral element that the geobotanical prospector strikes it rich. Biogeochem-
ical methods have now become standard practice for search for ore
deposits in the United States, Canada, Scandinavia, the U.S.S.R. and
elsewhere.

Let me relate a personal anecdote as a prologue to the description of
some of the results that the method has produced. During his nightly
rounds of our department, a faithful janitor would customarily linger in
the herbarium. Our suspicions were aroused by his preoccupation with
the contents of the herbarium cases, a conduct most unlikely for one of
his limited talents. His predilection for dried plants was, however, gen-
uine. He was scanning the contents of every case with a Geiger counter,
in the hope that somewhere in our Pacific Northwest collection, his coun-
ter would begin ticking at a runaway accelerated rate. His actions told
us that he was looking for uranium, at that time a much soughtafter ele-
ment. The outcome of his effort was, alas, unsuccessful, though the intent
was perfectly justified. Uranium deposits could be located by this
method!

More systematic and successful have been the operations of the Geo-
chemical Prospecting Methods Division of the U.S. Geological Survey.
Helen Cannon of the U.S. Geological Survey has published (1960) a
comprehensive review of geobotanical prospecting for ore bodies. Al-
though she points out that the recognition of absences of vegetation, or
unusual changes in appearance of plants also can yield “strikes,” it is the
plant indicator approach that concerns us here.

152 MADRONO [Vol. 20

The list of minerals which plant indicators can disclose reads like a
miner’s “Eldorado”: A conservative compilation would contain boron,
copper, gypsum, iron, lead, phosphorus, selenium, silver, uranium and
zinc. The copper indicators are both abundant and unusually reliable.
They “belong” mainly to three plant groups: the Caryophyllaceae or
pink family, the Labiatae or mint family, and the mosses. These copper
deposits have been located in Sweden by simply examining localities from
which the herbarium specimens of the ‘‘copper mosses” had been col-
lected. The copper indicators, Elsholtzia haichowensis from China, Acro-
cephalus robertu from Katanga, and Ocimium homblei from Rhodesia
all belong to the mint family and are very useful in prospecting. The
blue-flowered Ocimium homoblei will not grow in soil containing less than
100 parts of copper per million. The distribution of this plant has led to
the discovery of several ore deposits and is currently being mapped in
both Northern and Southern Rhodesia by the Rhodesian Selection Trust
(Cannon, 1960).

The well-known affinity of members of the loco-weed genus, Astrag-
alus, for selenium has led to uranium discoveries, since the occurrence of
the two elements is often highly correlated. A good example of the plant
indicator method comes from the work of Cannon’s group in western
United States. Several species, grasses, legumes, and composites, in the
shadscale—juniper vegetation of the Yellow Cat area in Grand Co., Utah,
proved to be consistent indicators of selenium. In this particular area,
selenium and molybdenum are useful pathfinder elements in prospecting
for uranium and vanadium. On mineralized soil indicator species con-
tained 6 to 11 times the amount of uranium found in unmineralized
ground.

As biogeochemistry becomes more sophisticated in technique, we would
predict additional rewarding mineral discoveries. Edaphic plant ecology
is certain to contribute to future mineral prospecting and as well, should
reap rewards for the student of plant distribution.

Epilogue

It is axiomatic in biology that complexity through factor interactions
breeds exceptions to consistent trends and that the analytic approach at
the community level must momentarily disregard complex interactions.
Our singling out of the soil factor in plant distribution has been just
such an over-simplification. The dwelling place of a particular species or
assemblage of species is the result of past and ongoing interplay between
biota and environment.

In this paper, we have taken the view that soil characteristics can often
have the dominating local or even regional impact of determining distri-
bution and/or pattern of plants in associations. Edaphic plant ecology,
then, becomes one useful key to the understanding of discontinuity in
vegetation.

A condensed version of this paper was presented at the American As-

1969 | KRUCKEBERG: SOIL DIVERSITY 153

sociation for the Advancement of Science Symposium, ‘Plant Biology
Today—Advances and Challenges,” Berkeley, California, December,
1965. The author’s studies on serpentine vegetation of Washington and
on the genus Streptanthus have been supported by N.S.F. Grants GS-
2792 and GB-4579.

Department of Botany, University of Washington, Seattle

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257-284.

PHYTOGEOGRAPHY OF NORTHWESTERN NORTH AMERICA:
BRYOPHYTES AND VASCULAR PLANTS

W. B. SCHOFIELD

INTRODUCTION

Within northwestern North America there are a number of fairly
natural phytogeographic regions, each characterized by a series of en-
demic and associated species with common affinities. The present survey
does not attempt to discuss all of these regions or to give exhaustive
lists of the flora that characterizes them. This study can be considered
very preliminary, to be amplified and improved with additional floristic
information and of detailed preparation of distribution maps. Those of
Hultén (1968) emphasize the importance of such maps.

Subspecific categories are not considered here. Although several spe-
cies are represented in western North America by endemic subspcies,
this is not indicated in the discussion.

In the present discussion the region covered is as follows: Alaska and
Yukon, British Columbia and the Rocky Mountains portion of Alberta,
and also including the area of Hitchcock, e¢ al., (1955-1969), Vascular
Plants of the Pacific Northwest: ‘““Washington State, the northern half
of Oregon (approximately north of the 44th parallel), Idaho north of
the Snake River Plains, the mountainous portion of Montana, and an
indefinite southern fringe of British Columbia.” For the distribution of
the vascular plants, therefore, the basic sources have been Hultén
(1968), Hitchcock, et al. (1955-1969), Henry (1915), and Eastham
(1947). Reference has been made also to the introductory portions of
Davis (1952), Peck (1941), Piper (1906), and Calder and Taylor
(1968). For the mosses the basic source of distributional data has been
Grout (1928-1939), although these data have been considerably ex-
panded. For the hepatics Frye and Clark (1937-1947) and Arnell
(1956) have provided general information, greatly amplified by more
recent literature.

The region covered is phytogeographically complex. It occupies an
area somewhat more than one third the area of Europe. The climate
varies from arctic to temperate, from oceanic to continental, from ex-
tremely humid to arid. Latitude ranges from north of the Arctic circle
(to somewhat beyond 71°N) southward to approximately 44°N, and
longtitude from 170°E to approximately 113°E. The elevation ascends
from sea-level to 20,320 ft., Mt. McKinley, Alaska, with numerous
mountain chains possessing peaks more than 10,000 ft. high. The geo-
logic substrata are equally complex, and widely dispersed through
various climatic extremes. Vegetation ranges from arctic and alpine

159

156 MADRONO [Vol. 20

tundras through boreal coniferous, montane coniferous, humid coastal
coniferous and drier coniferous forests, and arid grassland to semi-
desert. This is discussed by Daubenmire (1969).

Besides these factors, the present composition of the flora has been
moulded by historical circumstances. A major factor determining the
present ranges of species was the Pleistocene glaciations and the inter-
vening interglacials. Associated with climatic changes were variations in
the position and nature of corridors of migration, thus leading to ex-
pansion or restriction of floral boundaries. The flora available preceding
the Pleistocene glaciations is discussed by Wolfe and Leopold (1967)
and Wolfe (1969).

In Alaska and Yukon there existed, during the Pleistocene glacia-
tions, large unglaciated areas that served as refugia for the flora. This
flora consisted largely of circumboreal and circumpolar species, but also
possessed taxa surviving from floras of the more distant past, some of
them possibly from Tertiary time. These taxa are discussed in detail
later. Smaller unglaciated areas existed also in the Queen Charlotte
Islands, British Columbia, and possibly at higher elevations in the
Rocky Mountains of Canada.

South of the boundary of the continental glaciers, a considerable
portion of Idaho, Washington and Oregon has been colonized by plants
for many millions of years. Post-Pleistocene and recent variations in the
vegetational composition are discussed by Heusser (1960; 1965). Colin-
vaux (1967) has summarized the Quaternary vegetational history of
arctic Alaska.

Phytogeographic examination of the vascular flora of northwestern
North America appears to have lagged behind floristic studies. Although
Asa Gray (1859) compared the flora of western North America with
that of Japan, and Gray and Hooker (1880) analyzed the Rocky Moun-
tain flora, it is Piper (1906) who first attempted to summarize all of
the floristic elements. Harshberger (1911) also analyzed floristic ele-
ments, but concentrated on floristic composition of the vegetation in-
stead of the ranges of the species that make up the flora. Peck (1941)
has also summarized the floristic composition of vegetational areas in
Oregon, noting phytogeographic affinities. Weber (1965) has discussed
the phytogeography of the southern Rocky Mountains.

Although several studies of the phytogeography of California have
appeared (summarized in Stebbins and Major, 1965), the only other
major analysis of the total vascular flora of an extensive area in West-
ern North America is that of Cooper (1936) in his analysis of the strand
and dune flora of the Pacific coast. Unfortunately Detling did not com-
plete his comprehensive study of the flora of the Pacific Northwest, but
fragments of the manuscript have been published (Detling, 1968).
Northward, in Alaska, Hultén has published numerous studies, first his
beautiful synthetic study (1937) and culminating in his recent Flora of
Alaska and Neighboring Territories (1968). His atlases of vascular

1969] SCHOFIELD: PHYTOGEOGRAPHY 157

plant distributions (1958; 1962) have also contributed considerably to
the understanding of the ranges of plants in the boreal and arctic por-
tions of the Northern Hemisphere. Another publication of similar sig-
nificance is that of Meusel, et al. (1965). Distribution maps illustrating
ranges of arctic and boreal species in Canada have been published by
Raup (1947), Raymond (1950), and Porsild (1955; 1957; 1958; 1966),
and are particularly important in understanding taxa of circumboreal
and circumarctic distribution. In many cases these maps have shown
that earlier interpretation of species as disjunct have resulted from
inadequate collection. The check-list of Porsild and Cody (1968) adds
further such evidence, amplified somewhat in Cody and Porsild (1968).

A general phytogeographic synthesis for the bryoflora has not ap-
peared previously. Herzog (1926) has briefly summarized the affinities
of the bryoflora of western North America, with a concentration on
endemism. Irmscher (1929) has discussed disjunctions, and makes refer-
ence to western North American mosses. The most complete summaries
of bryophyte floristic elements are included in Evans (1914), Koch
(1954; 1956), Steere (1959; 1965), Persson (1949), Iwatsuki (1958),
Schofield (1965; 1968a; 1968b), and Ireland and Schofield (1967).
The maps of Szweykowski (1961-1969) are highly important in under-
standing the distribution of circumboreal hepatics.

In the following discussion the flora is treated initially by major
phytogeographic elements. Within each of these, more detailed distri-
butional patterns are considered. In each section hepatics are treated
first, followed by mosses and vascular plants. For the bryophytes the
order of taxa is basically that of Arnell (1965) for the hepatics, Crum,
et al. (1965) for the mosses, Hitchcock, et al. (1955-1969) for the vas-
cular plants peculiar to the region of that flora and Hultén (1968) for
Alaskan and other species of northern distribution.

ENDEMISM

In this category are treated those taxa that are either restricted to
the area of the study or extend into it, but are western North American
endemics. The latter designation includes taxa that are confined mainly
to areas in or west of the Rocky Mountains, infrequently extending east-
ward in the prairies or, occasionally, to the Black Hills of South Dakota.

ENDEMIC GENERA

In the bryophytes, although the number of endemic genera is not
great, northwestern North America shows greater richness than the re-
mainder of North America, north of Mexico. In the hepatics there is
the Family Gyrothyraceae with the genus Gyrothyra Howe (Schuster,
1955) which occurs from coastal northern California northward to
south coastal Alaska. This taxon has undoubtedly increased in abund-
ance with the increase in disturbance by man. It is particularly abundant
on roadside silts and clays.

158 MADRONO [Vol. 20

Endemic moss genera include: Crumia Schof., Roellia Kindb., Leuco-
lepis Lindb., Pseudobraunia (Lesq. & James) Broth., Alsza Sull., Den-
droalsia Britt., Bestia Broth., Tripterocladium (Mill.) Kindb., Trachy-
bryum (Broth.) Schof., and Rhytidiopsis Broth.

Of these genera Roellia, Trachybryum, and Rhytidiopsis are confined
largely to subalpine elevations, although Tvachybryum sometimes de-
scends to sea level and is occasionally associated with the oak woodland
from northern California to southwestern British Columbia. Leucole pis
is widespread at all elevations. Pseudobraunia, Alsia, Dendroalsia, Bestia,
and Tripterocladium are predominantly of lower elevations and occur
mainly west of the Cascade Mountains. Crumia is more widespread
(Schofield, 1966), being restricted by its calcareous seepage habitat
rather than by elevation. It it noteworthy that all endemic moss genera
except Pseudobraunia and Tripterocladium are dioecious and none
possess any special gemmae for vegetative reproduction, yet most show
very wide geographic range and often occur in great abundance.

In the vascular flora generic endemism is particularly notable.

Gramineae: Pleuropogon R.Br. and Scribneria Hack.

Liliaceae: Camassia Lindl., Leucocrinum Nutt., and Scoliopus Torr.

Orchidaceae: Eburophyton Heller.

Chenopodiaceae: Graywia H. & A., Nitrophila Wats., and Sarcobatus
Nees.

Portulacaceae: Calyptridium Nutt., Lewisia Pursh, and Spraguea
Torr.

Berberidaceae: Vancouveria Morr. & Dec.

Papaveraceae: Canbya Parry, Eschscholtzia Cham., and Meconella
Nutt.

Cruciferae: Anelsonia Macbr. & Pays., Athysanus Greene, Caulanthus
Wats., Chlorocambe Rydb., Idahoa Nels. & Macbr., Phoenicaulis Nutt.,
Physaria (Nutt.) Gray, Polyctenium Greene, Schoenocrambe Greene,
Stanleya Nutt., Streptanthella Rydb., Thelypodium Endl., and Thysano-
carpus Hook.

Sarraceniaceae: Darlingtonia Torr.

Saxifragaceae: Bolandra Gray, Conimitella Rydb., Elmera Rydb.,
Lithophragma Nutt., Peltiphyllum Engl., Suksdorfia Gray, Telesonix
Raf., Tellima R.Br., and Tolmiea T. & G.

Hydrangeaceae: Whipplea Torr.

Rosaceae: Chamaebatiaria (Porter) Maxim., Kelseyva (Wats.) Rydb.,
Luetkea Bong., Osmaronia Greene, Peraphyllum Nutt., Petrophytum
(Nutt.) Rydb., and Purshia DC.

Umbelliferae: Orogenia Wats., Perideridia Reichenb., RhAysopterus
Coult. & Rose, and Sphenosciadium Gray.

Cactaceae: Pediocactus Britt. & Rose.

Malvaceae: Sidalcea Gray.

Euphorbiaceae: Eremocar pus Benth.

1969] SCHOFIELD: PHYTOGEOGRAPHY 159

Ericaceae: Allotropa Torr. & Gray, Cladothamnus Bong., Hemitomes
Gray, Pitvopus Small, and Pleuricospora Gray.

Primulaceae: Douglasia Lindb.

Polemoniaceae: Eriastrum Woot. & Standl., Gymnosteris Greene,
Langloisia Greene Leptodactylon H. & A., and Linanthastrum Ewan.

Hydrophyllaceae: Ellisia L., Hesperochiron Wats., and Romanzo fia
Cham.

Boraginaceae: Coldenia L., and Dasynotus Johnst.

Scrophulariaceae: Chionophila Benth., Cordylanthus Nutt., Mimetan-
the Greene, Synthyris Benth., and Tonella Nutt.

Valerianaceae: Plectvitis DC.

Campanulaceae: Downingia Torr., Githopsis Nutt., Heterocodon Mitt.,
Howellia Gray, and Porterella Torr.

Compositae: A pargidium T. & G., Bahia, Laq., Balsamorhiza Nutt.,
Blepharipappus Hook., Chaenactis DC., Chrysothamnus Nutt., Cro-
cidium Hook., Dimeresia Gray, Eatonella Gray, Enceliopsis (Gray)
Nels., Eviophyllum Laq., Hulsea Torr. & Gray, Luinia Benth., Madia
Mol., Railardella Benth., Rigiopappus Gray, and W yethia Nutt.

More than half of these genera are monotypic. Many are widespread,
Camassia, Sarcobatus, Eschscholtzia, Lewisia, Romanzo fia, Lithophrag-
ma, Luetkea, Apargidium, etc., where others, Scribneria, Chlorocambe,
Darlingtonia, etc., are highly restricted in their range. Of particular in-
terest is the fact that there are no endemic genera of Pteridophytes or
Gymnosperms. A number of families are especially rich in endemic
genera: Cruciferae (13 genera), Saxifragaceae (19 genera), Rosaceae
(7 genera) and Compositae (17 genera). Many of these genera are re-
stricted to rather arid regions, although a number are of alpine and sub-
alpine habitats and others are of forests at lower elevations. As in the
bryophytes, most of the genera are clearly distinct from related genera.

Since there is such a richness of endemic species, these are treated
here in relation to thir distribution pattern in the region.

SPECIES ENDEMISM

Widespread at Elevations Below Subalpine

These species are conspicuous elements of both flora and vegetation.
The vascular plants give the vegetation its physiognomic character and
the bryophytes tend to dominate that flora in other strata. Within such
a wide range many species are environmentally restricted. For example,
some bryophytes are consistently on perennially dry and exposed rock
surfaces while others are confined to rocks perennially inundated. The
forest is entirely of endemic trees and mainly of endemic shrubs, and
the herbaceous vegetation is dominated by endemic species. Most of
the species do not extend beyond the crests of the Rocky Mountains and
many disappear with the boundary to the boreal coniferous forest of
northern latitudes or the arid portions of the interior regions. Thus the

160 MADRONO [Vol. 20

widespread element is found in regions of relatively high precipitation
on both the coast and lower elevations of the coastal mountains and also
at lower elevations of the mountains west of the Rockies. Many of the
species extend southward along the coast as far south as the southern
limits of the coastal redwood forest (Sequoia sempervirens (Don) Endl.)
in California and occasionally at lower elevations of the Sierra Nevada.

Within this same geographic area are other floristic elements; these
are discussed elsewhere in this paper. Their phytogeographic significance
is great, but their contribution to the vegetational cover is generally
smaller than that of the endemic species, particularly in the vascular
flora.

The distributions of the hepatics are not well understood since collec-
tion has been rather limited: Gyrothyra underwoodiana Howe., Plecto-
colea rubra (Gottsche) Evans, Scapania americana Miull., Bazzania am-
bigua (Lindenb.) Trevis., Radula bolanderi Gottsche, Porella roelliu
Steph., P. navicularis (Lehm. & Lindenb.) Lindb., Frullania nisquallen-
sis Sull., and F. franciscana Howe.

Among the mosses the details are somewhat clearer and the represen-
tation of endemic taxa is somewhat greater: Sphagnum mendocinum
Sull. & Lesq., Atrichum selwynii Aust., Pogonatum macouni (Kindb.)
Kindb. & Mac., Ditrichum ambiguum Best, D. schimperi (Lesq.) Kunze,
Dicranella n. sp., Amphidium californicum (Hampe) Broth., Dicranum
howellt Ren. & Card., Barbula rubiginosa Mitt., Scouleria aquatica
Hook., Racomitrium depressum Lesq., R. varium (Mitt.) Lesq. & James,
Pohlia longibracteata Broth., Leucolepis menziest (Hook.) Steere,
Plagiomnium insigne (Mitt.) Koponen, P. venustum (Mitt.) Koponen,
Rhizomnium glabrescens (Kindb.) Koponen, Ulota megalospora Vent.,
U. obtusiuscula Mull. & Kindb., Orthotrichum consimile Mitt., Fonti-
nalis neomexicana Sull. & Lesq., F. patula Card., Dichelyma uncinatum
Mitt., Neckera douglasit Hook., Porotrichum bigelovit (Sull.) Kindb.,
Thamnobryum leibergu (Britt.) Nieuwl., Jsothecium stoloniferum
(Hook.) Brid., Claopodium bolanderit Best, Homalothecium fulgescens
(Mitt.) Lawt. Brachythecium frigidum (Miull.) Besch., B. lampro-
chryseum Mull. & Kindb., Eurhynchium oreganum (Sull.) Jaeg. &
Sauerb., Scleropodium obtusifolium (Hook.) Kindb., and Hypnum cir-
cinale Hook.

These bryophytes occupy a diversity of habitats but the majority are
epiphytic on tree trunks and occur less commonly on rock. The remain-
der occupy various habitats, from splashed rock faces, for example
Scouleria aquatica and Scleropodium obtusifolium; humid cliff bases,
on rock or on soil, Pogonatum macounu, Pohlia longibracteata, Poro-
trichum bigelovii, and Thamnobryum leibergu; rotten logs or forest
floor, Dicranum howellii and Brachythecium frigidum ; or aquatic Sphag-
num mendocinum and Fontinalis neomexicana.

A number of species that have a wide range but are rare are:

1969] SCHOFIELD: PHYTOGEOGRAPHY 161

Hepatics: Blepharostoma arachnoideum Howe, Jungermannia alleni
Clark, Plectocolea rubra (Gottsche.) Evans, and Sphaerocarpos hians
Haynes.

Mosses: Fissidens ventricosus Lesq., F. pauperculus Howe, Crumia
latifolia (Kindb.) Schof., Scouleria marginata Britt., Brotherella roelliu
(Ren. & Card.) Fleisch., and Triperocladium leucocladulum (Mill.)
Kindb.

Based on their present ecology and distribution it can be inferred that
during glaciation, the bulk of these species persisted largely south of
the glacial boundary, probably in forested environments. The relative
scarcity of most of them in unglaciated Alaska indicates that they did
not persist there. A number of these species have probably increased in
abundance in recent times, largely with the expansion of the available
habitat, chief among these are Gyrothyra underwoodiana, an unde-
scribed Dicranella, and Pohlia longibracteata, all of which frequently
abound on moist road-cuts. Many others have probably been greatly
restricted by the elimination of their forest habitat, although gross dis-
tributional pattern probably has not been greatly altered.

The vascular flora of this widespread distribution is large. The woody
elements of this flora characterize the vegetation. Since these taxa are
endemic, the impression of great endemism results in spite of the very
considerable number of circumboreal and circumpolar species that make
up the total flora. The following list is far from complete, but will serve
to illustrate the diversity of taxa present:

Polypodiaceae: Polystichum munitum (Kaulf.) Presl, Polypodium
glycyrhiza Eat., and P. hespertum Maxon.

Taxaceae: Taxus brevifolia Nutt.

Pinaceae: Pinus contorta Dougl., Picea sitchensis (Bong.) Carr.,
Pseudotsuga menziesu (Mirb.) Franco, Tsuga heterophylla (Raf.) Sarg.,
and Abies grandis (Dougl.) Lindl.

Cupressaceae: Chamaecyparis nootkatensis (Lamb.) Spach.

Juncaceae: Juncus brachyphyllus Wieg. and J. oreganus Wats.

Liliaceae: Clintonia uniflora (Schult.) Kunth, Disporum hookert
(Torr.) Nichols, Erythronium oreganum Appleg., E. revolutum Sm.,
Trillium ovatum Pursh, Veratrum californicum Durand, and Zygadenus
elegans Pursh.

Orchidaceae: Cypripedium montanum Dougl. and Listera caurina
Piper.

Poaceae: Agropyron spicatum (Pursh.) Scribn. & Sm., Agrostis
aequivalvis Trin., A. diegoensis Vasey, A. idahoensis Nash., A. micro-
phylla Steud., Bromus pacificus Shear, Elymus innovatus Beal, Festuca
idahoensis Elmer, F. subulata Trin., Glyceria elata (Nash.) Hitchc.,
Melica subulata (Griseb.) Scribn., Poa laxiflora Buckl., P. stenantha
Trin., and Trisetum cernuum Trin.

162 MADRONO [Vol. 20

Cyperaceae: Carex atrostachya Olney, C. preslw Steud., C. phaeoce-
phala Piper, C. petasata Desv., C. microptera Mack., C. laeviculmis
Meinsch., C. phyllomanica Boott., C. scopulorum Holm, C. kelloggii
Boott., and C. sitchensis Prescott.

Araceae: Lysichiton americanum Hult. & St. John.

Salicaceae: Salix lasiandra Benth., S. scouleriana Benth., and S. sit-
Chensis Sanson.

Betulaceae: Alnus rhombifolia Nutt. and A. rubra Bong.

Aristolochiaceae: Asarum caudatum Lindl.

Portulacaceae: Montia parvifolia (Moc.) Greene and M. sibirica (L.)
Howell.

Caryophyllaceae: Silene menziesu Hook.

Ranunculaceae: Aconitum columbianum Nutt., Aquilegia formosa
Fisch., Coptis asplenifolia Salisb., Ranunculus alismaefolius Geyer, R.
occidentalis Nutt., and Thalictrum occidentale Gray.

Berberidaceae: Berberis aquifolium Pursh.

Saxifragaceae: Boykinia elata (Nutt.) Greene, Heuchera cvylindria
Dougl., H. glabra Willd., H. micrantha Dougl., Mitella trifida Grah.,
Saxifraga ferruginea Grah., Tellima grandiflora (Pursh.) Dougl., Tia-
rella trifoliata L., Tolmiea menziesu (Pursh.) T. & G., Ribes bracteosum
Dougl., and R. laxiflorum Pursh.

Rosaceae: Holodiscus discolor (Pursh.) Maxim., Osmaronia cerasi-
formis (T. & G.) Greene, Physocarpus capitatus (Pursh.) Kuntze,
Potentilla glandulosa Lindl., P. gracilis Dougl., Prunus emaginata
(Dougl.) Walpers, Pyrus fusca Raf., Rosa gymnocarpa Nutt., R. nut-
kana Presl., Rubus lastococcus Gray, R. leucodermis Dougl., R. nivalis
Dougl., and Spiraea douglas Hook.

Leguminosae: Lathyrus nevadensis Wats., Lotus purshianus ( Benth.)
Clements & Clements, Lupinus lepidus Dougl., and L. polyphyllus Lindl.

Oxalidaceae: Oxalis oregana Nutt.

Celastraceae: Pachystima mysinites (Pursh) Raf.

Aceraceae: Acer circinatum Pursh and A. glabrum Torr.

Balsaminiaceae: /mpatiens ecalcarata Blank.

Rhamnaceae: Ceanothus sanguineus Pursh and Rhamnus purshiana
DC.

Hypericaceae: Hypericum anagalloides C. &S.

Violaceae: Viola purpurea Kell.

Onagraceae: Boisduvalia densiflora (Lindl.) Wats., Epilobium glaber-
rimum Barbey, E. luteum Pursh, and E. minutum Lindl.

Cornaceae: Cornus nuttallu Aud.

Ericaceae: Allotropa virgata T. & G., Chimaphila menziesii (R.Br.)
Spreng., Gaultheria shallon Pursh, Menziesia ferruginea Sm., Pleuricos-
pora fimbriolata Gray, Pyrola aphylla Sm., P. dentata Sm., P. picta Sm.,
Vaccinium alaskaense Howell, and V. parvifolium Sm.

Primulaceae: Dodecatheon jeffreyi van Houtte.

1969] SCHOFIELD: PHYTOGEOGRAPHY 163

Gentianaceae: Gentiana sceptrum Griseb. and G. douglasiana Bong.

Convolvulaceae: Cuscuta occidentalis Millspaugh.

Hydrophyllaceae: Romanzo fia sitchensis Bong.

Caprifoliaceae: Lonicera ciliosa (Pursh) DC.

Campanulaceae: Heterocodon rariflorum Nutt.

Compositae: Agoseris grandiflora (Nutt.) Greene, Antennaria ana-
phaloides, Rydb., A. corymbosa Nels., A. dimorpha (Nutt.) T. & G.,
Arnica amplexicaulis Nutt., A. diversifolia Greene, A. latifolia Bong.,
Aster subspicatus Nees, Microseris laciniata (Hook.) Schultz-Bip., and
Prenanthes alata (Hook.) Dietr.

Subalpine and Alpine

Besides possessing a flora containing rich representation of circum-
polar species, the mountains of northwestern North America have many
bryophyte and vascular plant endemics. The woody flora is essentially
endemic but not confined to the mountains while endemism decreases
in the herbs and bryophytes. The subalpine forest probably possesses
more endemic bryophytes than the alpine portion, but in the vascular
flora endemism increases in alpine areas. While some mountains serve
as islands of endemism for vascular plants, the bryophytes are not so
confined. This is in spite of very narrow environment restriction of
many of them. Most of these bryophytes produce numerous sporophytes
annually although several are dioicous. Special vegetative reproductive
organs are not known for any of the endemic alpine bryophytes.

In the following discussion the widespread subalpine and alpine spe-
cies are treated first and various mountains are noted with their endemic
floras.

1. Widespread subalpine and alpine

Hepatics: Macrodiplophyllum imbricatum (Howe) Perss.

Mosses: Oligotrichum parallelum (Mitt.) Kindb., Polytrichadelphus
lyallii Mitt., Buxbaumia piperi Best, Ditrichum montanum Leib., Tre-
matodon boast Schof., Dicranoweisia roella Kindb., Dicranum pallidi-
setum (Bailey) Irel., Grimmia atricha Mull. & Kindb., Pohlia columbica
(Kindb.) Andr., Roellia roellu (Broth.) Crum, Lescuraea baileyi (Best
& Grout) Lawt., ZL. atricha (Kindb.) Lawt., L. stenophylla (Ren. &
Card.) Kindb., Heterocladium procurrens (Mitt.) Rau. & Herv., Hygro-
hypnum besti (Ren. & Bryhn.) Holz., Trachybryum megaptilum (Sull.)
Schof., Brachythecium leibergii Grout, B. hylotapetum Hig. & Hig.,
and Rhytidiopsis robusta (Hook.) Broth.

Vascular Plants:

Polypodiaceae: Pellaea bridgesii Hook.

Pinaceae: Larix occidentalis Nutt., L. lyallii Parl., Pinus albicaulis
Engelm., P. flexilis James, P. monticola Dougl., Tsuga mertensiana

164 MADRONO [Vol. 20

(Bong.) Sarg., Abzes amabilis (Dougl.) Forbes, A. lastocarpa (Hook.)
Nutt., and Picea engelmannii Parry.

Cupressaceae: Juniperus occidentalis Hook.

Juncaceae: Juncus drummondii Mey., J. mertensianus Bong., J. parryi
Engelm., and J. regelit Buch.

Cyperaceae: Carex anthoxanthea Presl., C. circinnata Mey., C. nigri-
cans Mey., C. albonigra Mack., C. atrata L., C. mertensi Prescott, and
C. petricosa Desv.

Liliaceae: Allium validum Wats., Erythronium grandiflorum Pursh,
E. montanum Wats., Lilium columbianum Hanson, Stenanthium occi-
dentale Gray, and Xerophyllum tenax (Pursh) Nutt.

Orchidaceae: Cypripedium montanum Dougl.

Poaceae: Agrostis humilis Vasey, A. thurberiana Hitchc., A. variabilis
Rydb., Bromus sitchensis Trin., B. suksdorfi Vasey, Calamagrostis
tweedw (Scribn.) Scribn., Festuca viridula Vasey, Melica spectabilis
Scribn., Oryzopsis exigua Thurb., Poa bolanderi Vasey, P. curta Rydb.,
P. curtifolia Scribn., P. gracillima Vasey, P. grayana Vasey, P. letter-
mani Vasey, P. nervosa (Hock.) Vasey, P. reflexa Vasey & Scribn., and
P. suksdorfu (Beal) Vasey.

Salicaceae: Salix barclayi Anderss., S. barrattiana Hook., S. cascaden-
sis Cockerell, S. dodgeana Rydb., S. geyveriana Anderss., S. nivalis Hook.,
S. tweedyi (Bebb) Ball, and S. wolfiz Bebb.

Polygonaceae: Eriogonum androsaceum Benth., E. chrysops Rydb., E.
pyrolifolium Hook., Polygonum bistortoides Pursh, P. minimum Wats.,
P. newberryt Small, P. phytolaccifolium Meisn., and Rumex paucifolius
Nutt.

Caryophyllaceae: Silene parryi (Wats.) Hitchc. & Maguire, S. scaposa
Robins., S. scoulert Hook., and Stellaria jamesiana Torr.

Ranunculaceae: Aquilegia jonesi Parry, Caltha biflora D.C., C. lep-
tosepala D.C., Delphinium glareosum Greene, D. glaucum Wats., D.
occidentale Wats., Ranunculus cardiophyllus Hook., R. cooleyae Vasey
& Rose, R. eschscholtzi Schlecht., R. inamoenus Greene, and R. vere-
cundus Robins.

Papaveraceae: Papaver pygmaeum Rydb.

Fumariaceae: Dicentra uniflora Kell.

Cruciferae: Anelsonia eurycarpa (Gray) Macbr. & Pays., Arabis
furcata Wats., A. lyallii Wats., A. microphylla Nutt., A. platyperma
Gray, Cardamine breweri Wats., Chlorocrambe hastata (Wats.) Rydb.,
Draba apiculata Hitche., D. aurea Vahl., D. crassifolia Nutt., D. densi-
folia Nutt., D. incerta Pays., D. lonchocarpa Rydb., D. paysonii Macbr.,
D. praealia Greene, D. stenoloba Ledeb., and D. ventosa Gray.

Crassulaceae: Sedum oregonense (Wats.) Peck.

Saxifragaceae: Elmera racemosa (Wats.) Rydb., Leptarrhena pyroli-
folia (Don) R. Br., Mitella brewert Gray, Parnassia fimbriata Konig.,
Saxifraga arguta Don, S. chrysantha Gray, S. debilis Engelm., S. occi-

1969] SCHOFIELD: PHYTOGEOGRAPHY 165

dentalis Wats., S. oregana Howell, S. tolmiei T. & G., Telesonix jamesi
(Torr.) Raf., Ribes howellii Greene, R. mogoelonicum Greene, and R.
montigenum McClatchie.

Rosaceae: /vesia gordoni (Hook.) T. & G., 7. tweedyi Rydb., Kelseya
uniflora (Wats.) Rydb., Luetkea pectinata (Pursh) Kuntze, Potentilla
brevifolia Nutt., P. drummondii Lehm., P. flabellifolia Hook., P. hookert-
ana Lehm., Rubus pedatus Sm., Sanguisorba sitchensis Meyer, and
Spiraea densiflora Nutt.

Leguminosae: Astragalus cottonii Jones, A. tegetarius Wats., A. whit-
neyi Gray, Hedysarum occidentale Greene, Oxytropis parryi Gray, Tri-
folium beckwithii Brew., T. dasyphyllum T. & G., T. nanum Torr., and
T. parryi Gray.

Haloragidaceae: Hippuris montana Ledeb.

Umbelliferae: Angelica roseana Henderson.

Ericaceae: Cassiope mertensiana (Bong.) Don, Gaultheria humifusa
(Grah.) Rydb., G. ovatifolia Gray, Phyllodoce empetriformis (Sw.)
Don, Rhododendron albiflorum Hook., and Vaccinium membranaceum
Dougl.

Gentianaceae: Gentiana calycosa Griseb.

Polemoniaceae: Polemonium elegans Greene.

Boraginaceae: Cryptantha nubigena (Greene) Pays.

Scrophulariaceae: Castilleja applegate: Fern., C. parviflora Bong., C.
rhexifolia Rydb., Mimulus lewisti Pursh, Pedicularis bracteosa Benth.,
P. contorta Benth., P. cvstopteridifolia Rydb., P. ornithorhyncha Benth.,
and Pentstemon davidsoni Greene.

Valerianaceae: Valeriana acutiloba Rydb.

Compositae: Antennaria lanata (Hook.) Greene, A. mollis Hook.,
A. nevadensis Gray, Arnica michauxiana Bess., A. scopulorum Gray,
A. alpigenus (T. & G.) Gray, Chaenactis alpina (Gray) Jones, Erigeron
asperugineus (Eat.) Gray, E. lanatus Hook., E. leiomerus Gray, E. sim-
plex Greene, EF. ursinus D. C. Eat., E. vagus Payson, Haplopap pus lyallii
Gray, H. pygmaeus (T. & G.) Gray, Hulsea algida Gray, Saussurea
americana Eat., Senecio megacephalus Nutt., S. subnudus DC., and S.
werneriufolius Gray.

2. Anumber of mountain areas posses their endemic species

a. Rocky Mountains (mainly)

Vascular Plants: Juncus halli Engelm., J. tweedyi Rydb., Allium
brevistylum Wats., Draba crassa Rydb., Sedum debile Wats., Conimi-
tella williamsu (Eat.) Rydb., Trifolium haydenu Porter, Primula parryi
Gray, Phacelia lyallu (Gray) Rydb., Synthyris canbyi Pennell, Cirsium
tweedyi Rydb., Erigeron pallens Crong., Hymenoxys grandiflora (T. &
G.) Parker, and Townsendia spathulata Nutt. )

b. Cascade Mountains (principally)

Mosses: Pohlia cardoti (Ren.) Broth.
Vascular Plants: Silene suksdorfit Robins., Draba aureola Wats.,

166 MADRONO [Vol. 20

Physaria alpestris Suksd., Smelowskia ovalis Jones, Tauschia strick-
landu (Coult. & Rose) Math. & Const., Castilleja cryptantha Greenm..,
C. rupicola Piper, C. suksdorfii Gray, Pedicularis rainierensis Pennell &
Warren, Aster gormanu (Piper) Blake, Erigeron cascadensis Heller,
Hulsea nana Gray, Luinia nardosmia (Gray) Cronq. and L. stricta
(Greene) Rob.

c. Olympic Mountains

Vascular Plants: Petrophytum hendersoniu (Canby) Rydb., Viola
flettu Piper, Campanula pipert Howell, Aster paucicapitatus Rob., Eri-
geron flettu Jones, and Senecio websteri Greenm.

d. Cascade Mountains, Coast and Insular Mountains, and
Olympic Mountains
Mosses: Dichodontium olympicum Ren. & Card. and Grimmia olym-
pica Britt.
Vascular Plants: Delphinium glareosum Greene, Erysimum arenicola
Wats., Smelowskia divergens Wats., Vaccinium deliciosum Piper, Arnica
nevadensis Gray, and Senecio fletti Wieg.

e. Wenatchee Mountains

Vascular Plants: Silene seelyi Morton & Thompson, Delphinium viri-
descens Leiberg, D. xantholeucum Piper, Lomatium cuspidatum Math.
& Const., Valeriana columbiana Piper, Chaenactis ramosa Stockwell, and
C. thom psonu Cronq.

f. Wallowa Mountains (sometimes also in Blue Mountains)

Vascular Plants: Lomatium greenmani Mathias, L. oreganum Coult.
& Rose, Castilleja chrysantha Greenm., C. fraterna Greenm., C. glanduli-
fera Pennell, C. owenbeyana Pennell, C. rubida Piper, Pentstemon
spathulatus Pennell, and Senecio porteri Greene.

There are numerous other subalpine and alpine endemics, in a number
of cases of very restricted distribution. The majority of the endemics
are of circumpolar genera and many are especially rich in species, for
example, Salix, Arabis, Draba, Saxifraga, Trifolium, Castilleja, Pedicu-
laris, Erigeron, and Senecio. Although many of these genera are notor-
iously polymorphic, the endemic species tend to be remarkably distinct.
Since many of the species are ecologically restricted, their discovery is
often by chance, and thus their total distribution through mountainous
western North America is not thoroughly known. Considerable botanical
exploration even in presumably well-known mountain areas, remains
to be done.

Dry Interior Plans

East of the Cascade and Coastal Mountains and west of the Rocky
Mountains there extends a lowland trough lying in the rain shadow of
the coastward mountains. This drier region possesses a vegetation that

1969] SCHOFIELD: PHYTOGEOGRAPHY 167

is composed predominantly of species endemic to western North America.
Many range southward into the cold deserts and some even to the
warmer arid regions as far south as Mexico. Many are found also east
of the Rocky mountains and the northern limits are largely in central
British Columbia, although occasionally some species extend into Yukon
and Alaska. Endemism is highest in perennial herbs although some are
woody or annual. No endemic hepatics have been reported and few
mosses, although little careful bryological exploration has been made
in this area.

Bryophytes: Barbula andreaeotdes Kindb., B. platyvneura Mull. &
Kindb., Pottia nevadensis Card. & Thér., Grimmia calyptrata Hook.,
Funaria americana Lindb., and Orthotrichum halli Sull. & Lesq.

Vascular Plants

Pinaceae: Pinus ponderosa Dougl. and P. flexilis James.

Lilliaceae: Allium nevadense Wats. Fritillaria pudica (Pursh) Spreng.,
and Leucocrinum montanum Nutt.

Iridaceae: [ris chrysophylla Howell and Calochortus bruneaunis Nels.
& Macbr.

Poaceae: Danthonia parryi Scribn., D. unispicata Munro, Melica bul-
bosa Geyer, M. fugax Boland., Muhlenbergia andina (Nutt.) Hitchc.,
Stipa letterman Vasey, S. thurberiana Piper, and Trisetum wolf Vasey.

Ulmaceae: Celtis douglasi Planch.

Polygonaceae: Chorizanthe brevicornu Torr., C. watsoni T. & G.,
Eriogonum angulosum Benth., E. caespitosum Nutt., FE. cernuum Nutt.,
E. chrysocephalum Gray, E. deflexum Torr., FE. douglas Benth., E.
elatum Dougl., E. heracleoides Nutt., LE. microthecum Nutt., E. niveum
Dougl., E. sphaerocephalum Dougl., E. thyvmoides Benth., and Poly-
gonum austiniae Greene.

Chenopodiaceae: Atriplex truncata (Torr.) Gray, Grayia spinosa
(Hook.) Moq., Kochia americana Wats., Monolepis pusilla Torr., M.
spathulata Gray, Nitrophila occidentalis (Moq.) Wats., Salicornia rubra
Nels., Sarcobatus vermiculatus (Hook.) Torr., Suaeda intermedia Wats.,
and S. spaldingii Wats.

Amaranthaceae: Amaranthus californicus (Moq.) Wats.

Portulacaceae: Calyptridium roseum Wats., Lewisia rediviva Pursh,
and Talinum spinescens Torr.

Caryophyllaceae: Avenaria aculeata Wats., A. franklinii Dougl., A.
hookeri Nutt., A. pusilla Wats., Silene douglasii Hook., S. oregana Wats.,
and S. spaldingiit Wats.

Paeoniaceae: Paeonia brownii Dougl.

Ranunculaceae: Clematis hirsutissima Pursh, C. ligusticifolia Nutt.,
Delphinium andersonii Gray, D. depauperatum Nutt., D. glaucescens
Rydb., D. multiplex (Ewan) Hitchc., D. stachyvdeum (Gray) Nels. &
Macbr., Myosurus aristatus Benth., Ranunculus andersonii Gray, R.
jovis Nels., and R. reconditus Nels. & Macbr.

168 MADRONO [Vol. 20

Papaveraceae: Canbya aurea Wats.

Cruciferae: Avabis cobrensis Jones, A. cusickii Wats., A. lignifera
Nels., Caulanthus crassicaulis (Torr.) Wats., C. pilosus Wats., Draba
douglasu Gray, Erysimum occidentale (Wats.) Robins., /dahoa scapigera
(Hook.) Nels. & Macbr., Lepidium dictyotum Gray, Lesquerella doug-
lasu Wats., L. kingit Wats., Phoenicaulis cheiranthoides Nutt., Physaria
didymocarpa (Hook.) Gray, Polyctenium fremontii (Wats.) Greene,
Schoenocrambe linifolia (Nutt.) Greene, Stanleya tomentosa Parry, S.
viridifolia Nutt., Streptanthella longirostris (Wats.) Rydb., Thely-
podium integrifolium (Nutt.) Endl., and 7. sagittatum (Nutt.) Endl.

Saxifragaceae: Lithophragma parviflora (Hook.) Nutt., L. tenella
Nutt., Rzbes aureum Pursh, and R. velutinum Greene.

Rosaceae: Cercocarpus ledifolius Nutt., Chamaebatiaria millefolium
(Torr.) Maxim., Holodiscus dumosus (Hook.) Heller, Peraphyllum
ramosissimum Nutt., and Purshia tridentata (Pursh) D.C.

Leguminosae: Astragalus adanus Nels., A. argophyllus Nutt., A. ar-
thuru Jones, A. atratus Wats., A. calycosus Torr., A. casez Gray, A.
cibarius Sheld., A. collinus Dougl., A. convallarius Greene, A. curvicar pus
(Sheld.) Macbr., A. cusickiit Gray, A. filipes Torr., A. geyert Gray, A.
howell Gray, A. inflexus Dougl., A. leibergi Jones, A. lyallii Gray, A.
malacus Gray, A. microcystis Gray, A. newberryi Gray, A. nudisiliquus
Nels., 4. obscurus Wats., A. palousensis Porter, A. reventus Gray, A.
salmonis Jones, A. scaphoides Jones, A. sinuatus Piper, A. spaldingu
Gray, A. speirocarpus Gray, A. stenophyllus T. & G., A. succumbens
Dougl., A. tegetarioides Jones, A. toanus Jones, A. tweedyi Canby, A.
umbraticus Sheld., Lathyrus lanszwertiu Kell., L. pauciflorus Fern., L.
rigidus White, Lupinus caudatus Kell., L. holosericeus Nutt., L. laxiflorus
Dougl., L. sabini Dougl., L. saxosus Howell, L. wyethiu Wats., Oxytro pis
lagopus Nutt., Petalostemon ornatum Dougl., Trifolium gymnocarpon
Nutt., 7. macrocephalum Pursh, and T. thom psoni Morton.

Malvaceae: /liamna longisepala (Torr.) Wiggins, Sidalcea neomext-
cana Gray, S. oregana (Nutt.) Gray, Sphaeralcea grossularufolia (H. &
A.) Rydb., and S. munroana (Dougl.) Spach.

Violaceae: Viola beckwithi T. & G. and V. trinervata Howell.

Loasaceae: Mentzelia albicaulis Dougl., M. dispersa Wats., and M.
laevicaulis (Dougl.) T. & G.

Cactaceae: Pediocactus simpsonu (Engelm.) Britt. & Rose.

Onagraceae: Oenethera alyssoides H. & A., O. andina Nutt., O. boothu
Dougl., O. claviformis Torr. & Frem., O. deltoides Torr. & Frem., O.
minor (Nels.) Munz, O. palmeri Wats., O. scapoidea Nutt., and O.
tanacetifolia T. & G.

Umbelliferae: Lomatium canbyi Coult. & Rose, L. farinosum (Hook.)
Coult. & Rose, L. gormanii (Howell) Coult. & Rose, L. hambleniae
Math. & Const., ZL. nudicaule (Pursh) Coult. & Rose, L. watsoni Coult.
& Rose, and Tauschia hooverit Math. & Const.

Gentianaceae: Frasera montana Mulford.

1969] SCHOFIELD: PHYTOGEOGRAPHY 169

Polemoniaceae: Gilia minutiflora Benth., Gymnosteris nudicaulis (H.
& A.) Greene, G. parvula Heller, Linanthus pharnaceoides ( Benth.)
Greene, Phlox aculeata Nels., and P. caespitosa Nutt.

Hydrophyllaceae: Hesperochiron californicus (Benth.) Wats., Nama
aretioides (H. & A.) Brand, N. densum Lemmon, and Phacelia bicolor
Torr.

Boraginaceae: P. glandulifera Piper, Cryptantha scoparia Nels., C.
simulans Greene, Hackelia arida (Piper) Johnst., H. ciliata (Dougl.)
Johnst., H. patens (Nutt.) Johnst., Pectocarya setosa Gray, and Plagio-
bothrys harknessii (Greene) Nels & Macbr.

Scrophulariaceae: Castilleja angustifolia (Nutt.) Don, C. cervina
Greenm., C. chromosa Nels., C. exilis Nels., C. flava Wats., C. inverta
(Nels. & Macbr.) Pennell & Ownbey, C. linarifolia Benth., C. longis pica
Nels., C. lutescens (Greenm.) Rydb., C. oresbia Greenm., C. pallescens
(Gray) Greenm., C. rustica Piper, C. thompsoniu Pennell, C. xantho-
tricha Pennell, Cordyvlanthus capitatus Nutt., C. ramosus Nutt., Mimu-
lus cusicku (Greene) Piper, Orthocarpus barbatus Cotton, Pentstemon
acuminatus Dougl., P. barrettiae Gray, P. cinicola Keck, P. cusicku
Gray, P. cvaneus Pennell, P. gairdneri Hook., P. humilis Nutt., P. laetus
Gray, P. lemhiensis (Keck) Keck & Cronq., P. peckit Pennell, P. pumilus
Nutt., P. radicosus Nels., P. rydbergi Nels., P. seorsus (Nels.) Keck,
and P. speciosus Dougl.

Orobanchaceae: Orobanche californica S.&S.

Compositae: Antennaria geyert Gray, Artemisia tridentata Nutt.,
A. tripartita Rydb., Brickellia microphylla (Nutt.) Gray, B. oblongifolia
Nutt., Carsium magnificum (Nels.) Petr., C. utahense Petr., Eatonella
nivea (Eat.) Gray, Erigeron aphanactis (Gray) Greene, E. chrysopsidis
Gray, E. linearis (Hook.) Piper, EF. piperianus Cronq., E. poliospermus
Gray, Haplopappus stenophyllus Gray, Madia minima (Gray) Keck,
Rigiopappus leptocladus Gray, Stephanomeria exigua Nutt., and S.
lactucina Gray.

Californian

A distinctive element in the flora of southwestern British Columbia
occupies the so-called ‘“‘Mediterranean”’ climatic portion of Southern
Vancouver Island, the islands of the southern Strait of Georgia and the
headlands of the adjacent mainland. The species occupy sites that are
edaphically similar to those occupied by the same taxa further south to
California in more conspicuously Mediterranean climates, and where
they are more widespread. All species are restricted to west of the
Cascade Mountains, occupy drier sites, but are not maritime. This ele-
ment possibly extended its range northward from California or Oregon
to southern British Columbia during the Hypsithermal Interval and
fragments persist only in edaphically suitable sites although the general
climate of the region is unfavourable.

170 MADRONO [Vol. 20

It is equally possible that the species have entered the region by
expanding their range stepwise via the available edaphically suitable
sites, and no Hypsithermal Interval need be involved as an initiating
cause. The element is conspicuous both in the bryoflora and vascular
flora and is represented by both western North American endemics and
by species of wider world distribution, but whose restriction is essen-
tially to Mediterranean climates. This element is discussed briefly by
Schofield (1965; 1968a; 1968b), and Ireland and Schofield (1967).

Hepatics: Fossombronia longiseta Aust. and Frullania californica
(Aust.) Evans.

Hornworts: Anthoceros hallii Aust.

Mosses: Fissidens ventricosus Lesq., Ditrichum ambiguum Best, Pleur-
idium bolanderi Mull., Timmiella crassinervis (Hampe) Koch, Tor-
tula amplexa (Lesq.) Steere, T. bolanderi (Lesq.) Howe, Physcomitrium
megalocarpum Kindb., Ptvchomitrium gardneri Lesq., Orthotrichum
papillosum Hampe, Pseudobraunia californica (Lesq.) Broth., Alsza
californica (Hook. & Arnott.) Sull., Dendroalsia abietina (Hook.) Britt.,
Bestia vancouveriensis (Kindb.) Wijk. & Marg., [sothecium cristatum
(Hampe) Robins., Homalothecium nuttallu (Wils.) Jaeg. & Sauerb., A.
pinnatifidum (Sull. & Lesq.) Lawt., and H. arenarium (Lesq.) Lawt.

Vascular Plants: Carex brevicaulis Mack., Juncus bolanderit Engelm.,
Brodiaea congesta Smith, Allium crenulatum Wieg., Disporum smithiu
(Hook.) Piper, Erythronium oreganum Appleg., E. revolutum Smith,
Sisyrinchium douglasii Dietr., Habenaria elegans Lindl., Poa confinis
Vasey, Quercus garryana Dougl., Montia diffusa (Nutt.) Greene, Del-
phinium menziesii D.C., Ranunculus lobbiu (Hiern) Gray, Berberis ner-
vosa Pursh, Meconella oregana Nutt., Corydalis scoulerti Hook., Ribes
sanguineum Pursh, Rosa pisocarpa Gray, Lotus micranthus Benth.,
Lupinus bicolor Lindl., Trifolium oliganthum Steud., Rhus diversiloba
T. & G,, Viola howellii Gray, V. sempervirens Greene, Clarkia amoena
(Lehm.) Nels. & Macbr., Lomatium utriculatum (Nutt.) Coult. & Rose,
Arbutus menziesii Pursh, Arctostaphylos columbiana Piper, Vaccinium
ovatum Pursh, Dodecatheon hendersoni Gray, Navarretia squarrosa
(Esch.) H. & A., Hydrophyllum tenuipes Heller, Amsinckia spectabilis
F.& M., Castilleja levisecta Greenm., Mimulus alsinoides Dougl., Ortho-
carpus attenuatus Gray, O. pusillus Benth., Galium cymosum Wieg.,
Plectritis congesta (Lindl.) D.C., Valeriana scoulert Rydb., Balsamorhiza
deltoidea Nutt., Madia madioides (Nutt.) Greene, Microseris bigelovit
(Gray) Schultz-Bip., and Senecio macounii Greene.

Other vascular plants, probably of the same element, extend north-
ward to the Puget Sound area in Washington, and occur southward to
California between the coastal mountains and the Cascades. The follow-
ing are representative: Castanopsis chrysophylla (Dougl.) D.C., Are-

1969] SCHOFIELD: PHYTOGEOGRAPHY 171

naria paludicola Robins., Anemone deltoidea Hook., Vancouveria hex-
andra (Hook.) Morr. & Dec., Lupinus albicaulis Dougl., and Trifolium
gracilentum T.& G.

Other elements, representing the same general distribution, extend
as far north as the Columbia Gorge, thence southward into California.
Still others have a restricted distribution in central Oregon: Brodiaea
hendersonii Wats., Pleuropogon oregonus Chase, Delphinium leuco-
phaeum Greene, /sopyrum halli Gray, Stanleya confertifolia (Robins. )
Howell, Sidalcea campestris Greene, Lomatium bradshawti (Rose)
Math. & Const., and L. halla (Wats.) Coult. & Rose.

Maritime

A number of species are confined to the sea-coast, mainly to sandy
shores, the latter elements having been discussed by Cooper (1936):

Polypodium scouleri Hook. & Grev., Juncus leseuru Bol., Agrostis
longiligula Hitchc., A. pallens Trin., Calamagrostis crassiglumis Thurb.,
C. nutkaensis (Presl) Steud., Poa confinis Vasey, P. howelli Vasey &
Scribn., P. macrantha Vasey, P. pachypholis Piper, Salix hookeriana
Barr., Abronia latifolia Eschsch., A. umbellata Lam., Sagina crassicaulis
Wats., Spergularia macrotheca (Hornem.) Heynh., Thelypodium lasio-
phyllum (H. & A.) Greene, Sedum spathulifolium Hook., Saxifraga
marshalli Greene, Filipendula occidentalis (Wats.) Howell, Potentilla
pacifica Howell, Sanguisorba menziesu Rydb., Lathyrus littoralis (Nutt.)
Endl., Lupinus littoralis Dougl., Vicia gigantea Hook., Sidalcea hender-
sonu Wats., S. hirtipes Hitche., Angelica hendersoni Coult. & Rose,
Conioselinum pacificum (Wats.) Coult & Rose, Lilaeopsis occidentalis
Coult. & Rose, Sanicula arctopoides H. & A., S. bipinnatifida Dougl.,
Garryva elliptica Dougl., Romanzoffia tracyvi Jeps., Castilleja litoralis
Pennell, Orthocarpus castillejoides Benth., Boschniackia hookeri Wal-
pers, Plantago macrocarpa C. & S., Lasthenia minor (D.C.) Ornduff,
Erigeron glaucus Ker, Ambrosia chamissonis (Less.) Greene, Jaumea
carnosa (Less.) Gray, and Senecio bolanderi Gray.

All of these species are not equally widespread, Poa pachypholis be-
ing restricted to the type locality. Others extend from California to
Alaska: Calamagrostis nutkatensis, Sagina crassicaulis, Pontentilla pa-
cifica, Vicia gigantea, Contoselinum pacificum, and Plantago macro-
carpa. Still others extend from southern British Columbia to California:
Salix hookeriana, Abronia latifolia, Spergularia macrantha, Sidalcea
hendersonu, and Sanicula arctopoides, etc. A number extend from Ore-
gon to California: Saxifraga marshallii, Garrya elliptica, Castilleja
litoralis, Erigeron glaucus, and Senecio bolanderi.

Alaska and Yukon

A considerable portion of Alaska and Yukon escaped glaciation dur-
ing the Pleistocene and served as a refugium for plants. Hultén (1937;

172 MADRONO [Vol. 20

1968) and Porsild (1951; 1966) have been the principal contributors
to the knowledge of this flora and Hultén (1937; 1968) in particular,
has discussed history of the flora. Although many species have expanded
their ranges well beyond the boundaries of Alaska and Yukon, many
others continue to be restricted to areas near the refugia.

Steere has done considerable bryological field work in Alaska and has
discussed this in various papers (Steere, 1938; 1958a; 1959, Schuster &
Steere, 1968) and has contributed most of the information concerning
Alaskan bryophyte endemics but many of his data remain unpublished.
Persson (1946a; 1946b; 1947; 1949, 1952a; 1946b; 1962; 1968) has
contributed richly to the knowledge of the bryoflora of the region.
Although his data have yielded no new information concerning the en-
demics, his detailed discussions have considerably clarified the bryo-
geography. Other publications concerning the bryophytes of Alaska are
Evans (1900; 1901; 1914), Howe (1901), Williams (1901; 1903),
Cardot and Thériot (1902), Cardot (1906), Holzinger and Frye (1921),
Bartram (1938), Clark and Frye (1942; 1946; 1948), Harvill (1947;
1950), Stair (1947; 1948), Thomas (1952), Sherrard (1955; 1957),
Ando, Persson and Sherrard (1957), Steere and Schofield (1956), Pers-
son and Gjaervoll (1957; 1961), Persson and Weber (1958), Schuster
and Steere (1958), Iwatsuki and Sharp (1967; 1968) and Hattori and
Sharp (1968). The most complete bryogeographic summaries are by
Evans (1914), Persson (1949) and Steere (1953; 1965).

Among the bryophytes the Hyegrolejeunea has closest affinities with
tropical species, the Pterigoneurum is largely a genus of arid regions,
the Fruliania is doubtfully distinct from the widely distributed North
American endemic F. bolanderi, and the Trichodon, of close affinity
with a circumboreal species, is known from a single collection and is
therefore not well understood. The Oligotrichum is clearly distinct, and
is not closely related to any western North American species.

Hepatics: Frullania chilcootensis Steph., Hvgrolejeunea alaskana
Schuster & Steere.

Mosses: Oligotrichum falcatum Steere, Trichodon borealis Williams,
and Pterigoneurum arcticum Steere.

Vascular Plants.

Poaceae: Arctagrostis poaeoides Nash, Poa eyerdamu Hult., Pucci-
nellia triflora Swallen, P. interior Sorens., and Agrophyron yukonense
Scribn. & Merr.

Cyperaceae: Carex jacob-peteri Hult. and C. microchaeta Holm.

Salicaceae: Salix setchelliana Ball, S. stolonifera Cov., S. arctolitoralis
Hult., and S. athabascensis Raup.

Betulaceae: Betula kenaica Evans.

Polygonaceae: Polygonum alaskanum (Small) Wright.

1969] SCHOFIELD: PHYTOGEOGRAPHY 173

Chenopodiaceae: Atriplex drymarioides Standl. and A. alaskensis
Wats.

Portulacaceae: Claytonia bostocku Porsild and C. scammaniana Hult.

Caryophyllaceae: Stellaria alaskana Hult. and Melandrium macro-
spermum Porsild.

Ranunculaceae: Ranunculus turneri Greene.

Papaveraceae: Papaver walpolei Porsild.

Cruciferae: Thlaspi arcticum Porsild, Draba exalata Ekman, D.
maxima Hult., D. olgiviensis Hult., Smelowskia pyriformis Drury &
Rollins, S. borealis (Greene) Drury & Rollins, Erysimum angustatum
Rydb., and Brava bartlettiana Jordal.

Saxifragaceae: Boykinia richardsonu (Hook.) Gray, Saxifraga spicata
Don, and S. reflexa Hook.

Leguminosae: Lupinus kuschei Eastw., Astragalus polaris Benth.,
A. nutzotinensis Rousseau, A. williamsii Rydb., Oxytropis kokrinensis
Porsild, O. scammaniana Hult., O. huddlesoniu Porsild, O. glaberrima
Hult., O. kobukensis Welsh, O. Royukukensis Porsild, and O. sheldonen-
sis Porsild.

Umbelliferae: Podistera yukonensis Math. & Const.

Primulaceae: Douglasia arctica Hook., D. gormani Constance, and
Androsace alaskana Cov. & Standl.

Gentianaceae: Gentiana platy petala Griseb.

Hydrophyllaceae: Phacelia mollis Macbr., Romanzoffia sitchensis
Bong., and R. unalaschensis Cham.

Boraginaceae: Eritrichium splendens Kearney and Mertensia drum-
mondu (Lehm.) Don.

Scrophulariaceae: Pentstemon gormani Greene, Synthyris borealis
Pennell, Castilleja unalaschcensis (C. & S.) Malte, C. hyetophila Pen-
nell, C. chryvmactis Pennell, C. vukonis Pennell, C. annua Pennell, C.
villosissima Penell, and Rhinanthus arcticus (Sterneck) Pennell.

Campanulaceae: Campanula aurita Greene.

Compositae: Haplopappus macleani Brandegee, Aster yukonensis
Cronq., Erigeron purpuratus Greene, E. hyperboreus Greene, Antennaria
pallida Nels., A. stolonifera Porsild, A. alborosea Porsild & Porsild, A.
leuchippi Porsild, Artemisia alaskana Rydb., Senecio yukonensis Porsild,
S. hyperborealis Grumm., S. sheldonensis Porsild, Saussurea angustifolia
(Willd.) D.C., and Taraxacum carneocoloratum Nels.

Most of the species are, predictably, of polymorphic circumpolar
genera, but the presence of a species of Boykinia suggests that it is a
Tertiary relict (Hultén, 1968). The endemics are most richly repre-
sented in alpine and subalpine habitats, but a number are maritime and
others in forests, testifying to the diversity or habitats available in the
Pleistocene refugia.

Aleutian Islands
Tatewaki (1963) has suggested ‘“Hultenia” to designate the phyto-

174 MADRONO [Vol. 20

geographic area encompassed by the Aleutian and Commander Islands.
He indicates that both flora and vegetation merit the recognition of this
area and gives a detailed analysis of the floristic composition and affini-
ties. He notes a “marked difference between the (flora of) the Com-
mander Islands and the Aleutian Islands. There is a decided floristic
depression between the first and second district.””’ The Commander
Islands flora is clearly of the Eastern Asiatic floristic Region while the
Aleutian Islands are of the North American floristic Region. He terms
the line between these ‘‘Tatewaki’s Line.” This arch of islands is en-
visioned as a migratory route, serving as a stepping stone corridor for
the expansion of Asiatic species eastward and North American species
westward. Ample floristic evidence is presented to support this concept.

No bryophyte endemics have yet been reported for the area although
unpublished results of Z. Iwatsuki and A. J. Sharp suggest that such
species may be present.

Except for the Polystichum all vascular plants are derivative species
of arctic and alpine areas. The Polvstichum has its affinities with Hima-
layan and Chinese species. The remaining species may be relatively
‘“voung,” belonging to notoriously polymorphic genera in some cases to
Taraxacum, Draba, and Artemisia.

Tatewaki notes the following: Polystichum aleuticum Christens.,
Calamagrostis bracteolata Vassiliev, Elymus aleuticus Hult., Cerastium
aleuticum Hult., Draba aleutica Ekman, Artemisia aleutica Hult.,
Taraxacum chromocarpum Hagl., T. eyverdamiu Hagl., and T. onco-
phorum Haegl.

Although noted for the Aleutian Islands by Tatewaki, Hultén (1968)
does not indicate the presence of the Elymus or the Taraxacum species.
In this flora however, the following species are essentially restricted to
the Aleutian Islands, although in all cases these species extend also to
the Alaskan mainland as well: Poa hispidula Vasey, Poa turneri Scribn.,
Salix cyclophylla Rydb., and Gentiana aleutica C. &S.

The Queen Charlotte Islands

The Queen Charlotte Islands of British Columbia have served as a
refugium for a number of species, both endemics and disjunct fragments
of a flora of pre-glacial times. The higher elevations, at least, escaped
glaciation, and the affinities of many bryophytes and vascular plants
imply that they are pre-Pleistocene relicts. Calder and Taylor (1968)
have thoroughly treated the vascular flora and Persson (1958), and
Schofield (1962; 1965; 1966b; 1968a; 1968b) have provided prelimi-
nary notes concerning the bryophytes.

Among the bryophytes only the endemic Acanthocladium carlottae
Schof. has been described although there remain undescribed species of
Seligeria, Brotherella, Acanthocladium, and Mastopoma (?). The latter
three genera suggest a montane flora of a subtropical latitude, the rela-

1969 | SCHOFIELD: PHYTOGEOGRAPHY Lis

tionship of each of the species being largely with the Malaysian area,
and suggesting great antiquity. There is a rich representation in the
Islands of species showing affinities either with East Asia or Western
Europe. These are discussed later under these disjunct elements.

The endemic vascular plants are confined largely to higher elevations
or to habitats of lower elevations on the flanks of the mountains. The
bryophyte disjuncts and endemics show a similar restriction. In all
cases relationships of the undescribed taxa is with taxa of distant un-
glaciated areas rather than with those of adjacent glaciated areas, em-
phasizing that the species are probably pre-glacial relicts.

Vascular Plants: /sopyvrum saviler Calder & Taylor, Saxifraga taylori
Calder & Savile, Geum schofieldu Calder & Taylor, Ligusticum calderi
Math. & Const., and Senecio newcombei Greene.

Columbia River Gorge

Piper (1906) noted that the gorge of the Columbia River and valleys
of adjacent tributaries served as an area of endemism. He noted that 16
species were endemic to the gorge. Since that time many have either
been found to be more widespread or have slipped into the synonymy
of more widespread species. Detling (1958), in discussing the flora of
the gorge noted 7 species endemic to the gorge. Douglasia laevigata
Gray is more widespread, and thus should be excluded. Perusal of Hitch-
cock, et al. (1955-1969) indicates that 17 species are indeed endemic to
the Columbia River Gorge, although a number do extend sometimes
into the Wilamette Valley or into some of the tributary watercourses
of the Columbia River.

Detling (1958) suggests that the gorge served as a corridor of migra-
tion for both lowland and highland species, supporting this concept
by noting disjunctions of species in the gorge and in these other areas.
He suggests that the lowland migrations probably occurred during the
Hypsithermal and that the montane elements migrated downwards from
higher elevations during the Pleistocene refrigeration. The endemics,
fragments of these floras, are suggested to be relics, restricted in their
range by rather narrow environmental tolerance. Unfortunately no ex-
perimental evidence is available to support or refute this hypothesis.

A single bryophyte has been noted as endemic to the Columbia River
Gorge (Hermann & Lawton, 1968): Desmatodon columbianus Hermann
& Lawt.

Vascular Plants: Agrostis howell Scribn., Calamagrostis howellii
Vasey, Poa leibergu Scribn., Allium robbinsu Henderson, A. pleianthum
Wats., Salix fluviatilis Nutt., Bolandra oregana Wats., Sullivantia ore-
gana Wats., Astragalus diaphanus Dougl., Eryngium petiolatum Hook.,
Lomatium columbianum Math. & Const., L. laevigatum (Nutt.) Coult.
& Rose, Cryptantha leucophaea (Dougl.) Pays., Pentstemon barrettae
Gray, Erigeron howellii Gray, E. oreganus Gray, and Hieracium longi-
berbe Howell.

176 MADRONO [Vol. 20

BOREAL

The Boreal flora is composed of four elements of particular signifi-
cance: circumboreal, circumboreal maritime, boreal American, and cir-
cumboreal through anthropogenic introduction. In the boreal bryoflora
the North American vegetation is dominated by circumboreal species,
with remarkably few endemic taxa while in the vascular flora the con-
spicuous elements of the vegetation are endemic to North America, thus
all tree species and most shrubby species are endemic to North America
(exception: Alnus crispa). Many circumboreal bryophytes and herba-
ceous vascular plants are also conspicuous elements in the Arctic flora.
Many of these species extend their ranges southward in the mountains
as far as Arizona, and, in some cases, into Mexico. The woody species, on
the other hand, are largely supplanted southward by western North
American endemics, even in the Northern Rocky Mountains.

Circum boreal

Hepatics: Riccardia sinuata (Dick.) Trevis., R. pinguis (L.) Gray,
Pellia endiviifolia (Dicks.) Dumort., P. neesiana (Gottsche.) Limpr.,
P. epiphylla (L.) Lindb., Metzgeria conjugata Lindb., Moerckia floto-
viana (Nees.) Schiffn., Blasia pusilla L., Fossombronia dumortiert (Hub.
& Genth.) Lindb., Ptilidium ciliare (L.) Hampe, P. pulcherrimum
(Web.) Hampe, Lepidozia reptans (L.) Dumort., Bazzania trilobata
(L.) Gray, B. tricrenata (Wg.) Trevis., Calvpogeia neesiana (Mass. &
Carest.) Miull., C. sphagnicola (Arn. & Perss.) Warnst. & Loeske, C.
trichomanis (L.) Corda, C. fissa (L.) Raddi, C. suecica (Arn. & Perss.)
Miull., Cephaloziella elachista (Jack.) Schiffn., C. hampeana (Nees.)
Schiffn., C. rubella (Nees.) Douin, Anastrophyllum michauxu (Web.)
Buch., Barbilophozia barbata (Schmid.) Loeske, B. lycopodioides
(Wallr.) Loeske, Gymnocolea inflata (Huds.) Dumort., Jamesoniella
autumnalis (D.C.) Steph., Jungermannia lanceolata Schrad., J. pumila
With., J. atrovirens Dumort., J. tristis Nees., J. sphaerocarpa Hook.,
Leiocolea heterocolpos (Thed.) Buch., L. gillmani (Aust.) Evans,
Lophozia excisa (Dicks.) Dumort., L. marchica (Nees.) Steph., L.
incisa (Schrad.) Dumort., Mylia taylori (Hook.) Gray, M. anomala
(Hook.) Gray, Nardia scalaris (Schrad.) Gray, N. geoscyphus (DeNot.)
Lindb., Orthocaulis kunzeanus (Hib.) Buch., Plectocolea obovata
(Nees.) Mitt., P. hyalina (Lyell) Mitt., Sphenolobus minutus (Crantz.)
Steph., Lophocolea heterophylla (Schrad.) Dumort., L. minor Nees.,
L. cuspidata (Nees.) Limpr., Chiloscyphus polyanthos (L.) Corda,
Harpanthus scutatus (Web. & Mohr.) Spr., Geocalyx graveolens
(Schrad.) Nees., Plagiochila asplenioides (L.) Dumort., Diplophyllum
taxifolium (Wahl.) Dumort., D. albicans (L.) Dumort., Scapania irri-
gua (Nees.) Dumort., S. paludicola Loeske & Miill., S. umbrosa (Schrad.)
Dumort., S. undulata (L.) Dumort., Cephalozia bicuspidata (L.) Du-
mort., C. connivens (Dicks.) Spr., C. catenulata (Hib.) Lindb., C.

1969] SCHOFIELD: PHYTOGEOGRAPHY Mee

media Lindb., C. macouniu Aust., Cladopodiella fluitans (Nees.) Spr.,
Odontoschisma denudatum (Nees) Dumort., O. elongatum (Lindb.)
Evans, Gymnomutrion concinnatum Corda, Marsupella sphacelata (Gies.)
Lindb., M. sparsifolia (Lindb.) Dumort., M. emarginata (Ehrh.) Du-
mort., Radula complanata (L.) Dumort., Porella platyphylla (L.)
Lindb., Preissia quadrata (Scop.) Nees., Conocephalum conicum (L.)
Dumort., Reboulia hemispherica (L.) Raddi, Riccia sorocarpa Bisch.,
R. crystallina L., R. fluitans L., and Ricciocarpus natans (L.) Corda.

Mosses: Sphagnum nemoreum Scop., S. rubellum Wils., S. fimbriatum
Wils., S. fuscum (Schimp.) Klinger., S. girgensohnii Russ., S. papillosum
Lindb., S. squarrosum Crome., Andreaea rupestris Hedw., Fissidens
adianthoides Hedw., F. bryoides Hedw., F. osmundioides Hedw., Tri-
chodon cvylindricus (Hedw.) Schimp., Ditrichum heteromallum (Hedw.)
Britt., Distichium capillaceum (Hedw.) B.S.G., Blindia acuta (Hedw.)
B.S.G., Trematodon ambiguus (Hedw.) Hornsch., Dicranella hetero-
malla (Hedw.) Schimp., D.rufescens (With.) Schimp., D. varia (Hedw.)
Schimp., Dicranodontium denudatum (Brid.) Britt., Amphidium lap-
ponicum (Hedw.) Schimp., Dichondontium pellucidum (Hedw.) Schimp.,
Oncophorus wahlenbergu Brid., Kiaeria starket (Web. & Mohr.) Hag.,
Dicranum elongatum Schleich., D. fuscescens Turn., D. scoparium
Hedw., Encalypta ciliata Hedw., E. vulgaris Hedw., Tortella fragilis
(Hook.) Limpr., T. tortuosa (Hedw.) Limpr., Bryoerythrophyllum
recurvirostrum (Hedw.) Chen, Barbula convoluta Hedw., B. unguiculata
Hedw., Pottia heim (Hedw.) Furnr., Tortula mucronifolia Schwaegr.,
T. norvegica (Web.) Wahlenb., 7. ruralis (Hedw.) Gaertn., Mey., &
Scherb., Grimmia alpicola Hedw., G. apocarpa Hedw., Racomitrium
aciculare (Hedw.) Brid., R. canescens (Hedw.) Brid., R. lanuginosum
(Hedw.) Brid., Tavyloria lingulata (Dicks.) Lindb., Tetraplodon an-
gustatus (Hedw.) B.S.G., Splachnum ampullaceum Hedw., Tetraphis
pellucida Hedw., Pohlia nutans (Hedw.) Lindb., P. wahlenbergii (Web.
& Mohr.) Andr., Leptobryum pyriforme (Hedw.) Wils., Bryum pallens
Sw., Plagiomnium affine (Bland.) Koponen, Mnium spinulosum (Voit.)
Schwaegr., Aulacomnium palustre (Hedw.) Schwaegr., Meesea trifaria
Crum, Steere, & Anderson, Paludella squarrosa (Hedw.) Brid., Catas-
copium nigritum (Hedw.) Brid., Plagiopus oederiana (Sw.) Limpr.,
Philonotis fontana (Hedw.) Brid., Timmia austriaca Hedw., Ortho-
trichum obtusifolium Brid., O. speciosum Nees., Ulota phyllantha Brid.,
Fontinalis antipyretica Hedw., Climacium dendroides (Hedw.) Web. &
Mohr., Neckera pennata Hedw., Myurella julacea (Schwaegr.) B.S.G.,
Leskea polycarpa Hedw., Thuidium recognitum (Hedw.) Lindb., A dieti-
nella abietina (Hedw.) Fleisch., Cratoneuron filicinum (Hedw.) Spruce,
Campylium stellatum (Hedw.) Jens., Leptodictyum riparium (Hedw.)
Warnst., Amblystegium serpens (Hedw.) B.S.G., Drepanocladus adun-
cus (Hedw.) Warnst., D. uncinatus (Hedw.) Warnst., Hygrohyvpnum
luridum (Hedw.) Jenn., Calliergon cordifolium (Hedw.) Kindb., Scor-

178 MADRONO [Vol. 20

pidium scorpioides (Hedw.) Limpr., Tomenthypnum nitens (Hedw.)
Loeske, Brachythecium albicans (Hedw.) B.S.G., B. plumosum (Hedw.)
B.S.G., Eurhynchium praelongum (Hedw.) B.S.G., E. pulchellum
(Hedw.) Jenn., Pterigynandrum filiforme Hedw., Orthothecium chry-
seum (Schwaegr.) B.S.G., Pleurozium schrebert (Brid.) Mitt., Plagio-
thecium denticulatum (Hedw.) B.S.G., Pylaisiella polyantha (Hedw.)
Grout, Hypnum callichrom Funck., H. revolutum (Mitt.) Lindb., /sop-
tervgium pulchellum (Hedw.) Jaeg. & Sauerb., Ptilium crista-castrensis
(Hedw.) DeNot., Rhytidiadelphus triquetrus (Hedw.) Warnst., Hylo-
comium splendens (Hedw.) B.S.G., Atrichum undulatum (Hedw.)
Beauv., Pogonatum alpinum (Hedw.) Rohl., P. urnigerum (Hedw.)
Beauv., and Polytrichum piliferum Hedw.

Vascular Plants.

Lycopodiaceae: Lycopodium annotinum L. and L. clavatum L.

Selaginellaceae: Selaginella selaginioides (L.) Link.

Equisetaceae: Equisetum variegatum Schleich., E. fluviatile L., and
i, arvense L.

Ophioglossaceae: Botrychium lunaria (L.) Sw.

Polypodiaceae: Pteridium aquilinum (L.) Kuhn, Thelypteris phegop-
teris (L.) Slosson, Athyrium filix-femina (L.) Roth., Cystoperis fragilis
(L.) Bernh., Woodsia ilvensis (L.) R.Br., Drvopteris dilatata (Hoffm.)
Gray, and Gymnocar pium drvyopteris (L.) Newm.

Cupressaceae: Juniperus communis L.

Typhaceae: Typha latifolia L.

Sparganiaceae: Sparganium augustifolium Michx.

Potamogetonaceae: Potamogeton natans L., P. gramineus L., and
P. filiformis Pers.

Scheuchzeriaceae: Scheuchzeria palustris L.

Poaceae: Phalaris arundinacea L., Hierochloe odorata (L.) Wahlenb.,
Alopocurus aequalis Sobol., Cinna latifolia (Trev.) Griseb., Agrostis
borealis Hartm., Calamagrostis neglecta (Ehrh.) Gaertn., Mey., &
Scherb., Tvisetum spicatum (L.) Richter, Beckmannia erucaeformis
(L.) Host, Poa glauca Vahl, P. palustris L., Glyceria maxima (Hartm.)
Holmb., and Bromus inermis Leyss.

Cyperaceae: Eriophorum angustifolium Honck., Trichophorum caes-
pitosum (L.) Hartm., Eleocharis uniglumis (Link.) Schult., Rhynchos-
pora alba (L.) Vahl., Carex pauciflora Lightf., C. diandra Schrank,
C. canescens L., C. disperma Dew., C. limosa L., and C. rostrata Stokes.

Araceae: Calla palustris L.

Juncaceae: Juncus alpinus Vill., J. articulatus L., and Luzula parvi-
flora (Ehr.) Desv.

Orchidaceae: Cypripedium calceolus L., Listera cordata (L.) R.Br.,
Platanthera obtusata (Pursh) Lindb., Goodyera repens (L.) R.Br.,
Corrallorhiza trifida Chatelain, and Calypso bulbosa (L.) Richb. f.

Salicaceae: Salix phylicifolia L.

1969] SCHOFIELD: PHYTOGEOGRAPHY 179

Myricaceae: Myrica gale L.

Betulaceae: Betula nana L.

Polygonaceae: Koenigia islandica L. and Polygonum amphibium L.

Caryophyllaceae: Chenopodium glaucum L., Stellaria longifolia Muhl.,
S. calycantha (Ledeb.) Bong., Cerastium arvense L., Sagina nodosa (L.)
Fenzl., and Moehringia lateriflora (L.) Fenzl.

Ceratophyllaceae: Ceratophyllum demersum L.

Ranunculaceae: Caltha palustris L., Ranunculus trichophyllus Chaix.,
and R. sceleratus L.

Cruciferae: Subularia aquatica L., Cardamine pratensis L., and Arabis
hirsuta (L.) Scop.

Droseraceae: Drosera rotundifolia L.

Crassulaceae: Sedum rosea (L.) Scop.

Rosaceae: Rubus chamaemorus L., Potentilla palustris (L.) Scop.,
P. fruticosa L., and Sanguisorba officinalis L.

Leguminosae: Hedysarum alpinum L.

Linaceae: Linum perenne L.

Callitrichaceae: Callitriche hermaproditica L.

Violaceae: Viola selkirki Pursh.

Onagraceae: Epilobium angustifolium L., E. palustre L., and Circaea
alpina L.

Haloragidaceae: Myriophyllum verticellatum L.

Cornaceae: Cornus suecica L.

Ericaceae: Pyrola secunda L., Moneses uniflora (L.) Gray, Mono-
tropa hypopitys L., Empetrum nigrum L., Ledum palustre L., Andromeda
polifolia L., Chamaedaphne calyculata (L.) Moench., Arctostaphylos
uva-urst (L.) Spreng., Vaccinium vitis-idaea L., V. uliginosum L., and
Oxycoccus microcar pus Turcz.

Primulacaceae: Androsace septentrionalis L., and Lysimachia thyrsi-
flora L.

Labiatae: Scutellaria galericulata L., Stachys palustris L., and Men-
tha arvensis L.

Scrophulariaceae: Limosella aquatica L. and Veronica scutellata L.
Lentibulariaceae: Utricularia intermedia Hayne and U. vulgaris L.
Rubiaceae: Galium boreale L. and G. triflorum Michx.

Caprifoliaceae: Sambucus racemosa L. and Linnaea borealis L.

Campanulaceae: Campanula rotundifolia L.

Compositae: Evigeron acris L. and Senecio congestus (R.Br.) D.C.

Circumboreal Maritime

No hepatics are restricted to sea-shores although several tolerate some
salinity. Among the mosses only two are essentially restricted to mari-
time habitats, both occurring on rocks affected by salt spray: Grimmua
maritima Turn. and Ulota phyllantha Brid. Other mosses are tolerant
of salt spray, but are not restricted to such habitats.

Among the vascular plants are a number of obligate halophytes. In

180 MADRONO [Vol. 20

some cases these are found away from the sea-coast, but generally in
saline or alkaline environments. In North America there are some ex-
ceptions, e.g., Lathyrus maritimus in the Great Lakes area, Armeria
maritima in the Rocky Mountains.

Vascular Plants: Zostera marina L., Ruppia spiralis L., Calamagrostis
deschampsioides Trin., Puccinellia phryganodes (Trin.) Scribn. & Merr.,
Elymus arenarius L., Corex glareosa Wahlenb., C. mackenziei Krecz.,
Stellaria humifusa Rottb., Honckenya peploides (L.) Ehrh., Cochlearia
officinalis L., Potentilla egedit Wormsk., Lathyrus maritimus L., Hip-
purts tetraphylla L. {., Ligusticum scoticum L., Armeria maritima (Mill.)
Willd., Mertensia maritima (L.) Gray, and Tripleurospermum phaeoce-
phalum (Rupr.) Pobed.

Boreal American

This element is composed of endemic species of wide distribution in
northern North America. The number and vegetational importance of
endemic bryophytes of this distribution pattern is not significant but
the vascular plants, particularly woody species, are main components
of the vegetation.

Hepatics: Plectocolea obscura Evans is the only species that can be
placed here and even this species is uncertain, being reported from
Northeastern United States and Oregon. The latter record needs verifi-
cation.

Mosses: Seligeria campylopoda Kindb., Grimmia dupretu Ther.,
Physcomitrium immersum Sull., Philonotis americana Dism., and Clima-
cium americanum Brid.

Few of these species are common, the exceptions being Philonotis
americana and Climacium americanum. The others are infrequent and
in rather specialized habitats.

Vascular Plants (dominant or conspicuous elements of the vegetation
are designated by an asterisk).

Pinaceae: Pinus banksiana Lamb.,* Larix laricina (DuRoi) Koch.,*
Picea glauca (Moench.) Voss.,* and P. mariana (Mill.) Britt., Sterns.,
Pogg.*

Cupressaceae: Juniperus horizontalis Moench.*

Sparganiaceae: Sparganium eurycarpum Engelm. and S. multipedun-
culatum (Morong.) Rydb.

Potamogetonaceae: Potamogeton epihydrus Raf. and P. foliosus Raf.

Alismataceae: Sagittaria cuneata Sheld.

Poaceae: Oryzopsis pungens (Torr.) Hitchc., Muhlenbergia richard-
sonis (Willd.) Trin., M. glomerata (Willd.) Trev., Agrostis geminata
Trin., Calamagrostis canadensis (Michx.) Beauv.,* Danthonia spicata
(L.) Beauv., Sphenopholis intermedia (Rydb.) Rydb., Glyceria borealis
(Nash) Batchelder, G. striata (Lam.) Hitche., Festuca saximontana

1969} SCHOFIELD: PHYTOGEOGRAPHY 181

Rydb., Agropyron smithii Rydb., A. subsecundum (Link.) Hitchc., and
A. pauciflorum (Schwein.) Hitchc.

Cyperaceae: Eriophorum viridi-carinatum (Engelm.) Fern., Scirpus
subterminalis Torr., S. americanus Pers., S. paludosus Nels., S. validus
Vahl.,* S. microcar pus Presl., Carex leptalea Wahlenb., C. bebbu Olney,
C. crawfordii Fern., C. aenea Fern., C. arcta Boott, C. interior Bailey,
C. deweyana Schwein., C. aurea Nutt., C. garberit Fern., C. deflexa
Hornem., C. concinna R.Br., C. eburnea Brott, and C. lanuginosa Michx.

Juncaceae: Juncus nodosus L.

Liliaceae: Smilacina racemosa (L.) Desf. and S. stellata (L.) Desf.

Iridaceae: Sisyrinchium montanum Greene.

Orchidaceae: Amerorchis rotundifolia (Banks) Hult., Platanthera
orbiculata (Pursh) Lindb., P. dilatata (Pursh) Lindb., Listera conval-
larioides (Sw.) Nutt., and Corallorhiza maculata Raf.

Salicaceae: Populus balsamifera L.,* P. tremuloides Michx.,* Salix
arctophila Cockerell.,* S. brachycarpa Nutt., S. pedicellaris Pursh, S.
mackenzteana Barratt, S. myrtillifolia Anderss., S. candida Flugge, and
S. interior Rowlee.

Betulaceae: Betula glandulosa Michx.* and B. papyrifera Marsh.*

Urticaceae: Urtica gracilis Ait.

Santalaceae: Geocaulon lividum (Richards.) Fern.

Polygonaceae: Rumex fenestratus Greene, Polygonum pennsylvani-
cum L., and P. achoreum Blake.

Caryophyllaceae: Arenaria dawsonensis (Britt.) Mattf.

Nymphaeaceae: Nuphar variegatum Engelm.

Ranunculaceae: Actaea rubra (Ait.) Willd., Anemone multifida Poir.,
Ranunculus abortivus L., R. pennsvilvanicus L. f., and, R. macounii
Britt.

Fumariaceae: Corydalis aurea Willd. and C. sempervirens (L.) Pers.

Cruciferae: Rorippa obtusa (Nutt.) Britt., Cardamine pennsvlvanica
Muhl., Draba aurea Vahl., Descurainia richardsoniu (Sweet) Schulz,
Arabis arenicola ( Richards.) Gelert, and Erysimum inconspicum (Wats.)
MacM.

Saxifragaceae: Saxifraga tricuspidata Rottb., Ribes oxycanthoides L.,
R. hudsonianum Richards., and R. glandulosum Grauer.

Rosaceae: Rubus pubescens Raf., R. arcticus L., Fragaria virginiana
Duchesne, Potentilla vahliana Lehm., P. argentea Pursh, and P. penn-
sylvanica L.

Leguminosae: Oxytropis deflexa (Pall.) D.C., Hedysarum mackenziet
Richards. and Vicia americana Miuhl.

Geraniaceae: Geranium bicknelli Britt.

Violaceae: Viola adunca Sm. and V. renifolia Gray.

Elaeagnaceae: Shepherdia canadensis (L.) Nutt.* and Elaeagnus com-
mutata Bernh.

Onagraceae: Epilobium leptophyllum Raf.

182 MADRONO [Vol. 20

Umbelliferae: Cicuta bulbifera L.

Ericaceae: Ledum groenlandicum Oeder,* Kalmia polifolia Wang.,
and Vaccinium caespitosum Michx.

Primulaceae: Primula mistassinica Michx.

Apocynaceae: A pocynum androsaemifolium L.

Labiatae: Dracocephalum parviflorum Nutt.

Scrophulariaceae: Euphrasia disjuncta Fern. & Wieg. and Pedicularis
groenlandica Retz.

Rubiaceae: Galium brandegei Gray.

Caprifoliaceae: Viburnum edule (Michx.) Raf., Svmphoriocarpus
albus (L.) Blake,* and Lonicera involucrata (Richards.) Banks.

Compositae: Solidago multiradiata Ait., S. canadensis L., Aster laevis
L., A. junciformis L., Erigeron compositus Pursh, E. hyssopifolius
Michx., EL. elatus Greene, E. philadelphicus L., Antennaria pulcherrima
(Hook.) Greene, Achillea lanulosa Nutt., Artemisia canadensis Michx.,
Petasites sagittatus (Banks) Gray, Senecio pauciflorus Pursh, S. pauper-
culus Michx., Taraxacum lacerum Greene, and Lactuca biennis (Moench.)
Fern.

Circumboreal by Anthropogenic Introduction

In the bryophytes it is rather difficult to determine anthropogenic
introductions since such a high proportion of the species show a natural
circumboreal distribution. Certainly the distribution of many circum-
boreal species have been anthropogenically expanded by destruction of
competing native vascular plants and by clearing sites, but in many
cases, if abandoned by man, such sites revert to a covering of vascular
plant vegetation and thus bryophytes are eliminated. In cities, however,
a number of presumably introduced species do persist in gardens and
on stone or concrete walls, and sometimes as lawn weeds. Most of the
species are also natural elements of the local flora, thus invasion of the
urban sites cannot be confidently attributed to anthropogenic intro-
duction. Among such bryophytes are:

Hepatics: Blasia pusilla L. and Marchantia polymorpha L.

Mosses: Ceratodon purpureus (Hedw.) Brid., Dicranowetsia cirrata
(Hedw.) Lindb., Barbula vinealis Brid., Pottia truncata (Hedw.) Furnr.,
Tortula ruralis Hedw., T. muralis Hedw., Grimmia apocarpa Hedw.,
Funaria hygrometrica Hedw., Pohlia annotina (Hedw.) Lindb., Lepto-
bryvum pyriforme (Hedw.) Wils., Bryum argenteum Hedw., Callier-
gonella cuspidata (Hedw) Loeske, Brachythecium albicans (Hedw.)
B.S.G., Eurhynchium praelongum (Hedw.) B.S.G., and Rhytidiadelphus
squarrosus (Hedw.) Warnst.

It should be noted that most of these bryophytes are common ele-
ments of the natural circumboreal flora, but their invasion of anthro-
pogenic environments has considerably expanded their local ranges. The
vascular plants, on the other hand, are mainly accidental introductions

1969] SCHOFIELD: PHYTOGEOGRAPHY 183

and in many cases are noxious weeds of arable land. Many species were
introduced first in eastern North America and have expanded their
ranges westward with the activity of man. Many were introduced in
ship’s ballast, others with seeds of domestic crops and a number have
escaped from cultivation.

Vascular Plants

Poaceae: Anthoxanthum odoratum L., Phleum pratense L., Alopecurus
pratensis L., Agrostis tenuis Sibth., A. stolonifera L., Holcus lanatus L.,
Avena fatua L., A. sativa L., Arrhenatherum elatius (L.) Presl & Presl,
Dactylis glomerata L., Poa trivialis L., P. pratensis L., P. annua L.,
Festuca arundinacea Schreb., Bromus tectorum L., B. secalinus L.,
Lolium perenne L., L. tementulum L., Agropyron pecteniforme Roem. &
Schult., and A. repens (L.) Beauv.

Urticaceae: Urtica urens L.

Polygonaceae: Rumex acetosella L., R. acetosa L., R. obtusifolius L.,
R. crispus L., Polygonum convolvulus L., P. persicaria L., P. hydro piper
L., and P. aviculare L.

Chenopodiaceae: Chenopodium rubrum L. and C. album L.

Caryophyllaceae: Stellaria media (L.) Vill., Spergularia rubra (L.)
Presl & Presl, Agrostemma githago L., Melandrium noctiflorum (L.)
Fries, and Vaccaria pyramidata Medic.

Ranunculaceae: Ranunculus repens L. and R. acris L.

Papaveraceae: Papaver rhoeas L.

Cruciferae: Lepidium sativum L., Thlaspi arvense L., Sisyrinchium
officinale (L.) Scop., S. altissimum L., Sinapsis arvensis L., Brassica
juncea (L.) Czern., B. rapa L., Raphanus sativus L., Rorippa nastur-
tium-aquaticum (L.) Hayek., Capsella bursa-pastoris (L.) Medic.,
Neslia paniculata (L.) Desv., Descurainia sophia (L.) Prantl, Turritis
glabra L., Ervsimum cheiranthoides L., andd Hesperis matronalis L.

Leguminosae: Medicago sativa L., M. lupulina L., Melilotus officinalis
(L.) Lam., WM. albus Desv., Trifolium hybridum L., T. repens L., T.
pratense L., Vicia angustifolia (L.) Reichard., and V. cracca L.

Gerianaceae: Geranium robertianum L.

Umbelliferae: Pastinaca sativa L.

Boraginaceae: Lappula myosotis Moench. and Myosotis palustris L.

Labiatae: Nepeta cataria L., Glechoma hederacea L., and Galeopsis
bifida Boenn.

Scrophulariaceae: Linaria vulgaris Mill., Veronica anagallis-aquatica
L., V. persica Poir., and V. arvensis L.

Plantaginaceae: Plantago lanceolata L. and P. major L.

Compositae: Guaphalium uliginosum L., Anthemis cotula L., Matri-
caria matricarioides (Less.) Porter, Tripleurospermum inodorum (L.)
Schultz-Bip., Chrysanthemum vulgare (L.) Bernh., Senecio vulgaris L.,
Cirsium arvense (L.) Scop., C. vulgare (Savi) Ten., Leontodon autumn-
alis L., Taraxacum officinale Weber, and Cre pis tectorum L.

184 MADRONO [Vol. 20

CIRCUMARCTIC

A number of species are restricted to arctic regions, rarely extending
into the subarctic. Steere (1953; 1965) has discussed the bryogeographic
element and Porsild (1957) has noted vascular plants of this distribu-
tional type.

Hepatics: Mesoptychia sahlbergu (Lindb., & Arn.) Evans, Lophozia
latifolia Schuster, and Plagiochila arctica Bryhn. & Kaalas.

Musci: Psilopilum laevigatum (Wahlenb.) Limpr., Distichium hagenii
Ryan, Blindia polaris ( Berger.) Hag., Haplodon wormsk joldii (Hornem.)
R.Br., Tetraplodon paradoxus (R.Br.) Hagen, Pohlia crudoides (Sull.
& Lesq.) Broth., Bryum wrighti Sull., Cyrtomnium hymenophyllum
(B.S.G.) Holmen, Cinclidium latifolium Lindb., C. subrotundum Lindb.,
and Aulacomnium acuminatum (Lindb. & Arn.) Par.

Vascular Plants: Arctagrostis latifolia (R.Br.) Griseb., Colpodium
vahlianum (Liebm.) Nevski, Arctophila fulva (Trin.) Anderss., Puc-
cinellia phryganodes (Trin.) Scribn. & Merr., Agropyron boreale (Turcz.)
Drobov, Eriophorum triste (T. Fries) Love & Hadac, Carex subs patha-
cea Wormskj., C. adelostoma Krecz., C. krauseit Beocl, Luzula arctica
Blytt., Salix arctica Pall., Cerastium regeli Ostenf., Minuartia strictat
(Sw.) Kiern., Ranunculus confervoides (Fries) Fries, R. pallasu
Schlecht., R. lapponicus L., R. sulphureus Soland., Cochlearia officinalis
L., Eutrema edwardsu R.Br., Draba subcapitata Simm., D. micropetala
Hook., D. alpina L., D. macrocarpa Adams, Braya purpurascens (R.Br.)
Bunge, Saxifraga hieracifolia Waldst. & Kit., S. foliolosa R.Br., Potentilla
hyparctica Malt, P. pulchella R.Br., Dryas octopetala L., Pyrola grandt-
flora Radius, Casstope tetragona (L.) Dvn., Lomatogonium rotatum
(L.) Fries., Pedicularis lapponica L., and Erigeron eriocephalus Vahl.

ARCTIC-ALPINE

Hultén (1937) has suggested that, for species of this distributional
pattern, Arctic-Montane is more appropriate, since this does not imply
that the species are present in the European Alps. Although this is true,
the term alpine has been used traditionally in a more general way, de-
noting any montane area above tree line. Arctic-alpine, as generally
used, indicates that a species is widespread in Arctic regions, i.e., north
of tree-line, and extends southward in higher elevations of mountains
or in sites edaphically equivalent (cliffs, bogs, headlands, etc.). It has
been shown (Mooney and Billings, 1961; Mooney and Johnson, 1965)
that, among the flowering plants, the alpine populations of arctic alpine
species represent ecotypes in those species that have been experimentally
examined. It is possible that the bryophytes of this distribution also
possess ecotypes. In the bryophytes, however, vegetative reproduction
decreases selection and thus segregation of ecotypes is greatly impeded.
Persistence of bryophytes in microenvironments that closely match the

1969] SCHOFIELD: PHYTOGEOGRAPHY 185

macroenvironment of arctic regions would also work against the type
of selection that leads to alpine ecotypes in vascular plants.

Hepatics: Anthelia julacea (L.) Dumort., A. juratzkana (Limpr. )
Trevis, Cephaloziella arctica Bryhn. & Douin, Arnellia fennica
(Gottsche.) Lindb., /sopaches bicrenatus (Schmid.) Buch., Lziocolea
badensis (Gottsche.) Joerg., L. bantriensis (Hook.) Joerg., L. muelleri
(Nees.) Joerg., Lophozia longiflora (Nees.) Schiffn., L. ventricosa
(Dicks.) Dumort., L. alpestris (Schleich.) Evans, L. wenzeli (Nees.)
Steph., L. longidens (Lindb.) Macoun, Orthocaulis binsteadi (Kaal.)
Buch., O. attenuatus (Mart.) Evans, O. quadrilobus (Lindb.) Buch.,
Saccobasis polita (Nees.) Buch., Tritomaria exsecta (Schmid.) Schiffn.,
T. exsectiformis (Breidler) Schiffn., T. quinquedentata (Huds.) Buch.,
Harpanthus flotowianus Nees., Scapania cuspiduligera (Nees.) Mull.,
S. uglinosa (Sw.) Dumort., S. subalpina (Nees.) Dumort., S. paludosa
(Mull.) Mull., Cephalozia pleniceps (Aust.) Lindb., C. ambigua Mass.,
C. striatula Jens., Pleuroclada albescens (Hook.) Spr.,Gyvmnomitrion
coralloides Nees., Marchantia alpestris Nees., Mannia pilosa (Hornem. )
Frye & Clark, Asterella ludwigit (Schwaegr.) Underw., Peltole pis qua-
drata (Sauter) Miill., Clevea hyalina (Sommerf.) Lindb., and Sauteria
alpina Nees.

Mosses: Trematodon brevicollis Hornsch., Arctoa fulvella ( Dicks.)
B.S.G., Dicranum acutifolium (Lindb. & Arn.) Jens., D. elongatum
Schleich., Encalypta affinis Hedw. f., EF. brevicolla (B.S.G.) Bruch.,
Molendoa tenuinervis Limpr., Barbulaicmadophila Schimp., Didvmodon
rufus Lor., Pottia heimiu (Hedw.) Furn., Stegonia latifolia (Schwaegr. )
Vent., Desmatodon svstylius B.S.G., D. laureri (Schulz.) B.S.G., Voztia
nivalis Hornsch., Tavloria froelichiana (Hedw.) Lindb., T. splachnoides
(Schleich.) Hook.,Pohlia schimperi (C.M.) Andr., P. drummondii
(C.M.) Andr. Plagiobryum demissum (Hoppe & Hornsch.) Lindb.,
Bryum obtustfolium Lindb., Mnium blytti B.S.G., Cyrtomnium hymeno-
phylloides (Hutb.) Koponen, Cinclidium styvgium Sw., Aulacomnium
turgidum (Wahl.) Schwaegr., Amblvodon dealbatus (Hedw.) B.S.G.,
Conostomum tetragonum (Hedw.) Lindb., Bartramia ithyphyila Brid.,
Myurella tenerrima (Brid.) Lindb., Drepanocladus tundrae (H. Arnell)
Loeske, Cirriphvllum cirrosum (Schwaegr.) Grout, Hypnum bambergeri
Schimp., H. vaucheri Lesq., H. procerrimum Mol., and Rhytidium
rugosum (Hedw.) Kindb.

Vascular Plants: Huperzia selago (L.) Bernh., Lycopodium alpinum
L., Woodsia alpina (Bolton) Gray, Hierochloe alpina (Sw.) Roem. &
Schult., Phleum commutatum Gandoger, Alopecurus alpinus Sm.,
Phippsia algida (Soland.) R.Br., Poa alpina L., P. arctica R.Br., Erio-
phorum scheuchzeri Hoppe, Kobresia myosuroides (Vill.) Fiori & Paol.,
K. simpliciuscula (Wahlenb.) Mack., Carex capitata Soland., C. micro-
glochin Wahlenb., C. bicolor All., C. glacialis Mack., C. misandra R.Br.,

186 mMADRONO [Vol. 20

Juncus biglumis L., Luzula confusa Lindeb., L. spicata (L.) D.C. To-
fieldia pusilla (Michx.) Pers., Salix reticulata L., Oxyria digyna (L.)
Hill, Polygonum viviparum L., Sagina saginoides (L.) Karst., Minuartia
rubella (Wahlenb.) Graebn., Silene acaulis L., Melandrium apetalum
(L.) Fenzl., Ranunculus hyperboreus Rottb., R. nivalis L., Thalictrum
alpinum L., Cardamine bellidiflora L., Draba nivalis Liljebl., D. flad-
nizensis Wulf., Ervsimum pallasu (Pursh) Fern., Saxifraga oppositifolia
L., S. flagellaris Willd., S. nivalis L., S. caespitosa L., Parnassia palustris
L., Sibbaldia procumbens L., Astragalus eucosmus Robins., A. alpinus
L., Oxytropis campestris (L.) D.C., Epilobium latifolium L., E. horne-
mannii Rchb., Rhododendron lapponicum (L.) Wahlenb., Lotseleuria
procumbens (L.) Desv., Phyllodoce caerulea (L.) Bab., Arctostaphylos
alpina (L.) Spreng., Diapensia lapponica L., Pedicularis sudetica Willd.,
Pinguicula vulgaris L., Campanula uniflora L., Achillea borealis Bong.,
Arnica alpina (L.) Olin, Taraxacum ceratophorum (Ledeb.) D.C., and
Crepis nana Richards.

CIRCUMALPINE

A number of plants are predominantly alpine in distribution, and
not essentially arctic, although occasionally they are found in mountain-
ous parts of the arctic. These species occur in many mountain ranges
throughout the Northern Hemisphere, sometimes extending to edaphic-
ally suitable sites associated with cliffs, canyons, and river gorges. It is
possible that a number of the bryophytes may ultimately prove to be
arctic-alpine in distribution, but current information would place them
in the present category.

Hepatics: Haplomitrium hookeri (Sm.) Nees., Jungermannia cordi-
folia Hook., Nardia compressa (Hook.) Gray, Tritomaria scitula (Tayl.)
Joerg., Cephalozia leucantha Spr., Hygrobiella laxiflora (Hook.) Spr.,
Marsupella brevissima (Dumort.) Grolle, and Gvmnomitrion obtusum
(Lindb.) Pears.

Mosses: Oreas martiana (Hoppe & Hornsch.) Brid., Aongstroemia
longipes (Sommerf.) B.S.G., Oligotrichum hercynicum (Hedw.) Lam.
& D.C., Grimmia mollis B.S.G., Oedipodium griffithianum (Dicks.)
Schwaegr., Hygrohypnum smithii (Swartz.) Broth., H. alpestre (Hedw.)
Loeske, Calliergon stramineum (Wahl.) Kindb., Brachythecium tur-
gidum B.S.G., and B. collinum (Schleich.) B.S.G.

Vascular Plants: Athyrium distentifolium Tausch., Cystopteris mon-
tana (Lam.) Bernh., Vahlodea atropurpurea (Wahlenb.) Fries, Lloydia
serotina (L.) Rchb., Sagina saginoides (L.) Karst., Anemone narcissi-
flora L., Thalictrum alpinum L., Sibbaldia procumbens L., Myosotts
alpestris Schmidt, Aster alpinus L., Senecio fuscatus (Jord. & Fourr.)
Hayek., and S. resedifolius Less.

1969] SCHOFIELD: PHYTOGEOGRAPHY 187

Many of the bryophytes are widespread in mountainous western
North America and absent from Eastern North America: MJoerckia
blyttu, Nardia compressa, Hygrobiella laxiflora, Oreas martiana, Aong-
stroemia longipes, etc., which is the case also for several vascular plants:
Lloydia serotina, Mvyosotts alpestris, Senecio fuscatus, and Aster alpinus.

DISCONTINUOUS DISTRIBUTIONS

In the flora of northwestern North America there are several striking
disjunct elements. For most local disjunctions the details are presently
not apparent, particularly in the bryoflora. Only further collections will
expose these if they do exist. For the more dramatic disjunctions, how-
ever, the evidence is clear and, in many cases, the species involved are
environmentally restricted. Thus the western European disjuncts in west-
ern North America are predominantly confined to oceanic environments
and are unlikely to be found across North America since the environ-
ment is unavailable there. The situation for coastal and semi-arid ele-
ments with affinities in southern South America is similar.

Western American Bicentric Alpine

A number of species, independent of their gross distributional pat-
tern, show a disjunction within western North America suggesting that
in this geographic area, at least, the Pleistocene glaciations eradicated
the intervening portions of their range, leaving only those portions that
survived and later expanded outward from their glacial refuges. Since
suitable habitats are available in the intervening areas it must be
assumed that the species are in some way prevented from merging the
two western American fragments of their distribution. All species show-
ing this pattern are alpine; they are segregated here under their general
distributional element. Weber (1965) has discussed this disjunction for
the Southern Rocky Mountains.

1. Arctic-alpine: Alopecurus alpinus Sm., Poa vaseyochloa Scribn.,
Salix polaris Wahlenb., Minuartia biflora (L.) Schinz. & Thell., Sax-
fraga hirculus L., S. foliolosa R.Br., and Gentiana tenella Rottb.

2. Endemic Western American alpine: Poa nevadensis Vasey, Salix
dodgeana Rydb., Silene douglasii Hook., Draba densifolia Nutt., Arabis
lemmonii Wats., Potentilla virgulata Nels., Phlox hoodu Richards.,
Townsendia hookeri Beaman, Erigeron pumilus Nutt., and Artemisia
cana Pursh.

3. Circumarctic: Phippsia algida (Soland.) R.Br., Carex rupestris
All., and Draba fladnizensis Wulf.

4. Circumalpine: Swertia perennis L.

5. Eurasia—Western American: Silene repens Patrin., Anemone
narcissiflora L., Viola biflora L., and Gentiana algida Pall.

188 MADRONO [Vol. 20

6. Asia—Western American: Kobresia sibirica Turcz., Ranunculus
gelidus Karel & Kiril, Smelowskia calycina (Steph.) Mey., Bupleurum
triradiatum Adams, and Androsace filiformis Retz.

Details of bryophyte distributions are presently insufficient to de-
termine whether this distribution pattern is followed by these plants.
Affinities with Asia

This floristic element has probably received more attention than any
other in the geographic region under consideration. Gray (1859) had
noted the relationships in the vascular flora and these have been treated
in greater detail by Hultén (1928; 1937), Hara (1939), Li (1952(,
Tatewaki (1963) and briefly by Schofield (1965). The bryoflora has
been considered in greatest detail by Persson (1946a; 1946b; 1947;
1052a; 1952b; 1958; 1962; 1963). Other discussions have been by
Holzinger & Frye (1921), Persson and Gjaervoll (1957; 1961), Persson
and Weber (1958), Steere (1959), Steere and Schofield (1956), Steere
and Schuster (1960), Schofield (1965; 1966; 1968a; 1968b) and
Iwatsuki and Sharp (1967; 1968). Other floristic treatments are also
included in the literature cited, and it is from these that the following
details have been derived.

The Asiatic affinities can be segregated into several distinct elements:
Amphi-—Beringian, North Pacific, East Asian—North American and
Eurasian—Western American. Further subdivisions could be made,
particularly in the vascular flora. In the bryoflora, however, even many
of the above categories are not clearly demonstrable.

Amphi-Beringian

In this element are included species found on both sides of the Bering
sea, extending into Siberia on the Asian side and into Alaska in North
America. Some species found in China have also be included. In all
cases the distribution appears to expand both eastward and westward
from the Bering Sea area.

Hepatics: Pseudolepicolea fryei Perss. (Grolle & Ando), Ascidota ble-
blepharophyvlla Mass., and Radula prolifera Arnell.

Pseudolepicolea fryei is also found in a single locality on the west
coast of Hudson Bay (Schuster, 1966).

Mosses: Gollania turgens (C. Mull.) Ando might be placed here, al-
though its distribution is in mountains of Alaska and locally in China.

Vascular Plants:

Selaginellaceae: Selaginella siberica (Milde) Huron.

Poaceae: Agrostis trinit Turcz., Calamagrostis holmiu Lange, Koeleria
asiatica Domin., Poa lanata Scribn. & Merr., P. malacantha Kom., P.
pseudoabbreviata Roshev., Colpodium wrightu Scribn. & Merr., Pucci-
nellia borealis Swallen, P. geniculata (Turcez.) Krecz., and Agropyron
macrourum (‘Turcz.) Drobov.

1969] SCHOFIELD: PHYTOGEOGRAPHY 189

Cyperaceae: Carex lugens Holm., C. podocarpa Clarke, and C. neso-
phila Holm.

Juncaceae: Luzula rufescens Fisch. and L. tundricola Gorodk.

Salicaceae: Salix phlebophylla Anderss., S. rotundifolia Trautv., S.
sphenophylla Skvortz., S. fuscescens Anderss., S. ovalifolia Trautv., S.
chamissonis Anderss., and S. pulchra Cham.

Polygonaceae: Rumex arcticus Trautv. and R. stbiricus Hult.

Portulacaceae: Claytonia tuberosa Pall., C. acutifolia Pall., and C.
sarmentosa Mey.

Caryophyllaceae: Cerastium maximum L., C. jenisejense Hult., Minu-
artia arctica (Stev.) Aschers. & Graebn., M. yukonensis Hult., A. cha-
missonis Maguire, Wilhelmsia physodes (Fisch.) McNeill, Melandrium
taylorae {Robins.) Tolm., and M. taimyrense Tolm.

Ranunculaceae: Delphinium brachycentrum Ledeb. and Aconitum
delphinifolium D.C.

Fumariaceae: Corydalis pauciflora (Steph.) Pers.

Cruciferae: Cardamine hyperborea Schulz, C. microphylla Adams,
C. purpurea C. & S., Draba caesia Adams, D. eschscholtzu Pohle, D.
pilosa D.C., D. pseudo pilosa Pohle, D. stenopetala Trautv., D. kamtscha-
tica (Ledeb.) Bush., and D. chamissonis Don.

Saxifragaceae: Saxifraga eschscholtzu Sternb., S. serpyllifolia Pursh,
S. exilis Steph., S. nudicaulis Don., S. davurica Willd., S. unalaschensis
Sternb. and Chrysosplenium wrighti Fr. & Sav.

Rosaceae: Spiraea beauverdiana Schneid. and Potentilla elegans C.&S.

Leguminosae: Astragalus umbellatus Bunge, Oxytropis mertensiana
Turcz., and O. arctica R.Br.

Umbelliferae: Cnidium ajanense (Regel & Tiling) Drude and C.
cnidiufolium (Turcz.) Schischk.

Primulaceae: Primula tschuktschorum Kjellm., P. cunetfolia Ledeb.,
P. borealis Duby, Douglasia ochotensis (Willd.) Hook., and Dode-
catheon frigidum C. &S.

Gentianaceae: Gentiana barbata Froel. and G. glauca Pall.

Polemoniaceae: Phlox sibirica L., Eritrichium aretioides (Cham.)
D.C., and E. chamissonts D.C.

Scrophulariaceae: Lagotis glauca Gaertn., Castilleja elegans Malte,
C. caudata (Pennell) Rebr., and C. hyperborea Pennell.

Orobanchaceae: Boschniakia rossica (C. & S.) Fedtsch.

Plantaginaceae: Plantago canescens Adams.

Valerianaceae: Valeriana capitata Pall.

Compositae: Artemisia globularia Bess., A. glomerata Ledeb., A.
senjavinensis Bess., A. laciniatiformis Kom., A. furcata Bieb., Arnica
lessingit Greene, A. frigida Mey., Senecio atropurpureus (Ledeb.)
Fedtsch., Saussurea nuda Ledeb., S. viscida Hult, Taraxacum lateritium
Dahlst., and T. kRamtschaticum Dahlst.

190 MADRONO [ Vol. 20

North Pacific

In this category are placed species that range around the North
Pacific Basin. In most cases the species do not extend into continental
regions. It seems likely that many of these species did not expand their
range via the Bering land bridge, but by the Aleutian Chain. In other
cases these species appear to be ancient relict populations of Tertiary
times. Several species persist in regions where they survived the Pleisto-
cene and preceding glaciations. In others of wider range, the species
have expanded since glaciation, but from their refugia on either side
of the Pacific.

Hepatics: Takakia lepidozioides Hatt. & Inoue, 7. ceratophylla
(Hook.) Grolle, Herberta himalayana (Steph.) Miller, Ptilidium cali-
fornicum (Aust.) Underw., Bazzania ambigua Lindenb., Lepidozia fila-
mentosa (Lehm. & Lindenb.) Gottsche, Lindenb., & Nees, Chandonan-
thus hirtellus (Web.) Mitt., C. pusillus Steph., Gymnomitrion pacificum
Grolle, Macrodiplophyllum plicatum (Lindb.) Perss., M. microdontum
(Mitt.) Perss., Scapania bolanderi Aust., Plagiochila satoi Hatt., P.
rhizophora Hatt., P. semidecurrens Lehm. & Lindenb., Porella vernicosa
Lindb., Radula obtusiloba Steph., R. auriculata Steph., Cololejeunea
macountu (Spruce) Evans, and A potreubia nana (Hatt. & Inoue) Hatt.
& Mizut.

Mosses: Sphagnum guwassanense Warnst., S. subobesum Warnst.,
Oligotrichum parallelum (Mitt.) Kindb., O. aligerum Mitt., Bartramiop-
sis lyellit (James) Kindb., Pogonatum laterale (Brid.) Brid., Pohlia
columbica (Kindb.) Andr., Trachycystis flagellaris (Sull. & Lesq.)
Lindb., Rhizomnium nudum (Williams) Koponen, Ulota japonica (Sull.
& Lesq.) Mitt., U. repens Mitt., Climacium japonicum Lindb., Pleuro-
210 psis ruthenica (Weinm.) Kindb., Bryknia hulteni Bart., Mvuroclada
maximowiczu (Borosz.) Steere & Schof., Campylium adscendens (Lindb.)
Perss., Hypnum subimponens Lesq., H. dieckit Ren. & Card., Clao podium
crispifolium (Hook.) Ren. & Card., C. pellucinerve (Mitt.) Best., Les-
curaea julacea Besch. & Card., Hypoptervgium fauriei Besch., and
Habrodon leucotrichus (Mitt.) Perss.

Vascular Plants (those marked with an * are maritime): Mecodium
wright (Bosch.) Copeland, Deschampsia beringensis Hult., Poa macro-
calyx Trautv. & Mey.,* Puccinellia pumila (Vasey) Hitchc.,* P. hulteniu
Swallen,* P. kamtschatica Holmb.,* Carex macrocephala Willd.,* C.
elusinoides Turcz., C. ramenskii Kom., C. gmelini H. & A.,* C. macro-
chaeta Mey., C. spectabilis Wew., Juncus ensifolius Wikstr., J. merten-
sianus Bong., Fritillaria camschatcensis (L.) Ker.-Gawl., Maianthemum
dilatatum (How.) Nels. & MacBr., Streptopus streptopoides (Ledeb.)
Frye & Rigg., Dactylorhiza aristata (Fisch.) Soo., Platanthera conval-
larufolia (Fisch.) Lindb., P. chorisiana (Cham.) Rchb., Atriplex gmelini

1969] SCHOFIELD: PHYTOGEOGRAPHY 191

Mey.,* Stellaria ruscifolia Pall., Cerastium fischerianum Ser., Sagina
crassicaulis Wats.,* Minuartia macrocarpa (Pursh) Ostenf., Aconitum
maximum Pall., Ranunculus eschscholtzu Schlecht., Oxygraphis glacialis
(Fisch.) Bunge, Papaver alboroseum Hult., Cardamine umbellata
Greene, Draba borealis D.C., D. hyperborea (L.) Desv., Saxifraga
bronchialis L., S. bracteata Don, Rubus pedatus Sm., R. spectabilis
Pursh, Geum calthifolium Menzies, G. rossit (R.Br.) Ser., G. pentapeta-
lum (L.) Makino, Sanguisorba stipulata Raf., Geranium erianthum
D.C., Viola langsdorfu Fisch., Epilobium behringianum Haussk., E.
sertulatum Haussk., Angelica genuflexa Nutt., Rhododendron camtschatt-
cum Pall., Phyvllodoce aleutica (Spreng.) Heller, Cassiope stelleriana
(Pall.) D.C., C. lycopodioides (Pall.) Don, Fauria crista-galli (Menzies)
Makino, Plagiobothrys orientalis (L.) Johnston, Pentstemon fruticosus
(Pursh) Greene, Veronica stellerit Pall., Euphrasia mollis (Ledeb.)
Wettst., Pedicularis chamissonis Stev., Pinguicula macroceras Link., and
Hieracium triste Willd.

A number of vascular plant genera are present only in East Asia and
Western North Amreica, but are represented by different species in
each of the areas: Pseudotsuga, Phyllospadix, Lysichiton, Castanopsis,
Achlys, and Echinopanax.

East Asia—North American

A number of species of vascular plants are widespread in North Amer-
ict, particularly in boreal and arctic regions and extend into the eastern
portion of Asia, occasionally westward nearly to Europe. The only bryo-
phyte of comparable range appears to be the moss Hypnum plicatulum
(Lindb.) Jaeg. & Sauerb.

Vascular Plants: Lycopodium obscurum L., Hierochloe pauciflora
R.B., Calamagrostis purpurascens R.Br., Danthonia intermedia Vasey,
Schizachne purpurascens (Torr.) Swallen, Bromus ciliatus L.,Elymus
mollis Trin., Eriophorum callitrix Cham., Carex stipata Muhl., C. viridula
Michx., C. membranacea Hook., Smilacina trifolia (L.) Desf., Salix
fuscescens Anderss., S. alaxensis (Anderss.) Cov., S. depressa L., Alnus
crispa (Ait.) Pursh, Stellaria longipes Goldie, S. edwardsit R.Br., Ceras-
tium beeringianum C. &S., Brasenia schrebert Gmel., Caltha natans, Pall.,
Coptis trifolia (L.) Salisb., Anemone richardsoni Hook., A. parviflora
Michx., Rorippa hispida (Desv.) Britt., Lesquerella arctica (Wormsk.)
Wats., Arabis lyrata L., A. drummondiu Gray, A. divaricarpa Nels., A.
holboelli Hornem., Mitella nuda L., Parnassia kotzebuei C. & S., Ribes
lacustre (Pers.) Poir., R. triste Pall., Geum macrophyllum Willd., Oxy-
tropis nigrescens (Pall.) Fisch., Stwm suave Walt., Conioselinum chinense
(L.) B.S.G., Angelica lucida L., Heracleum lanatum Michx., Cornus
canadensis L., Pyrola asarifolia Michx., Monotropa uniflora L., Arcto-
staphylos rubra (Rehd. & Wilson) Fern., Lycopus lucidus Turcz., L.
uniflorus Michx., Veronica americana Schwein., Pedicularis labradorica
Wirsing., P. langsdorfu Fisch., P. capitata Adams., Galium kamtchaticum

192 MADRONO [Vol. 20

Steller., Antennaria friesiana (Trautv.) Ekman, Anaphalis margaritacea
(L.) B. & H., Artemisia frigida Willd., Petasites palmatus (Ait.) Gray,
Senecio resedifolius Less., and S. pseudo-arnica Less.

Eurasia—Western America

A number of species are widespread through both Asia and Europe,
either in the arctic or in boreal regions, sometimes both, and extend into
western North America. In most cases these plants do not extend east of
the Rocky Mountains, but in some cases reach the west coast of Hudson
Bay or the Great Lakes region. These are represented by only vascular
plants.

Cryptogramma crispa (L.) R.Br., Thelypteris limbosperma (All.)
Fuchs., Ruppia spiralis L., Agrostis clavata Trin., Scolochloa festucacea
(Willd.) Link., Carex obtusata Lilj., C. pyrenaica Wahlenb., C. lapponica
Lang., C. rhynchophysa Mey., Cypripedium guttatum Sw., Hammarbya
paludosa (L.) Ktze., Salix hastata L., Rumex graminifolius Lamb, Silene
repens Patrin, Anemone narcissiflora L., Pulsatilla patens (L.) Mill.,
Thalictrum minus L., Aruncus sylvester Kostel, Hedysarum hedysaroides
(L.) Schinz & Thell., Zimpatiens noli-tangere L., Viola biflora L., V.
epipsila Ledeb., Ligusticum mutellinoides (Crantz.) Willar, Primula
sibirica Jacq., Androsace chamaejasme Host, A. filiformis Retz., Trien-
talis europaea L., Gentiana algida Pall., G. prostrata Haenke, Swertia
perennis L., Polemonium acutiflorum Willd., P. boreale Adams, M yosotis
alpestris Schmidt, Pedicularis verticellata L., P. oederi Vahl, Aster alpinus
L., A. sibiricus L., Artemisia laciniata Willd., Petasites frigidus (L.)
Franck., and Senecio fuscatus (Jord. & Fourr.) Hayek.

Western North America—Southern Hemisphere Disjunctions

In the western North American flora two different discontinuities are
exhibited by species that reappear in the Southern Hemisphere: bi-polar
disjuncts and Pacific North American—South American disjuncts.

Bipolar Disjuncts

Du-Rietz (1940) has thoroughly discussed the problem of bipolar
plant distribution, summarizing both pertinent literature and basic in-
formation. A bipolar disjunct pattern is that in which species occur in
the temperate Northern Hemisphere, and again in the temperate South-
ern Hemisphere but are essentially absent from tropical latitudes.

To explain this pattern DuRietz (1940) concluded that “it seems
necessary to look for epeirogenetic transtropical highland bridges older
than the mountain-chains of the Alpine Orogen. Such highland bridges
may have existed not only in Africa, but also bordering the transtropical
Alpine geosynclines (i.e. the Andean and the Malaysian geosynclines),
partly passing over present deep sea bottom.”

In many cases the plants of this disjunction are circumboreal in the
Northern Hemisphere, several being ubiquitous through that range.

1969] SCHOFIELD: PHYTOGEOGRAPHY 193

Sainsbury (1952) briefly discussed some of the mosses of New Zealand
that showed this distributional pattern. The discussions of Martin (1946;
1949; 1952a; 1952b) have also contributed to the understanding of this
disjunction.

Hepatics: Fossombronia pusilla (L.) Dum., Metzgeria furcata (L.)
Dum., Moerckia blytti (Moerch.) Brockm., Anthelia juratzkana
(Limpr.) Trevis, Ptilidium ciliare (L.) Hampe., Barbilophozia hatcheri
(Evans) Loeske, Jungermannia cordifolia Hook., Orthocaulis floerket
(Web. & Mohr.) Buch., and Diplophyllum obtusifolium (Hook.) Du-
mort.

Mosses: Sphagnum centrale C. Jens., S. fimbriatum Wils., S. magel-
lanicum Brid., S. palustre L., S. papillosum Lindb., S. subnitens Russow
& Warnst., Andreaea rupestris Hedw., Tetrodontium brownianum (Dicks.)
Schwaegr., Pogonatum alpinum (Hedw.) Rohl., Polytrichum formosum
Hedw., Buxbaumia aphylla Hedw., Fissidens adianthoides Hedw.,D1-
trichum flexicaule (Schwaegr.) Hampe, Saelania glaucescens (Hedw.)
Broth., Dicranum scoparium Hedw., Pottia heimu (Hedw.) Furnr.,
Desmatodon convolutus (Brid.) Grout, Tortula muralis Hedw., T. papil-
losa Wils., T. laevipila (Brid.) Schwaegr., Encalypta vulgaris Hedw.,
Grimmia donniana Sm., G. trichophylla Grev., Racomitrium lanuginosum
(Hedw.) Brid., Funaria microstoma B.S.G., Tetraplodon mnioides
(Hedw.) B.S.G., Bryum angustirete Kindb., B. pseudotriquetrum
(Hedw.) Gaertn., Mey. & Scherb., B. caespiticium Hedk., B. micro-
erythrocarpum C,. Mull. & Kindb., Aulacomnium palustre (Hedw.)
Schwaegr., Bartramia halleriana Hedw., B. pomiformis Hedw., B. ithy-
phylla Brid., Orthotrichum alpestre Hornsch., Climacium dendroides
(Hedw.) Web. & Mohr., Neckera pennata Hedw., Campylium poly-
gamum (B.S.G.) Jens., Leptodictyon riparium (Hedw.) Warnst., Drepan-
ocladus uncinatus (Hedw.) Warnst., Calliergon cordifolium (Hedw.)
Kindb., C. sarmentosum (Wahlenb.) Kindb., Calliergonella cuspidata
(Ren.) Grout, Brachythecium albicans (Hedw.) B.S.G., B. plumosum
(Hedw.) B.S.G., B. rutabulum (Hedw.) B.S.G., B. salebrosum (Web. &
Mohr) B.S.G., B. velutinum (Hedw.) B.S.G., Eurhynchium praelongum
(Hedw.) B.S.G., Pleurozium schreberi (Brid.) Mitt., Plagiothecium
denticulatum (Hedw.) B.S.G., P. roeseanum B.S.G., Hypnum revolutum
(Mitt.) Lindb.,Jsopterygium pulchellum (Hedw.) Jaeg. & Sauerb., and
H ylocomium splendens (Hedw.) B.S.G.

Vascular Plants: Botrychium lunaria (L.) Sw., Pteridium aquilinum
(L.) Kuhn., Asplenium trichomanes L., Potamogeton filiformis Pers., P.
praelongus Wulf., P. natans L., Triglochin palustris L., Vahlodea atro-
purpurea (Wahl.) Fries., Carex buxbaumi Wahl., C. capitata Soland.,
C. canescens L., C. diandra Schrank, C. lachenalii Schkuhr., C. magel-
lanica Lam., C. microglochin Wahl., C. pyrenaica Wahl., Juncus filiformis
L., Koenigia islandica L., Chenopodium glaucum L., Montia fontana L.,

194 MADRONO [Vol. 20

Honkenia peploides (L.) Ehrh., Anemone multifida Poir, Cochlearia
officinalis L., Gentiana prostrata Kaenke, and Hieracium gracile Hook.

Pacific North American—South American Disjuncts

In arid and coastal areas of Pacific North America are a number of
species that reappear again in South America in Argentina and Chile,
generally as elements of the same environment. These species have at-
tracted the attention of a number of botanists: Gray and Hooker (1880),
Bray (1898; 1900), Campbell (1944), Campbell and Wiggins (1947),
Raven and Lewis (1959), and Cruden (1966). In a symposium concern-
ing this disjunction, Raven (1963) provided the summary. In the same
publication were detailed studies of particular species by Chambers
(1963), Constance (1963), Heckard (1963), and Ornduff (1963).

No information concerning bryophytes was included and the details
concerning the bryoflora are inadequate to make any valid generalizations.

Raven (1963) suggests the following theory to explain this disjunc-
tion: “The great majority of the plants reached their disjunct areas by
long-distance dispersal relatively recently. For the bipolar species, the
Pleistocene seems the most likely time of dispersal, for the temperate
species, the late Pliocene or Pleistocene, and for the desert species, ex-
cluding those that have differentiated from common ancestors that
spanned the tropics, no time has probably been more likely than the
recent past. Both bipolar and temperate disjuncts have come mostly from
the north and are almost entirely herbaceous. The desert disjuncts, on
the other hand, often appear to have originated in the south, or have
diverged from a common tropical ancestor. Many of them are woody.”

As was noted earlier, DuRietz (1940) did not hold this opinion.
Cruden (1966) also suggests another alternative, noting that for the
examples given by Raven, birds could not have served as the dispersal
agents and no other agency is likely for such great distances. He sug-
gests that birds, other than shorebirds, may have been important in
stepwise dispersal of the species for relatively short distances. ‘“‘Moun-
tain hopping provides a reasonable explanation for the movement of a
large segment of the parental gene pool across the tropics through the
buildup of large intermediate populations.”

Unfortunately such mountains have not been available during the
time suggested and one would be forced to imply a change in the
ecology of the species during “migration” and reversion to the original
ecological requirements on “arrival.”

Species exhibiting this disjunction are (those marked * are essen-
tially maritime): Palleae atropurpurea (L.) Link, Lilaea_ scilloides
(Poir.) Haum., Triglochin concinna Burtt—Davy,* Agrostis idahoen-
sis, Nash, Bromus trinii Desv., Danthonia californica Boland., Des-
champsia danthonioides (Trin.) Munro, D. elongata (Hook.) Munro,
Festuca megalura Nutt., Poa stenantha Trin., P. secunda Pres, Carex

1969] SCHOFIELD: PHYTOGEOGRAPHY 195

praegracilis Boott, Scirpus cernuus Vahl.,* S. nevadensis Pers., Juncus
leseurtt Boland,* Calandrinia ciliata (R. & P.) D.C., Oxytheca dend-
roidea Nutt., Polygonum punctatum EIl., Cardionema ramosissima
(Weinm.) Nels. & Macbr.,* Anemone multifida Poir., Myosurus apeta-
lus Gay, Lepidium nitidum Nutt., Fragaria chiloensis (L.) Duch.,*
Trifolium macraei H. & A., T. microdon H. & A., Boisduvalia glabella
(Nutt.) Walpers, Gavophytum humile Juss., G. diffusum T. & G.,
Osmorhiza chilensis H. & A., O. depauperata Phil., Sanicula crassicaulis
Poepp., S. graveolens Poepp., Microsteris gracilis (Hook.) Greene,
Polemontum micranthum Benth., Coldenia nuttallii Hook., Cryptantha
circumscissa (H. & A.) Johnst., Heliotropium curassavicum L., Lappula
redowskiu (Hornem.) Greenm., Plectocarya linearis (R. & P.) D.C.,
Plagtobothrys scoulert (H. & A.) Johnst., Veronica peregrina L., Plan-
tago patagonica Jacq., Convolvulus soldanella L.,* Ambrosia chamis-
sonis (Less.) Greene,* Madia gracilis (Sm.) Keck., M. sativa Mol., and
Psilocar phus brevissimus Nutt.

Western North America—W estern Europe

Most of the species of this element are oceanic in their distribution
although a number are alpine. In both cases they appear to be per-
sistent remnants of a circumboreal flora, possibly dating back as early
as Tertiary time. To imply long-distance dispersal from Europe is illogi-
cal since suitable habitats for the species also exist in eastern North
America, but the species do not occur there. Evans (1900), and Persson
(1949) have discussed this element in the hepatics in particular, and
Schofield (1965; 1968a; 1968b) has considered all bryophytes. The
vascular flora of this disjunction was briefly discussed by Schofield
(1965). Several of the species are widespread and abundant in both
parts of their range, others are highly restricted.

Hepatics: Herberta straminea (Dumort.) Trevis, Mastigophora
woodsit (Hook.) Nees., Bazzania pearsoni Steph., Cephaloziella phylla-
cantha (Mass. & Carest.) Mull., C. turnert (Hook.) Muull., Anastrepta
orcadensis (Hook.) Schiffn., Anastrophyllum donianum (Hook.) Spr.,
A. assimile (Mitt.) Steph., Gymnocolea acutiloba (Kaal.) Miull., Junger-
mannia caespiticia Lindenb., Plagiochila major (Nees.) Arnell, Diplo-
phyllum obtusifolium (Hook.) Dum., Scapania scandica (Arn. & Bach.)
MacVicar, Marsupella alpina (Gottsche.) Bernet., M. brevissima (Du-
mort.) Grolle, M. commutata (Dumort.) Grolle, Pleurozia purpurea
(Lightf.) Lindb., Porella cordaeana (Hueb.) Evans., Metzgeria fruti-
culosa (Dicks.) Evans., Moerckia blyttii (Moerch.) Brockm., and
Bucegia romanica Radian.

Among the hepatics, and several of the mosses, a number of species

are found in scattered localities in mountainous Japan and in the Him-

alayas.

196 MADRONO [Vol. 20

Mosses: Andreaea nivalis Hook., Ditrichum zonatum (Brid.) Kindb.,
Cynodontium jenneri (Schimp.) Stirt., Kiaeria falcata (Hedw.) Hag.,
Dicranum tauricum Sapeh., D. spadiceum Zett., Dicranodontium unci-
natum (Harv.) Jaeg., Campylopus schwarz Schimp., C. schimperi
Milde, C. subulatus Milde, Paraleucobryum enerve (Thed.) Loeske,
Encalypta affinis Hedw., f., E. longicollis Bruch., Leptodontium recurvi-
folium (Tayl.) Lindb., Barbula vinealis Brid., Geheebia gigantea
(Funck.) Boul., Pottza lanceolata (Hedw.) Mull., Tortula subulata
Hedw., T. laevipila (Brid.) Schwaegr., T. latifolia (Spreng.) Hartm.,
T. princeps DeNot., Grimmia pulvinata (Hedw.) Sm., G. decipiens
(Schultz.) Lindb., G. hartmanniu Schimp., Micromitrium tenerum (B.S.
G.) Crosby, Funaria muhlenbergu Hedw. f., Tavloria hornschuchiana
(Gre. & Arn.) Lindb., T. froelichiana (Hedw.) Mitt., Pohklia erecta
Lindb., P. vexans (Limpr.) Lindb. f., P. gracilis (B.S.G.) Lindb.,;
Epipterygium tozeri (Grev.) Lindb., Bryum miniatum Lesq., B. canari-
ense Brid., Bartramia halleriana Hedw., Zygodon rupesiris (C. Hartm.)
Milde, Z. gracilis Wils., Orthotrichum rupestre Schleich., O. laevigatum
Zett., O. speciosum Nees., O. rivulare Turn., O. cupulatum Brid., O.
alpestre Hornsch., O. tenellum Bruch., O. pulchellum Brunt., Antitrichia
curtipendula (Hedw.) Brid., Pterogonium gracile (Hedw.) Sm., Neckera
menziesu Hook., Hookeria lucens (Hedw.) Sm., Fabronia pusilla Raddi,
Lescuraea stenophylla (Ren. & Card.) Kindb., Drepanocladus tricho-
phyllus (Warnst.) Podp., Hvgrohypnum molle (Hedw.) Loeske, Cal-
liergon megalophyllum Mik., Brachythecium trachypodium (Funch)
B. & S., B. tromsoense (Kaur. & Arn.) Limpr., Scleropodium caes pitans
(Mull.) L. Koch., S. tourette: (Brid.) Koch., Plagiothecium piliferum
(Sw.) B.S.G., P. platyphyllum Monk., and P. undulatum (Hedw.)
B.s.G.

Vascular Plants: Equisetum telmateia Ehrh., Blechnum spicant (L.)
Roth, Thelypteris oreopteris (Ehrh.) Slosson, Carex stenophylla
Wahlenb., C. foetida All., and Saxifraga adscendens L., also the vicariant
species (the European in parentheses): Anemone drummondu Wats.
(A. baldensis L.), Pulsatilla occidentalis (Wils.) Freyn. (A. alpinus
(L.) Debartre), Trifolium nanum Torr. (T. alpinum L.), and Astragalus
goniatus Nutt. (A. danicus Retz.).

Cordilleran Disjuncts in Eastern America

Fernald (1924; 1925), in his discussion of vascular plant distribution
in northeastern North America, noted a conspicuous element of western
North American affinity. Many of these species are North American
endemics, while others show a broken circumboreal distribution, or in
some cases, affinities with Asia. In all of these, however, there is a
marked disjunction between northeastern North America and Western
North America. In many the western North American portion of the
distribution does not extend beyond the Rocky Mountain chain.

1969] SCHOFIELD: PHYTOGEOGRAPHY 197

Since the publication of the earlier papers, Fernald (1926; 1933);
1935; 1942) published the results of his considerable field-work further
emphasizing this element. His explorations concentrated on areas in
eastern North America that had previously yielded disjuncts and in
which he located further taxa of this distribution pattern. These regions
included Newfoundland, Gaspé and the shores of Lake Superior. Most
frequently these plants occur in river canyons or at high elevations.
Thus, many of the species appear to be part of a broken circumalpine
or circumboreal distribution, or are species endemic to mountains of
North America. The contributions of Marie-Victorin (1935; 1938), in
his explorations of the islands of the St. Lawrence River, Anticosti and
Mingan Islands, contributed further information concerning this dis-
junction. Abbe (1936) briefly discussed cordilleran disjuncts in Labra-
dor peninsula, noting that they are few in number and that in many the
ranges are not as interrupted as was previously assumed.
Further discussions of this element are those of Stebbins (1935),
Wynne-Edwards (1937; 1939), Griggs (1940), Raymond (1950), Scog-
gan (1950), Bocher (1951), Butters & Abbe (1953), Rousseau (1953),
Rune (1953; 1954), Dutilly, Lepage and Duman (1958), Schuster
(1958), and Schofield (1959). The check-list of Newfoundland vascular
plants of Rouleau (1956) is also a valuable source of basic information,
as is that of Porsild and Cody (1968) for Mackenzie District.
Explanations for this disjunction are rather varied. The basic prob-
lems have been threefold: the inadequacy of details of glaciation in
the pertinent areas in eastern America, the ecology of the disjuncts in
their eastern outposts, and the uncertainties concerning the total ranges
of the species due to inadequate collections from intervening areas.
Changes in the status of these avenues of information have led to
changes in the theory explaining the plant disjunctions.
Fernald (1925) felt that ‘Cordilleran” and particularly arctic-alpine
species in eastern North America had survived on nunataks during the
Pleistocene glaciations. The nunataks he considered to be essentially
coincident with the areas rich in these disjunct plants. Arguments against
this hypothesis have been strongest and most convincing concerning the

arctic-alpine elements. Wynne-Edwards (1937; 1939), Rousseau (1950)
-and Dammann (1965) have noted that these species could certainly
have immigrated into their present sites following glaciation and now
persist in only those microclimatic sites that are not invaded by the
| general flora that is more adapted to the macroclimate.

Stebbins (1935) suggested that each disjunct “‘migrated eastward in
_ post-glacial times, following near the front of the retreating ice-sheet.
| Widespread, though local, in its western distribution, it becomes rarer
' and rarer eastward, with a more limited range north and south, until at
its eastern limit there are (few) widely separated stations for it.”

_ Abbe (1936) to explain the presence of this element in Labrador,

——

198 MADRONO [Vol. 20

where the cordilleran taxa are largely now near sea-level suggested:
‘Possibly then, the balance of all factors were such that in areas, as at
the heads of fiords, protected from wind, warmed by occasional fohn
winds and by the action of direct insolation, and with melt-water avail-
able from the ice-fields above, plants may have survived through the
peak of Wisconsin glaciation in the lee of the Torngat Mountains in
northeastern Labrador.”

Wynne-Edwards (1937) suggested what is termed by Victorin (1938)
the “‘rainbow hypothesis”: “Some of these plants have wide limits of
climatic tolerance, occurring through a wide latitudinal range, in which
case their American distribution takes the form of an arch spanning the
continent from the Cordillera to the St. Lawrence by way of the Arctic,
while others are more confined, the hardiest occupying the crown of the
arch and least hardy its two ends, whereby their ranges are disrupted
into western and eastern centres.”

Griggs (1940), to explain the distribution of rare plants, suggested
that, for disjuncts: ‘“‘rare plants have been eliminated from the older
adjacent barren areas by competition with the more competent common
vegetation but persist in the refuges more recently opened to coloniza-
tion because the ecological succession there has not run quite so far as
elsewhere.” This would assume a continuity in the past distribution of
the disjunct species.

Rousseau (1950) summarizes his explanation as follows: ‘‘(a) some
(of the Cordilleran disjuncts) . . . could be indifferent arctic alpine
plants, now absent from the Arctic proper through elimination by var-
ious historical factors (b) The remaining species after this elimination
could be considered, at least hypothetically, as pre-glacial plants, though
this is not the only probable explanation (c) The remaining species
could, as well, be considered as simply alpine species, living on alpine
formations constituted after the recess of the glacier. The plants could
have taken shelter there during the “‘pre-sylvatic period,” after having
travelled from the Canadian Rockies to the Gaspé Peninsula, along
the “Arctic Corridor” bordering the receding continental glacier. During
glaciation these plants in the Rockies could have sought refuge either
south of the glacier or on nunataks. The relicts would then be preglacial
in the Rockies and postglacial in Gaspé.”

Gaspé has been explored bryologically by both F. Leblanc and J.
Kucyniak but the results have not yet been published. From the exten-
sive bryological collections made in Newfoundland, Tuomikoski has
published results only on the hepatics (Buch & Tuomikoski, 1955). In
this paper he notes only two “‘Cordilleran” hepatics: Frullania bolanderi
Aust. and Cephalozia catenulata (Hueb.) Spruce. He notes, however,
that a number of species show this disjunct pattern in North America
although they belong to the circumboreal element. These species are
widely distributed through eastern North American, extend westwards

1969] SCHOFIELD: PHYTOGEOGRAPHY 199

frequently to the Great Lakes and southward in the Appalachian Moun-
tains, but are absent until west of the Rocky Mountains in many cases.
They are therefore excluded here as “Cordilleran disjuncts” largely
because they represent a different distributional pattern to those dis-
cussed arlier.

Steere (1937; 1938) drew attention to the bryophytes to Cordilleran
disjunction in the Great Lakes areas. Schuster (1958) provided further
information for this region.

Of the hepatics cited by Steere (1937; 1938), Schuster (1958) sug-
gested that they could not be considered Cordilleran but were of much
wider distribution, being found in intervening areas. He suggested that
those that have not been collected in intervening areas are largely incon-
spicuous and will undoubtedly be found with further exploration.

It seems possible that the mosses may be similarly distributed, but
the presently known ranges would support Steere’s contention that they
are Cordilleran disjuncts: Racomitrium patens (Hedw.) Hiibn., Grim-
mia hartmanni Schimp., Lescuraea incurvata (Hedw.) Lawt., and
Hygrohypnum molle (Schimp.) Loeske. It should be noted that these
mosses are essentially circumboreal in their world-wide distribution.

The Cordilleran disjunct vascular plants can be divided into three
categories (those marked with an asterisk are maritime).

1. Endemic to North America: Chedlanthes siliquosa Maxon, Woodsia
oregana Eat., Polystichum scopulinum (Eat.) Maxon, Poa canbyi
(Scribn.) Piper, Festuca scabrella Torr., Agropyron violaceum (Hornem.)
Lange, Eleocharis nitida Fern., Carex filifolia Nutt., Goodyera oblongi-
folia Raf., Platanthera unalaschensis (Spreng.) Kurtz., Salix vestita
Pursh, Polygonum fowleri Robins.,* Salicornia virginica L.,* Minuartia
obtusiloba (Rydb.) House, Spergularia canadensis (Pers.) Don,* Dryas
drummondiu Richards., Vaccinium ovalifolium Sm., Campanula latise pala
Hult., and Arnica cordifolia Hook., A. mollis Hook., and Cirsium folio-
sum (Hook.) D.C.

2. East Asian—western American—FEastern American disjunct: Po-
tamogeton subsibiricus Hagstr., Poa eminens Presl,* Eleocharis kamt-
schatica (Mey.) Kom.,* Carex lyngbaei Hornem.,* C. stylosa Mey., C.
franklinu Boott, Juncus ensifolius Wikstr., Epilobium glandulosum
Lehm., Conioselinum chinense (L.) B.S.P., Angelica lucida L., Galium
kamtschaticum Steller, Arnica frigida Mey., Senecio resedifolius Less.,

and S. pseudo-arnica Less.

3. Essentially circumboreal, but in North America with western North

America and eastern America disjunction: Lycopodium inundatum L.,
Athyrium distentifolium Tausch., Cystopteris montana (Lam.) Bernh.,
Polystichum lonchitis (L.) Roth., P. braunii (Spenn.) Fee., Tricho-
phorum pumilum (Vahl.) Schinz. & Thell., Thalictrum alpinum L..,

Ligusticum scothicum L., and Cornus suecica L.
In summary, there is a marked disjunct Cordilleran element in the

200 MADRONO [Vol. 20

flora of eastern North America, centered largely around the Gulf of St.
Lawrence region and the Great Lakes. The species are largely alpine in
distribution and ecology although several are maritime. In most cases
they are restricted to sites relatively free from invasion by the species
that dominate the general local vegetation, and are essentially in sites
at a persistent pioneering phase.

The most plausible explanation of their disjunction is that the eastern
representatives are remnants of a more widespread flora of the past,
possibly of pre-Pleistocene arctic-alpine distribution in North America.
The Pleistocene glaciations can be assumed to have eliminated the north-
central portion of the range, but since habitats were available in north-
eastern and western North America the species survived, probably south
of the glacial boundary, but possibly in nunataks or coastal refuges,
moving to their present sites following retreat of the ice sheet but being
eliminated from their Pleistocene refugium by the encroaching vegeta-
tion and by a succession toward more mesophytic temperate vegetation.
Evidence for nunataks and refugia in eastern North America has been
disputed.

SUMMARY

The flora of northwestern North America is composed of a rich repre-
sentation of endemics, both in genera and species. Highest concentration
of these is in areas that are environmentally diverse and escaped the
Pleistocene glaciations. Although the glaciated areas also possess many
endemics, their flora tends to be dominated by circumboreal taxa, great-
est in Alaska and northern British Columbia, and decreasing southward.

That much of the western North American flora is a fragment of a
more widespread flora of Eurasian affinities appears indisputable. Why,
in many cases, these taxa have not extended their range across North
America, or remain as persistent remnants in suitable environments on
both sides of the continents, can probably never be adequately explained.
Reconstruction of the climatic and accompanying geologic events as
reflected by the records of past floras will considerably aid in the propo-
sition of a working hypothesis. However, new information will always
make necessary modification of the historical details that have led to
present distributions.

In vascular plants, in particular, the ranges of taxa have been greatly
altered by man’s activities. The introduction of exotic species of vascu-
lar plants has also greatly affected the species that formed the native
flora that preceded the advent of man. The effects of aboriginal man
on plant introductions have never been adequately studied.

In bryophytes such introductions are presumed to be relatively in-
frequent but man’s influence on distribution patterns has been drastic
in environments that are readily exploited by man. Since bryophytes
are frequently found in rather extreme environments and persist in
small microclimatic sites, they may remain while the more vulnerable

1969] SCHOFIELD: PHYTOGEOGRAPHY 201

vascular flora succumbs. Thus the bryophytes serve as a valuable tool
to interpret past floristic history. From the hypotheses derived from
bryophyte distributions one can extrapolate to vascular plant distri-
butions.

The bryoflora of western North America consists of a considerably
higher percentage of elements of circumboreal distribution than does the
vascular flora. The presence of remarkably wide disjuncts is also gener-
ally more conspicuous in the bryophytes than in the vascular flora. This
is emphasized by disjunction of species of western Europe—western
North America and also of a number of species of southeast Asia—west-
ern North America. Especially notable about these taxa is the fact that
most lack sexual reproduction and have no special means of asexual
reproduction. To explain their distribution based on chance long dis-
tance dispersal is hazardous and creates more problems than it solves.

Bipolar distributions and affinities with South America imply a past
continuity of floras, although attempts have been made to explain this
disjunction by long-distance dispersal.

The Cordilleran disjuncts in eastern North America are considered
to be fragments of the continuous flora; the time of the continuity is
uncertain.

ACKNOWLEDGMENTS

Support for bryological field work, the results of which have been
incorporated in this review, has been by the National Research Council
of Canada. I am grateful to C. Leo Hitchcock who generously permitted
me to use the unpublished manuscript of Volume I of Vascular Plants
of the Pacific Northwest.

Department of Botany, University of British Columbia, Vancouver

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206 MADRONO [Vol. 20

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“Se

ONO

VOLUME 20, NUMBER 4 OCTOBER 1969

Contents
A NEw SPECIES OF CASTILLEJA FROM THE SOUTHERN SIERRA
NevapA, Lawrence R. Heckard and Rimo Bacigalupi 209

Two NEw SPECIES OF UMBELLIFERAE FROM THE SOUTHWESTERN
UniTEp States, Mildred E. Mathais, Lincoln Constance,

and William L. Theobald 214
AGROSTIS PERENNANS (POACEAE), A REMOTE DISJUNCTION IN
THE Pacrric NortHwWEST, Curi G. Carlbom 220

A Mutant OF LITHOCARPUS DENSIFLORUS, John M. Tucker
Wiliam E. Sundahl, and Dale O. Hail 221

CHROMOSOME NUMBERS IN SOME NortH AMERICAN SPECIES OF
THE GENUS CirstuM. II. WESTERN UNITED STATES,

Gerald B. Ownbey and Yu-tseng Hsi 225
A NEw CopROPHILOUS SPECIES OF CALONEMA (MYXOMYCETES),

Donald T. Kowalski 229
A NEw CAMPANULA FROM NORTHERN CALIFORNIA,

Lawrence R. Heckard 231
NoTEs AND News: Maprono 213

PROPOSED CALIFORNIA TREE ATLAS, James R. Griffin; THE

CLASSIFICATION SOCIETY 236

Reviews: T. G. Tutin, V. H. Heywood, N. A. Burges, D. M.
Moore, D. H. Valentine, S. M. Walters, and D. A. Webb,
Flora Europaea, Vol. 2 (Wallace R. Ernst); V. H. Hey-
wood, Plant Taxonomy (Dennis R. Parnell) ; C. Leo Hitch-
cock, Arthur Cronquist, Marion Ownbey, and J. W.
Thompson, Vascular Plants of the Pacific Northwest,
Part 1 (Robert Grnduff) ; Philip A. Munz, Supplement to
A California Flora (Peter H. Raven) 237

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KENTON L. CHAMBERS, Oregon State University, Corvallis
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WALLACE R. ERNST, Smithsonian Institution, Washington, D. C.
ArTuRO GoMEZ PomPA, Universidad Nacional Autonoma de México
EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia
MIpreD E. MATHIAS, University of California, Los Angeles
RoBERT OrNvDvuFF, University of California, Berkeley
Marion OwnBeEy, Washington State University, Pullman
Duncan M. Porter, Missouri Botanical Garden, St. Louis
REED C. Rot.ins, Harvard University, Cambridge, Massachusetts
IrA L. WiccINs, Stanford University, Stanford, California

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Cholame, California. Recording Secretary: John West, Department of Botany, Uni-
versity of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Depart-
ment of Botany, University of California, Berkeley. Treasurer: June McCaskill, De-
partment of Botany, University of California, Davis.

A NEW SPECIES OF CASTILLEJA
FROM THE SOUTHERN SIERRA NEVADA

LAWRENCE R. HECKARD and RIMo BACIGALUPI

Castilleja praeterita, Heckard and Bacigalupi, sp. nov. (fig. 1, a-h).
Planta perennis omnino praesertim insuper septatim villoso-hirsuta, caul-
ibus costato-angulatis 1-3.5 dm altis, insuper adscendente ramosis pler-
umque pluribus ex radice lignosa arcuato-adscendentibus. Folia 3—5 cm
longa, linearia vel lineari-lanceolata, obtusa vel acuminata, superiora in
lobis duobus (interdum 4) angustissimis adscendentibus dissecta. In-
florescentia angusta, conferta, villosa glanduloso-puberulentaque, 1.5—2
cm lata, demum 8-14 cm longa, bracteis eius plerumque 1.5—2 cm longis,
5-8 mm latis, parte centrali oblonga, ad basem versus paulo angustata,
ad apicem truncato-rotundatum versus sensim latescente, lobis laterali-
bus plerumque duobus angustatis adscendentibusque apice rotundatis ex
medio instructa. Calyx 14-18 mm longus, plus minusve quadrangularis,
sagittaliter aequaliterque 7-11 mm fissus, lobis terminalibus duobus in
quoque latere 0.5—2 mm longis, acutis vel rotundatis, quam latis paulo
longioribus. Corolla 13-16 mm longa, calyci plus minusve aequilonga
vel paulo exserta, galea ca. 1.5 mm longa, dorso minute puberulenta, labio
inferiore ca. 1.5 mm longo, sacculis tribus angustis conniventibus, apice
brevissime in lobulis involutis apiculatisque ca. 0.5 mm longis terminanti-
bus instructo. Inflorescentiae quod ad colorem attinet bimodales: brac-
teae calycesque aut pallide virides, ad apices versus citrini, galeae dorso
viridis membranis lateralibus stramineis; aut ei glaucescentes, ad apices
versus lateriti vel plerumque ei omnino pallide phoenicei, galeae mem-
branis lateralibus violaceo-porphyreis.

Perennial with one to several (to 15) arcuate-ascending annual costate-
angulate stems arising from a woody root-crown. Stems 1—3.5 dm tall,
occasionally with subordinate ascending branches arising mostly above
their mid-points. Stems and leaves villous-hirsute with septate tri-
chomes throughout, occasionally with some inconspicuous glandular tri-
chomes, becoming increasingly villous just below and in the inflores-
cence. Leaves 3-5 cm long and 2—5 mm broad, linear to linear-lanceo-
late or narrowly oblong, blunt or attenuate to a pointed tip, the lower
entire and nearly petioleless, the upper sessile, broadened below a pair
(rarely 2 pairs) of much narrower, divergent or ascending lobes. In-
florescence villous and glandular-puberulent, dense and narrow, 1.5—2
cm broad, becoming 8-14 cm long, the bracts and calyces scaberulous
towards the tips. Bracts shorter and broader than the upper leaves,
15-20 (to 25) mm long, 5-8 mm broad, with a pair (rarely 2 pairs) of
narrow lobes with rounded or apiculate apices ascending from near the
middle, the much broader central lobe oblong but slightly narrowed be-

Maprono, Vol. 20, No. 4, pp. 209-240. March 25, 1970.

209

[Vol. 20

Fic. 1, Castilleja praeterita: a, habit, x 1%; b, inflorescence, K ™%; leaf, x 2;
d, detail of bract margin, X 10; e, bract and flower, * 2; f, corolla, side view, xX 2;
g, corolla, front view, X 2; h, chromosomes, n = 12, prophase II, * 800 ( Baciga-
lupi & Heckard 9213). Drawings a-g are from the type collection, Bacigalupi &
Heckard 9190.

low and with a broadened truncate or truncately rounded distal portion.
Pedicels 1-2 mm long. Calyx 14-18 mm long, somewhat quadrangulate,
the corners formed by four major veins, cleft medianly 7-11 mm, lat-
erally 0.5—2 mm into lobes which are a little longer than broad and either
pointed or rounded and sometimes apiculate. Corolla 13—16 mm long, in-
cluded or exserted 1-2 mm beyond the calyx, its tube glabrous or sparse-
ly pilose (the trichomes sometimes gland-tipped) along the ventral and
lateral surfaces, its galea ca. 5 mm long, minutely puberulent along the
back, its lower lip ca. 1.5 mm long, formed of 3 narrow pouches, the
outer folds of which are connivent and each of which terminates in an
inconspicuous incurving hooded and apiculate lobe. Inflorescence of two
color-forms: (1) bracts and calyces pale green and tipped with lemon
yellow, the corolla with the membranous lateral flaps of the galea yel-
lowish white; (2) bracts and calyces grayish green and tipped with pale

1969 | HECKARD & BACIGALUPI: CASTILLEJA 211

brick red, or often pale reddish with anthocyanin pigment throughout,
the corollas with the lateral flaps of the galea maroon-purple; the back
of the galea and lower lip green in both forms but the lower lip of the
second color-form often suffused with dull purple. Anther-pairs sparsely
pilose along the dehisced margins, connivent at anthesis and forming a
tier in the opening between the lateral margins of the galea, later sep-
arating, the dimorphic anther-cells of the upper anther-pair ca. 1.5—2
mm and 1.0-1.5 mm long respectively, the lower anther-pair with the
corresponding cells ca. 0.3 mm shorter. Stigma capitate but very shal-
lowly bilobed, a little over 0.5 mm broad, protruding 0-2 mm beyond
the galea at anthesis. Ovary cylindric-ovoid, 2—2.5 mm long, maturing
into a many-seeded capsule ca. 8-10 mm long. Seeds 1—1.5 mm long and
0.75 mm broad, variously angled, with a tan, reticulate outer testa.
Chromosome number: n = 12.

Type. Bacigalupi & Heckard 9190, northern edge of Horse Meadow
on Salmon Creek, about 7 air miles east southeast of Fairview, southern
Sierra Nevada, Tulare Co., California, elevation 7400 ft., Aug. 10, 1966,
n = 12 (JEPS-holotype; isotypes to be distributed).

Other representative specimens (UC or JEPS unless otherwise indi-
cated): Tulare Co.: Big (Brown) Meadow, ca. 3 miles southeast of
Horse Meadow, Bacigalupi & Heckard 9206 (chromosome voucher: n
— 12); Monache Meadows, Munz 15067 (RSA); Bakeoven Meadows,
Howell 26764 (CAS, DS); southwest of Templeton Meadow, C. NV.
Smith 1515; Tunnel to Ramshaw Meadow, Howell 25909 (CAS, UC);
Little Whitney Meadow, Ferris & Lorraine 10779; Cottonwood Pass,
Peirson, Aug. 8, 1911; north of Crabtree Meadows, Raven 7594; below
Timberline Lake, Robbins 3623. Inyo Co.: Horseshoe Meadow, Paciga-
lupi & Heckard 9212 (chromosome voucher: n = 12); above Cot-
tonwood Sawmill, canyon of Cottonwood Creek, Bacigalupi &» Heckard
9213 (chromosome voucher: n = 12); Cottonwood Lakes, Alexander &
Kellogg 3315.

The specific name of this species comes from the Latin word, praeter-
ita, meaning overlooked, neglected, or passed by without notice.

Usually associated with and probably parasitic on the roots of Arte-
misia rothrocku and, less frequently, with Artemisia tridentata (Baci-
galupi 9213); dryish, sandy or rocky slopes, commonly bordering mea-
dows; in areas of Pinus contorta ssp. murrayvana, elevation 7,400—11,000
it

It will be seen that the cited specimens fall into two groups—a south-
ern one which includes the type of the species, and a more northerly
one—between which there is a gap of approximately 24 air miles.

_ Whether this interval represents an actual gap in distribution or merely

reflects minimal collecting in this area is conjectural.
There are some morphological differences between the northern and

_ southern populations of C. praeterita. The two southern populations

(Big, formerly Brown, Meadow and Horse Meadow) consistently have

212 MADRONO [Vol. 20

the yellow inflorescence with yellow-tipped bracts and calyces rather
than the pale brick red inflorescence which predominates in northern
populations. There is also a tendency in these southern populations for
the plants to be larger in all respects, including bracts and floral parts.
Especially noticeable is the broader inflorescence. Chromosome counts
indicate that these differences are not related to level of polyploidy as
samples both from southern populations and from two of the northern
populations all proved to be diploid.

Although collections of this species have been accumulating in her-
baria over the past seventy years, they have either remained unidentified
or have been incorrectly referred to Castilleja pilosa (Wats.) Rydb. The
superficial resemblance to C. pilosa may be attributed to similarities in
habit, foliage, and indument. Castilleja pilosa is sharply differentiated
from C. praeterita, however, by its calyx, which is more or less equally
4-cleft rather than relatively deeply slit only in its sagittal plane, as in
most castillejas. The 4-cleft calyx has been used by some taxonomists
(Watson, 1871; Gray, 1878; Jepson, 1925) as a basis for referring C.
pilosa and closely related species to Orthocar pus.

Judging from the absence of annotations on collections of C. praeter-
ita in California herbaria (CAS, DS, JEPS, RSA, UC), it would seem
that Pennell never saw specimens of our new species. Had he done so,
he doubtless would have devised a scheme of subgeneric subdivisions
very different from the one he proposed in 1951. Castilleja praeterita has
a lower lip somewhat pouched but less than half the length of the galea,
a combination of characters which precludes its inclusion in any of the
19 sections proposed by Pennell (1951) or as modified by Ownbey
(1959). The nearest affinity would seem to be with sections Chrysanthae
and Pallescentes, but the teeth of the lower corolla-lip are far less de-
veloped than in most members of those groups, being reduced to mere
apiculations. The nature of the apex of the calyx-lobes is used by Pen-
nell to separate section Chrysanthae (lobes obtuse) from section Palles-
centes (lobes acute). Shape of apex of the calyx-lobes of C. praeterita
varies considerably and hence is of doubtful value in relating the species
to either of these two sections. The calyx-lobes of C. praeterita are never
linear-triangular (as in most species of section Pallescentes), but are
broader and conform more closely with section Chrysanthae. On the
other hand, C. praeterita occupies ecological sites quite different from
the moist meadows preferred by all species of section Chrysanthae: it
favors drier and better drained slopes surrounding the lower portions of
mountain meadows, a habitat more similar to that occupied by species of
section Pallescentes. In view of these points of non-conformity, one
might be tempted to propose a new section to accommodate C. praeteriia,
but we believe there is need for a complete re-evaluation of the sections
as proposed by Pennell (1951). Noel Holmgren (1968) is in agreement
with this opinion and has not given nomenclatural recognition to super-
specific groupings. He suggests that the reticulate nature of species re-

1969 | HECKARD & BACIGALUPI: CASTILLEJA 213

lationships has resulted from speciation through hybridization between
members of relatively unrelated groups. A program of artificial hybridi-
zations currently in progress (Heckard, 1964) indicates that hybridiza-
tion is indeed possible not only between quite unrelated species, but even
between members of differing polyploid levels.

Our conclusion is that there are no species particularly closely related
to Castilleja praeterita. The very short apiculate teeth of the lower co-
rolla-lip occur in no species of either section Chrysanthae or Pallescentes.
The only species which approaches this reduced condition of teeth is C.
culbertsoni (section Chrvsanthae), where the teeth, though still longer
than those of C. praeterita, are reduced to about 0.5 mm in length. De-
spite this floral similarity, the sum of characters does not indicate a
close relationship between the two species. The only other species of
Castilleja which, to our knowledge, has the teeth of the lower lip as
reduced as in C. praeterita is C. cinerea Gray. This restricted endemic
of arid, sagebrush slopes at elevations above 6,000 ft. in the San Ber-
nardino Mts. has a deeply 4-cleft calyx which places it far from C.
praeterita and among the Orthocar pus-like species.

Jepson Herbarium, Botany Department, University of California, Berkeley

LITERATURE CITED

Gray, A. 1878. Synoptical flora of North America. Vol. 2, Part 1, New York.

Heckarp, L. R. 1964. [Abstract.] Causes of taxonomic complexity in Castilleja.
Amer. J. Bot. 51:686.

HormcreNn, N. H. 1968. A taxonomic revision of the Castilleja viscidula group.
Ph.D. Thesis, Columbia University, New York.

Jepson, W. L. 1925. A manual of the flowering plants of California. Berkeley.

Ownsey, M. 1959. Castilleja. Jn C. L. Hitchcock, et al., Vascular plants of the
Pacific Northwest. Vol. 4. Univ. Washington Press, Seattle.

PENNELL, F. W. 1951. Castilleja. Jn L. R. Abrams, Illustrated flora of the Pacific
States. Vol. 3. Stanford Univ. Press.

Watson, S. 1871. Vol. 5, Botany. Jn C. King, Report on the geological exploration
of the Fortieth Parallel. Washington, D.C.

NOTES AND NEWS

MADRONO. Starting with the January 1970 issue (Vol. 20, No. 5), Madrono
will be increased to 48 pages per issue from the current 32. An increasing number
of submitted papers and the recent demise of two California journals, Leaflets of
Western Botany and the Contributions from the Dudley Herbarium, makes this
desirable. Unfortunately, it will be necessary to increase the subscription rate to
Madrono and the dues to the Society as follows: institutional subscriptions, $12.00
per year and individual membership, $8.00 per year. Student membership remains
at $4.00 per year.

TWO NEW SPECIES OF UMBELLIFERAE FROM THE
SOUTHWESTERN UNITED STATES

Mivprep E. Matuias, LINCOLN CONSTANCE, and WILLIAM L. THEOBALD

For a number of years the senior authors have recognized the exis-
tence of anomalous populations of Umbelliferae in western Texas and
adjacent New Mexico. Herbarium specimens of these puzzling plants,
usually in flower or immature fruit, have been referred to Pseudocymop-
terus montanus, a polymorphic ‘‘catch-all.” A restudy of the Umbelli-
ferae for the treatment of the family for the Manual of the Texas Flora
has necessitated a review of these discordant elements. Adequate fruit-
ing material is now filed in herbaria and it is apparent that recognition
of two species is warranted, one referrable to Aletes and the other to
Pseudocvmo pterus.

Aletes filifolius Mathias, Constance, and Theobald, sp. nov. Fig. 1.
Plantae acaules vel caulescentes; foliis ternato-pinnatisectis, divisionibus
filiformibus, 0.5—5.6 cm longis; pedunculis 7-38 cm longis, foliis longi-
oribus; involucri bracteis plerumque nullis; radiis 4-21, 6-20 mm longis,
involucelli bracteolis linearis vel lanceolatis; pedicellis 6-15, 1.5—5 mm
longis; calycis dentes evidentibus, lanceolatis; umbelluli fructis 2-10,
oblongis vel ovoideo-oblongis, 2.4—8 mm longis, 1.8-4 mm latis, costis
prominentis et alatis, alis suberosis; vittae plerumque in valleculis soli-
tariae, in commissuri 2.

Plants 2—4 dm tall, caespitose from a branching woody root crowned
with old leaf sheaths, acaulescent or with 1—2 stem leaves; leaves petio-
late, broadly ovate in general outline; blades 2.5—20 cm long, 2.5—14
cm broad, ternately-pinnately decompound, the ultimate divisions fili-
form, 0.5—5.6 cm long, 1-2 mm broad; petioles 2.5-15 cm long; pe-
duncles 7-38 cm long, longer than the leaves, scaberulent at the base of
the umbel; bracts of involucre usually wanting; rays 4-21, 6-20 mm
long, spreading-ascending; bractlets of involucel linear to lanceolate, 2—5
mm long, free to slightly connate at base, rarely reduced to one; pedicels
6-15, 1.5-5 mm long; calyx-teeth evident, lanceolate; petals yellow,
ovate with a narrower inflexed apex; styles slender, spreading, stylo-
podium lacking; disk present; fruits in each umbellet 2—10, oblong to
ovoid-oblong, 2.4-8 mm long, 1.8-4 mm broad, the dorsal and lateral
ribs prominent and corky-winged, wings pale yellow to white, rarely
inconspicuous; vittae large, mostly solitary in the intervals, 2 on the
commissure; seed subterete in transection, at times slightly channeled
under the vittae; the face more or less plane; sclerenchymatous cells
inconspicuous.

Type: Moist soil on ledge along stream, north fork, North Mckit-
trick Canyon, Guadalupe Mts., Culberson Co., Texas, 18 Aug. 1946,
Correll 13961 (US 2178785—holotype, LL-isotype).

214

1969 | MATHIAS, CONSTANCE, & THEOBALD: UMBELLIFERAE 215

Distribution: Mountains of western Texas and southern and south-
central New Mexico.

Representative Specimens. TEXAS: Brewster Co.: frequent on lime-
stone north slopes of Altuda Mt., 10 mi SE of Alpine, Del Norte Mts.,
4500 ft., 8 June 1948, Warnock 7833 (LL, MICH, SMU, UC); abun-
dant near top of Baldy Peak, Glass Mts., 13 July 1940, Warnock W297
(UC). Culberson Co.: moist shaded bluffs, upper McKittrick Canyon,
Guadalupe Mts., 2140 m, 22 July 1931, Moore & Stevermark 3573
(GH, MICH, MO, UC, US); scarce among boulders, bed of Ist narrow
canyon off west side of North McKittrick Canyon, Guadalupe Mts.,
1575 m, 16 July 1945, McVaugh 7413 (LL, MICH, UC); vicinity of
Frijole Post Office, 5000-9500 ft, 4 Aug. 1930, Grassl 166 (MICH);
Pine Springs Canyon, Guadalupe Mts., 7 Sept. 1961, Correll & John-
ston 24272 (LL); numerous and scattered on the banks and in the bed
of the creek in Pine Spring Canyon and north McKittrick Canyon, 6800
ft, 2 June 1949, Hinckley & Hinckley 11 (US); McKittrick Can., 28
June 1939, Lehman (GH); growing in sand about rock, narrow canyon
floor, North Fork of McKittrick Canyon, Guadalupe Mts., 25 July 1957,
Correll & Johnstone 18496 (LL). Jeff Davis Co.: dry rocky places,
Little Aguja Canyon, Davis Mts., 1765 m, 17 June 1931, Moore & Stev-
ermark 3136 (GH, MO, UC).

NEW MEXICO: Dona Ana Co.: deep east-west canyon above Drip-
ping Springs, 6300-7300 ft, 28 July 1952, Dunn 8436 (UC); Filmore
Canyon, Organ Mountains, 26 May 1905, Wooton (UC, US); Van Pat-
ten’s, Organ Mts., 9 June 1906, Standley (MO, US), 16 July 1902,
Wooton (US), 29 Aug. 1894, Wooton (US). Eddy Co.: Carlsbad, 4
June 1924, Lee 154 (US). Grant Co.: Big Hatchet Mts., 17 May 1892,
Mearns 39 (US); Sycamore Creek, 13 Aug. 1902, Wooton (US). So-
corro Co.: lower valley of Tulerosa River, 30 Aug. 1905, Hough (US).

The genus Aletes was revised recently by Theobald, Tseng, and Ma-
thias (1964) who recognized five species occurring in the southwestern
United States. Aletes filifolius is readily distinguishable from the other
members of the genus on the basis of habit and basic leaf pattern. It is
the only caulescent species, usually with one or two stem leaves, and a
ternately-pinnately decompound leaf blade. The other five members of
the genus are acaulescent and have pinnate or bipinnate leaves. Never-
theless, on the basis of other vegetative characteristics, and especially in
floral and fruit characters, all of these taxa form a distinct and coherent
genus.

Aletes acaulis, the most widespread taxon and type species of the
genus, also occurs in western Texas. Both it and A. filifolius are known
from similar habitats on canyon slopes, canyon bottoms, and in rocky
crevices in the mountains and in several instances have been reported
from the same vicinity. As noted above they are readily distinguishable
from each other by their foliage characters.

[Vol. 20

MADRONO

216

1969 | MATHIAS, CONSTANCE, & THEOBALD: UMBELLIFERAE 217

It is expected that 4. filifolius will be recorded from adjacent Mexico.
Several collections from that area are possibly referable to it but con-
firmation must await the collection of more mature fruiting material.

Pseudocymopterus longiradiatus Mathias, Constance, and Theo-
bald, sp. nov. Fig. 2. Plantae caulescentes, foliis caulis 1-3, ternato-
pinnatisectis, divisionibus ovatis vel oblongis, lobatis, lobis obovatis vel
lineari-oblongis, 2-14 mm longis; pedunculis 13-49 cm longis, foliis
longioribus; radii 8-18, patentibus, 17-58 mm longis, pedicellis 12—25,
2-6 mm longis; umbelluli fructis 2-12, ovoideo-oblongis vel oblongis,
6-9 mm longis, costis lateralibus late alatis, alis membranaceis vel spon-
giosis; vittae in valleculis solitariae, in commissuris 2.

Plants 3-9 dm tall from a long taproot, caulescent with 1-3 stem
leaves, rarely acaulescent; leaves petiolate, ovate-oblong to broadly
ovate in general outline, 4.5-15 cm long, 3.5—13 cm broad, ternate-pin-
nately decompound; the ultimate divisions ovate to oblong in outline,
pinnately lobed to parted, the lobes obovate to linear-oblong, 2-14 mm
long, 1.5—4 mm broad, acute to distinctly acuminate; petioles 4.5-15 cm
long; peduncles terminal or axillary, 13-49 cm long, longer than the
leaves, scaberulent to hirtellous-pubescent at the base of the umbel;
bracts of involucre usually wanting; rays 8-18, 17-58 mm long, spread-
ing; bractlets of involucel linear-lanceolate, 3-11 mm long, free to slight-
ly connate at base, longer or shorter than the flowers; pedicels 12-25,
2—6 mm long; calyx teeth evident, ovate to deltoid, often with a pale or
colored margin; petals pale cream-yellow, ovate with a narrower in-
flexed apex; styles slender, spreading, stylopodium lacking; disk pres-
ent; fruits in each umbellet 2-12, ovoid-oblong to oblong, 6—9 mm long,
3—5 mm broad, flattened dorsally or appearing terete due to wings,
wings membranous or spongy, linear to triangular in transection; vittae
large, solitary in the intervals, 2 on the commissure; the seed face more
or less plane; sclerenchymatous cells absent.

Type: In sandy soil under maples and oaks by dry stream bank, Up-
_ per McKittrick Canyon, 6000 ft, Guadalupe Mts., Culberson Co., Texas,
22 June 1947, Meyer & Meyer 2186 (UC 758246-holotype; MO-
Isotype).

Distribution: Mountains of western Texas and southern New Mexico.
Representative specimens. TEXAS: Brewster Co.: infrequent on
_ northeast limestone slopes of Mt. Ord, 15 mi S of Alpine, Gage Estate,
4650 ft, 23 May 1949, Warnock & Turner 8645 (LL). Culberson Co.:
_ wooded bluff, locally abundant, Ist narrow canyon off west side of
_ North McKittrick Canyon, Guadalupe Mts., 1575 m, 16 July 1945,
_ McVaugh 7414 (LL, UC); moist soil in cool canyon, Guadalupe Moun-
tains, steep canyon on southeast slope of Pine Top Mountain, 15 Aug.

Fic. 1. Aletes filifolius: a, habit, x ™%; b, flower at anthesis, x 10; c, mature
fruit, lateral view, X 5; d, mature fruit, transection, x 8.

218 MADRONO

[Vol. 20

Fic. 2. Pseudocymopterus longiradiatus: a, habit, X
dorsal view, X 3; c, mature fruit, transection, X 8.

¥%; b, mature’ fruit,

1969 | MATHIAS, CONSTANCE, & THEOBALD: UMBELLIFERAE 219

1946, Correll 13904 (LL); under maples, south fork of McKittrick Can-
yon, Guadalupe Mountains, 30 April 1962, Correll & Ogden 25055
(LL); under oaks, South Fork of McKittrick Canyon, Guadalupe Mts.,
2 July 1958, Correll & Johnstone 19161 (LL); crevices of cliffs, north-
facing canyon, about 9 miles north of Van Horn, 24 April 1961, Correll
& Rollins 23805 (LL); on flats, vicinity of Frijole Post Office, 8000 ft,
10 Aug. 1930, Grassl 134 (MICH); woods near spring, canyon, east
side of Guadalupe Peak, 7500-8000 ft, 29 May-4 June 1912, Chase
5980 (MICH). Jeff Davis Co.: dry grassy shaded canyon slope, Little
Aguja Canyon, Davis Mountains, 1550 m, 17 June 1931, Moore & Stey-
ermark 3131 (GH, MICH, MO, UC); abundant on slopes in shaded
bottom, branch canyon to east just above pass, Wild Rose Pass, about
15 mi NE of Ft. Davis, 10 April 1947, McVaugh 7891 (MICH).

NEW MEXICO: Otero Co.: west of Mt. Park in an old apple or-
chard surrounded by pinyons and rather dense oak thicket about 12 ft.
tall, along the road to Cloudcroft, ca 6800 ft, 28 June 1952, Dunn 8097
(LA); Alamo (Lincoln) National Forest, Haynes Canyon, in forest of
Douglas spruce and rock pine, 10 Aug. 1911, Barlow (MO).

Pseudocymo pterus has been a difficult genus to delimit. For a num-
ber of years it has been treated as monotypic with the single highly
variable species, P. montanus. Studies now in progress indicate that sev-
eral taxa may warrant recognition within this complex. Pseudocymop-
terus longiradiatus can be distinguished by the much longer rays, the
larger fruit with larger vittae, and the ternate-pinnately decompound
leaves. Pseudocymopterus montanus in western Texas is reported only
from Mt. Livermore at elevations from 6000 to 8000 feet while P.
longiradiatus occurs at generally lower elevations in both the Davis and
Guadalupe Mountains.

Department of Botanical Sciences, University of California, Los Angeles
Department of Botany, University of California, Berkeley
Department of Biology, Occidental College, Los Angeles

LITERATURE CITED
THEOBALD, W. L., C. C. TsenNc, and M. E. Maruias. 1964. A revision of Aletes
and Neoparrya (Umbelliferae). Brittonia 16:296—-315.

AGROSTIS PERENNANS (POACEAE), A REMOTE
DISJUNCTION IN THE PACIFIC NORTHWEST

Curt G. CARLBOM

Agrostis perennans (Walt.) Tuckerm. is an extremely polymorphic
taxon in the eastern United States, with many different ecotypic and
morphological forms having been described. While it has been reported
from all states east of the Mississippi River and three states immedi-
ately to the west (Hitchcock, 1950), I am reporting it here for the first
time from the Pacific Northwest.

A collection of an Agrostis (Carlbom Ac-73, OSC) from a mesic, semi-
suaded site in the dense coastal Pseudotsuga—Tsuga forest approxi-
mately two miles east of Copalis, Washington, and an early Thomas
Howell collection from Tillamook, Oregon (Howell, s.n., 1884, ORE),
identical to it, were compared with a range of specimens from the east-
ern United States, including ten holotypes of names considered synonyms
with A. perennans by Hitchcock. These two western collections were
identical with the eastern material in the usual spikelet characters as
well as in the trichodium-net development.

Five S, plants (OSC) and cloned material of Ac-73 were morphologi-
cally indistinguishable from one another when grown under uniform
garden conditions. Cytological studies of meiosis in pollen mother cells
of Ac-73 and two S, plants revealed no meiotic irregularities, with nor-
mal bivalent pairing (21 IT) in all cells investigated. The haploid chro-
mosome number, n — 21, agrees with root-tip counts (2n = 42) re-
ported by Bjorkman (1960) for collections of A. perennans from east-
ern Canada.

Agrostis perennans is possibly a relict species in the Pacific Northwest,
having had previously a more or less continuous east-west distribution
as late as the Pleistocene.

Department of Botany, Oregon State University, Corvallis
Present address: Institute of Genetics, University of Lund, Sweden

LITERATURE CITED
ByORKMAN, S. O. 1960. Studies in Agrotis and related genera. Symb. Bot. Upsal. 17:
1-112.
Hircucock, A. S. 1950. Manual of the grasses of the United States. Ed. 2, revised
by Agnes Chase, U.S. Dept. Agr. Misc. Publ. 200.

220

A MUTANT OF LITHOCARFUS DENSIFLORUS

Joun M. Tucker, WILLIAM E. SUNDAHL, and DALE O. HALL

A peculiar, low, shrubby, oak-like plant was discovered by Dale Hall
in January, 1962, on the Challenge Experimental Forest (which is main-
tained by the Pacific Southwest Forest and Range Experiment Station,
Berkeley) approximately one mile north of Challenge, Yuba Co., Califor-
nia. It was quite unlike any species known to the senior author and did
not appear to be a hybrid of any California oak.

About 25 more were found a few weeks later. They weve about 1 to 1%
feet tall and were located in a strip 100 feet long and 50 feet wide. Many
of these new seedlings had germinated in squirrels’ seed caches. There
was about one ‘‘odd oak” in each group of five to 15 seedlings, the others
being tanoak. About half of the ‘odd oak” seedlings in these caches even-
tually died.

Some of the remaining plants were transplanted to pots. One was
brought to Davis; however, it soon died. Another, in a container with its
native soil, was brought to Davis in February, 1964. This one survived,
although its growth has been extremely slow. Altogether, there are about
20 of these plants living that we know of. They have been found in sev-
eral different places, about 2 miles apart (fig. 1). All were in close prox-
imity to Lithocarpus densiflorus. A brief note, and a photograph of one
of them, appeared in the Sacramento Bee (p. 10, County Life Section),
July 21, 1963 (by Melvin Gagnon, at that time a staff writer for the Bee).

Still another occurrence came to our attention recently (after this man-
uscript had been submitted for publication). Two individuals—one re-
portedly 8 feet tall (Smith No. 91, DAV), which is larger than any we
have seen—were noted at 2150 feet elevation, center of Sec. 10, T. 18 N.,
R. 7 E., in Yuba Co., by B. F. Smith, Sept. 20, 1968.

Close inspection in the field with regard to possible parent trees, and
re-examination of fine morphological details, led us to suspect that these
peculiar little plants must be odd forms of Lithocarpus densiflorus. Al-
though radically different from typical arborescent forms of this species
in size, stature, and leaf form (fig. 2), they are very similar in certain
other characters such as leaf pubescence and form of the stipules.

The several sites where these plants have been discovered so far lie be-
tween 2,150 and 3,425 feet elevation in the ponderosa pine belt (Transi-
tion Zone). Here the dominant vegetation is a forest of Pinus ponderosa,
Pinus lambertiana, Pseudotsuga menziesii, Abies concolor, Libocedrus
decurrens, Arbutus menziesii, Quercus kelloggii, Lithocarpus densiflorus,
Acer macrophyllum, Cornus nuttallii, and occasionally other woody
species.

At first, it was considered that these plants might be monosomics or tri-
somics. Squash preparations were made on several occasions in a search
for visible chromosomal aberrations. All such attempts were unsuccessful.

220

bo
Bo
Bo

MADRONO [Vol. 20

Challenge Experimental Forest

Yuba County, California
T.19N., R.7E.,MDM

LEGEND

location of mutant
section number
compartment number
paved road

dirt road

Fic. 1. Location of tanoak mutants.

A more likely hypothesis was advanced by Robert M. Echols, Genet-
icist, Pacific Southwest Forest and Range Experiment Station. He sug-
gested that we are dealing with a sublethal recessive mutation. If this is
true, the rather frequent occurrence of these odd plants over an area
more than two miles in extent indicates that this mutation spread, and at
present, occurs with a fairly high frequency in the heterozygous condi-
tion. A tree heterozygous for the mutation would appear to be a normal
tanoak, yet could be expected to produce an occasional homozygous re-
cessive acorn due to pollination by some other heterozygous tree nearby,
or possibly by selfing. With this thought in mind, acorns were collected
from several normal-appearing trees in the vicinity of the mutants, and
germinated at Davis during the fall of 1965.

From a total of 45 acorns that germinated (which represents the prog-
eny of five different trees), it was apparent that one seedling was a mutant

1969] TUCKER, SUNDAHL, & HALL: LITHOCARPUS 228

Fic. 2. A large shrubby mutant of Lithocarpus densiflorus.

(fig. 3). This was in the progeny of tree number 0-5. There can be no
doubt, therefore, that these peculiar little plants are forms of Lithocarpus
densiflorus, and, provided the hypothesis is correct, it is thus established
that tree number 0-5 is heterozygous.

224 MADRONO [Vol. 20

Fic. 3. Progeny from Lithocarpus densiflorus parent tree 0O-5-—a normal-appearing
seedling (left), and the mutant (right).

A program of experimental self-pollination may well be attempted. If a
sizeable number of experimentally-produced acorns can thus be obtained,
we would be in an excellent position to test our hypothesis that the aber-
rant forms are the result of a single gene mutation. If, on germination,
approximately 14 of them proved to be mutant forms and 34 normal, the
hypothesis would be confirmed. A strikingly different ratio would require
some other explanation.

If the tree proves to be completely self-incompatible, an attempt could
be made to locate a second heterozygous tanoak by progeny tests similar
to those already carried out. A series of controlled crosses, using the two
trees as parents, would then be attempted. In this way we may make a
small contribution to the knowledge available for the genus Lithocar pus.

Since this distinctive mutant is so different from typical L. densiflorus,
a formal taxonomic designation is justified. Although this case does not
seem to be adequately accommodated by the present International Code
of Botanical Nomenclature, it seems logical to us to accord it the status
of forma.

Lithocarpus densiflorus {. attenuato-dentatus Tucker, Sundahl, and
Hall, f. nov. A Lithocarpo densifloro tipico foliis lineari-oblongis apicibus
anguste, acuminatis, ad basem versus cuneatis, marginibus attenuato-
dentatis, dentibus exilibus discedit.

Small woody plants, the largest to ca. 8 feet in heigit; although
commonly with a single stem or trunk from the base, tending to become
branched, spreading, and shrub-like; twigs slender and persistently gray

1969 } OWNBEY & HSI: CIRSIUM 225

hirsute-tomentose; buds hirsute; leaves to 13 cm long and 1.5 cm wide,
linear-oblong, apex narrowly acuminate, sub-aristate, base narrowly cu-
neate, margins minutely and irregularly revolute, attenuate-dentate, the
teeth narrow, sub-aristate, and directed forward; upper surface glabrous,
dark green and slightly glossy, lower surface pale green, dull, and sparse-
ly stellate-pubescent; secondary veins 6-9 on a side; petiole 10-15 mm
long, sparsely stellate to glabrate, the petiole and midrib (especially on
the under side of the leaf) yellowish; stipules caducous, linear, 7-8 mm
long, lightly hirsute. No flowers or fruit have been observed.

Holotype: California, Yuba Co., Challenge Experimental Forest, ca. 1
mile N of Challenge, elevation 2675 ft., J. M. Tucker s.n., May 22, 1963
(DAV).

Department of Botany, University of California, Davis
Pacific Southwest Forest and Range Experiment Station, Challenge, California
Intermountain Forest and Range Experiment Station, Boise, Idaho

CHROMOSOME NUMBERS IN SOME NORTH AMERICAN
SPECIES OF THE GENUS CIRSIUM. II. WESTERN
UNITED STATES

GERALD B. OWNBEY and YU-TSENG HSI

As was suggested in the previous paper of this series (Ownbey and
Hsi, 1963), chromosome numbers in the North American species of
Cirsium may contribute substantially to an understanding of the taxon-
omy and evolution of the genus. At the very least they will be of sig-
nificant value in the initial arrangement of the species into alliances
which will with certain reservations represent natural groups. Our ex-
perience to date indicates that morphologically similar species now
grouped together frequently have the same or only slightly varying
chromosome numbers.

Due to the small and intergrading sizes of the chromosomes of Cir-
sium, i.e., 0.6—3.0 microns in length when fully contracted in the species
discussed here, it has not been possible adequately to characterize the
karyotypes of the species examined. It is safe to assume that at least one
pair of satellite chromosomes can be observed in all species and fre-
quently one or two additional satellite chromosomes are seen. Karyo-
type evolution in Cirsium may lead either to a loss or gain in numbers,
but reduction in numbers seems to be the rule. It is usually assumed that
17 is the primitive number in the genome as the preponderance of living
species have retained this number. Accessory chromosomes, when pres-
ent, cannot be identified morphologically in our preparations and for this
reason it has been concluded that they are intact or nearly so. A few ex-
amples of chromosomal fragments have also been seen.

226 MADRONO [Vol. 20

All of the chromosomal data recorded here is based upon the study of
root tip preparations. With few exceptions the root tips were obtained
from 7-14 day old seedlings grown under sterile conditions in the lab-
oratory.

Counts for Cirsium acanthodontum, C. rydbergu, C. utahense, C. wal-
lowense, C. fastoris and C. arizonicum are published here for the first
time. Earlier counts for the remaining species have been published as
follows: C. brevistylum, Moore & Frankton (1962); C. scopulorum,
Moore & Frankton (1965); C. tweedvi, Moore & Frankton (1965); C.
coloradense, Ownbey & Hsi (1963, under C. foliosum), Moore & Frank-
ton (1967); C. scariosum, Ownbey & Hsi (1963, under C. foliosum),
Moore & Frankton (1967); C. tioganum, Ownbey & Hsi (1963, under
C. foliosum), Moore & Frankton (1967); C. ochrocentrum, Hsi (1960),
Ownbey & Hsi (1963); C. undulatum, Hsi (1960), Frankton & Moore
(1961), Ownbey & Hsi (1963); C. subniveum, Ownbey & Hsi (1963) ;
C. californicum, Moore & Frankton (1963); C. occidentale, Moore &
Frankton (1963). The new counts for these species agree closely with
the earlier ones except that greater variation in the number of accessory
chromosomes is sometimes reported here.

We have arranged the species in the text in the sequence and in the
groups proposed by Petrak (1917). Species not known to Petrak, viz.,
C. acanthodontum Blake, C. brevistylum Cronq., C. subniveum Rydb.,
C. wallowense Peck and C. pastoris Howell, are placed in the groups to
which they appear to be most closely allied. All American thistles, both
native and introduced, belong to the subgenus Eucirsium.

Sect. Echenais, Subsect. Americana
Cirsium acanthodontum Blake. 2n = 32 (2 plants). Oregon, Curry Co., 11.7 miles
N of Agness, Ownbey & Ownbey 3054, MIN. This collection came from near
the type locality.

Sect. Onotrophe, Subsect. Crassifolia
Cirsium rydbergii Petrak. 2n = 34 (1 plant). Utah, Grand Co., Salt Wash, about
one-half mile N of Turnbow Cabin, Arches National Mcnument, Welsh &
Moore 2742, MIN.

Subsect. Minutiflora
Cirsium brevistylum Cronq. 2n = 34 (3 plants). Oregon, Coos Co., 15.5 miles N of
Agness, Ownbey & Ownbey 3055, MIN.

Subsect. Globosa
Cirsium scopulorum (Greene) Cock. 2n = 34, 35, 36, 37. Colorado, Clear Creek
Co., Mt. Evans, alt. ca. 11,000 ft., Ownbey 3671, MIN, 2n = 34 (4 plants),
2n = 35, (1 plant), 2n = 36 (2 plants), 2n = 37 (4 plants).
Cirsium tweedyi (Rydb.) Petrak. 2n = 34 (1 plant). Wyoming, Yellowstone Na-
tional Park, Sylvan Pass, Ownbey & Ownbey 3071, MIN.

Subsect. Acaulia
Cirsium coloradense (Rydb.) Cock. 2n = 34, 36? Colorado, La Plata Co., 21.9
miles E of Durango, Route 160, alt. ca. 7200 ft., Ownbey & Hsi 2642, MIN,
2n = 34 (2 plants), 2n = 36? (2 plants).

1969 | OWNBEY & HSI: CIRSIUM tA |

Cirsium scariosum Nutt. 2n = 34, 36. Idaho, Clark Co., 2.4 miles N of Spencer,
Ownbey & Ownbey 3067, MIN, 2n = 36 (1 plant); Montana, Powell Co., 5
miles SW of Avon, Ownbey & Hsi 2908, MIN, 2n = 34 (3 plants), 2n = 36?
(1 plant) ; Wyoming, Johnson Co., 13.5 miles W of Buffalo, Route 16, Ownbey
& Ownbey 3030, MIN, 2n = 34 (1 plant).

Cirsium tioganum (Congd.) Petrak. 2n = 34 (1 plant). Colorado, Jackson Co., 1.3
miles N of Walden, Ownbey & Ownbey 1497, MIN.

Cirsium canescens Nutt. X C. tioganum (Congd.) Petrak. 2n = 34 (2 plants).
Colorado, Jackson Co., 1.3 miles N of Walden, Ownbey G Ownbey 1497a, MIN.
The data for this hybrid were earlier published by Ownbey & Hsi (1963) under
C. canescens Nutt. & C. foliosum (Hook.) DC.

Subsect. Acanthophylla

Cirstum ochrocentrum Gray. 2n = 32, 34. Arizona, Apache Co., 3 miles N of
Concho, Baker & Baker 2512, MIN, 2n = 32 (2 plants), 2n = 34 (4 plants) ;
Texas, Tom Green Co., 4.4 miles NE of Tankersly Ownbey & Baker 2994, MIN,
2n = 32 (7 plants). The Arizona collection represents the southwestern race
cf the species distinguished by its scarlet-red ccrollas and scarcely decurrent
leaves. The Texas collection also appears to represent a recognizable race having
unusually small heads, phyllaries and phyllary spines.

Cirsium undulatum (Nutt.) Spreng. 2n = 26 (4 plants). Texas, Terrell Co., Inde-
pendence Creek bottoms near Pecos River, Demaree 48442, MIN. This collec-
ticn ccmes from the southern periphrey of the range of C. undulatum. It differs
morpholcgically from typical material in having extensively branched stems and
small heads. The phyllaries and anthers are also smaller than usual for the
species.

Subsect. Campylophylla

Cirsium subniveum Rydb. 2n = 34, 35, 36. Idaho, Bonneville Co., 0.5 mile W of
the Snake River bridge, just W of Swan Valley, Ownbey & Ownbey 3043, MIN,
2n = 34 (2 plants), 2n = 36 (1 plant); Lincoln Co., 19 miles S of Carey,
Rceute 26, Ownbey & Ownbey 3046, MIN, 2n = 34 (3 plants) ; same locality,
Ownbey & Ownbey 3047, MIN, 2n = 35 (2 plants). The voucher fcr Ownbev
& Ownbey 3047 was aberrant in having glabrous and shining upper surface cf
the leaves and in the breadly auriculate bases of the principal cauline leaves. The
other voucher specimens cited compare closely with the type specimen of C.
subniveum (Nelson 1070, US).

Cirstum utahense Petrak. 2n = 30, 32. Arizona, Coconino Co., along Route 180,
about 30 miles NW of Flagstaff, Baker & Baker 2511, MIN, 2n = 32 (5
plants) ; Yavapai Co., between Cordis Junction and Mayer, Deaver 5962, MIN,
2n = 30 (2 plants). The distinctions between C. wtahense and C neomexi-
canum are not always clear but, following the treatment of Cirszum by Howell
(1960) for the Pacific States, the voucher specimens are identified as C. utah-
ense. The voucher of Baker & Baker 2511 is not typical C. utahense in being
more thinly pubescent throughout and in having basal and cauline leaves re-
motely pinnatifid, the segments lanceolate.

Cirstum wallowense Peck. 2n = 34 (4 plants). Oregon, Wallowa Co., 31 miles N of
Enterprise, Ownbey & Ownbey 3060, MIN.

Cirstum californicum Gray. 2n = 28, 29, 30. California, Mariposa Co., Route 41,
Yosemite National Park, 11.8 miles N of the junction of the road to Mariposa
Grove, Baker & Baker 2503, MIN, 2n = 28 (4 plants), 2n = 29 (1 plant),
2n = 30 (2 plants).

Cirsium occidentale (Nutt.) Jepson. 2n = 28, 29, 30. California, Marin Co., along
Mt. Tamalpais road, 0.2 mile beyond Mt. Tamalpais State Park, Baker & Baker
2493, MIN. 2n = 28 (2 plants), 2n = 29 (1 plant), 2n = 30 (5 plants).

228 MADRONO [Vol. 20

Cirsium pastoris Howell. 2n = 30, 31, 32, 33. California, Mendocino Co., 1.6 miles
N of Cummings, Baker & Baker 2491, MIN, 2n = 30 (1 plant), 2n = 32 (2
plants), 2n = 33 (1 plant) ; Oregon, Josephine Co., 10 miles N of Grants Pass,
Route 99, Ownbey & Ownbey 3058, MIN, 2n = 30 (1 plant, 2n = 31 (1
plant), 2n = 32 (5 plants), 2n = 33 (1 plant).

Sect. Erythrolaena, Subsect. Subcoriacea
Cirsium arizonicum (Gray) Petrak. 2n = 30 (5 plants). Arizona, Coconino Co., a
few miles NE of Strawberry on the Long Valley road, Baker & Baker 2509,
MIN.

This study was supported by National Science Foundation grants
G9071 and GB2727 made to the senior author. We wish to thank the
following people for supplying us with collections of Cirsium seeds and
vouchers used in these studies: Gary R. Baker, Chester F. Deaver and
S. L. Welsh. We are indebted to Mrs. Siu-tsun Hsi for many excellent
cytological preparations.

Botany Department, University of Minnesota, Minneapolis
Biology Department, Southwest State College, Marshall, Minnesota

LITERATURE CITED

FRANKTON, C., and R. J. Moore. 1961. Cytotaxonomy, phylogeny, and Canadian
distribution of Cirsium undulatum and Cirsium flodmanii. Canad. J. Bot. 39:
21233;

Howe Lt, J. T. 1960. Cirsium. Jn Abrams, L. and R. S. Ferris, IHlustrated flora of
the Pacific States, Vol. IV. Stanford Univ. Press.

Hs, Y. 1960. Taxonomy, distribution and relationships of the species of Cirsium
belonging to the series Undulata. Ph. D. Thesis, University of Minnesota, Minne-
apolis.

Moorg, R. J., and C. FRANKTON. 1962. Cytotaxonomy and Canadian distribution of
Cirsium edule and Cirsium brevistylum. Canad. J. Bot. 40:1187-1196.

—_———., and —————-. 1963. Cytotaxonomic notes on some Cirsium species of the
western United States. Canad. J. Bot. 41:1553-1567.

—., and —————.. 1965. Cytotaxonomy of Cirsium hookerianum and related
species. Canad. J. Bot. 43:597-613.

.. and ——_——.. 1967. Cytotaxonomy of foliose thistles (Cirsium spp. aff. C.
foliosum) of western North America. Canad. J. Bot. 45:1733-1749.

Ownsey, G. B., and Y. Hsi. 1963. Chromosome numbers in some North American
species of the genus Cirsium. Rhodora 65:339-354.

Petrak, F. 1917. Die nordamerikanischen Arten der Gattung Cirsium. Beih. Bot.
Centralbl. 35:223-567.

A NEW COPROPHILOUS SPECIES OF CALONEMA
(MYXOMYCETES)

DONALD T. KOWALSKI

At present, the genus Calonema is monotypic, the single species be-
ing C. aureum Morgan. It is very similar to the genus Oligonema. The
only difference between the two genera is that in Oligonema the capilli-
tium is composed of short, free elaters, while in Calonema it consists of
long threads more or less united into a net. Some authors, like Lister
(1925) and Hagelstein (1944), believed that C. aureum was nothing
more than a form of Oligonema flavidum Peck. They both retained
Calonema in their monographs, however, but only for the sake of
convenience.

The species to be described below is common on cow dung through-
out the Sacramento Valley, wherever natural forage is present. It fruits
abundantly in cavities embedded in the dung or on the lower surface in
contact with the soil. Since both of these niches are characterized by a
high relative humidity, almost all of the collections consist of perfectly
matured sporangia.

Calonema luteolum Kowalski, sp. nov. Sporangiis dissipatis, gre-
gariis vel agglomeratis, sessilibus, globosis vel subglobosis, 0.1-O0.5 mm
diam; peridio simplici, membranaceo, luteo, iridescenti; capillitio luteo,
filamentis ramosis et anastomosis formandis reticulum, laevigatus, tubu-
laris, 2.0 crassis; sporis globosis, spinulosis, luteis, 12-13 diam;
plamodio ignoto.

Type. Near intersection of Butte Creek and U.S. Highway 99E, 2
miles south of Chico, Butte Co., California, April 22, 1967, D. T. Ko-
walski 5998 (1A-holotype, MICH, TEX.).

Collections examined: Kowalski 2554, 5362, 5393, 5400, 5407, 5420,
5508, 5514, 5521, 5526, 5529, 5536, 5540, 5544, 5549, 5552, 5613, 5614,
5620, 5623, 5625, 5636, 5639, 5992, 5993, 5994, 5995, 5996, 5997, 5998.

This species is known from the Sacramento Valley of California, where
it is found only on cow dung.

Sporangia (fig. 1) scattered, clustered, to often heaped, sessile, irregu-
larly shaped, globose or subglobose, occasionally slightly elongated, 0.1-
0.5 mm in diameter; peridium single, thin, membranous, transparent,
spores clearly visible through the peridium, iridescent, smooth, entirely
lacking any distinctive markings, brilliant yellow; hypothallus lacking;
capillitium (fig. 2) composed of branching and anastomosing tubular
threads, forming a distinct net, threads of uniform thickness, averaging
about 2.0 » thick, weakly attached to the peridium over the entire sur-
face, surface of threads smooth or minutely ornamented, but completely
lacking any sign of spiral ornamentation, internal thickenings present,
dividing up the threads into numerous chambers 1-4 , in diameter,
yellow, few free ends, but when free ends present, not noticeably in-

229

230 MADRONO [Vol. 20

Fics. 1-3. Calonema luteolum: 1, sporangia, * 33; 2, capillitium, 670; 3,
spores, X 1440.

flated; spores: (fig. 3.) globose, spinulose; vellow in mass, bright yellow
by transmitted light, 12-13 » in diameter; plasmodium unknown.

This species is easy to identify in the field. The restricted nature of
the substrate and the fact that the sporangia are bright yellow and form
in small heaped clusters is distinctive. The major microscopic charac-
teristics are the smooth capillitium, forming a distinct reticulum, and
the spinulose spores,:12—I3. microns in diameter. It can easily be sep-
arated from C. aureum on these features. Calonema aureum also has a
reticulate capillitium, but its surface bears rings or fragmentary spirals.
In none of the 30 collections listed above, did I observe any rings or
spirals on the capillitium of C..dJuteolum. The major difference, how-
ever, is that C. aureum-has spores which are 13—15 microns in diameter
and they bear a distinct, ‘coarse reticulum.

Whether or not ©. luteolum belongs in the genus Calonema is de-
batable. While it has characteristics resembling Calonema, in other ways
it is similar to the genus Perichaena: It resembles Perichaena in regards
to the-capillitium and-spores., The capillitium of Perichaena is also re-
ticulate and lacks rings or spirals and the spores of this genus are never

1969 | HECKARD: CAMPANULA 251

reticulate, being either minutely warted or spinulose. Species of Pevi-
chaena, however, have a two-layered peridium and the sporangia, while
they may be clustered, are never heaped. Calonema luteolum is similar
to the genera Calonema and Oligonema in that in both of these genera
the peridium is single, membranous and often iridescent and the spor-
angia can be heaped. It differs from these genera, however, in that their
capillitium often has spiral markings and the spores are reticulate, while
C. luteolum has a smooth capillitium and spinulose spores.

The problem arises in deciding which characteristics are the most im-
portant taxonomically, or, which characteristics are the most important
in showing phylogenetic relationships, I believe the presence of a single
peridium and heaped sporangia indicate that the affinities of C. luteolum
are with Calonema and Oligonema even though the capillitial and spore
characteristics are reminiscent of the genus Perichaena. Perhaps this is
one area in which cultural studies can be of immense importance in de-
termining evolutionary relationships.

This study was supported by Lng National Science pouncetien (Grant
GB-5799).

Department of Biology, Chico State College, Chico, California

LITERATURE CITED

HacELstTeEIn, R. 1944. The Mycetozoa of North America. Mineola, N.Y.
ListErR, A. 1925. A monograph of the Mycetozoa. Ed. 3. Revised by G. Lister.
Brit. Mus. Nat. Hist., London.

A NEW CAMPANULA FROM NORTHERN CALIFORNIA

LAWRENCE R. HECKARD

An undescribed Campanula has turned up in area where one would
not have expected to find a new species of flowering plants—Castle
Crags, the spectacular and conspicuous mass of spires and domes which
rises 4,000 ft. above the Sacramento River southwest of Dunsmuir. A
trail in Castle Crags State Park leads up to and among the granitic pin-
nacles where the Campanula grows fairly abundantly in the crevices
of sloping and even vertical walls. The plant was first collected in 1948
by the late Freed Hoffman whose private herbarium was given to the
University of California. The specimen, identified as C. scabrella, came
to my attention in connection with a review of the genus Campanula in
California. I am pleased to name this plant for Stanwyn G. Shetler of
the Smithsonian Institution, Washington, D.C., student of Campanula
and author of a useful conspectus of the genus in North America.

Campanula shetleri Heckard, sp. nov. Fig. 1. Herba perennis rosu-
lata dense caespitosa tota scabro-hispidula; folia breve spathulata pari-
bus dentium oppositorum duobus instructa; caules floriferi 2—5 cm alti,

bo
Ww
is)

MADRONO [Vol. 20

Fic. 1. Campanula shetleri: a, habit, & 114; b, rosette leaf, * 3; c, detail of
leaf lobe, & 10; d, detail of peduncle, « 20; e, flower, x 5; f, flower at fruiting
stage showing the pore of dehiscence, X 5; g, seed, X15; h, meiotic chromosomes
(camera lucida of MII, Heckard 1524), n= 17, ca. X 1300. (Voucher specimens
for the drawings: a-e, Heckard 1524; f,g, Heckard 1731).

1969] HECKARD: CAMPANULA RX:

inflorescentiis 1—5-floribus terminantes; corolla infundibuliformis 9—10.5
mm longa, ore similiter lata lobis eius ovato-deltoideis 4.5—5 mm longis,
quam tubo leviter brevioribus; hypanthium cupulatum super medium
latissimum lobis eius integris subulatis; capsulae pori in partem eius
super medium collocati.

Mat-forming perennial with densely clustered rosulate shoots arising
from long and slender, sparingly branched underground stems with ad-
ventitious roots arising singly or in clusters in axillary regions where the
inflorescence-branches have abscissed; rosettes with loose to dense clus-
ters of spirally arranged leaves, the stem clothed below for varying dis-
tances with withered leaves or leaf-bases, some rosettes giving rise in the
leaf-axils to 1 to 3 flowering shoots; rosette-leaves scabrous-hispidulous
throughout, the lamina short-spatulate (roughly hexagonal), about 6—7
mm long and 4—5 mm broad, with 2 pairs of opposite teeth, the apex
cuneate with a sharp to blunt tip, the base also cuneate, tapering gradu-
ally into a short petiole 1-2 mm long; flowering shoots ascending to
erect, 2-5 cm high, bearing few to ca 15 leaves which are gradually re-
duced towards the shoot-apex, the herbage scabrous-hispidulous through-
out, the cauline trichomes slightly retrorse; inflorescence with one ter-
minal flower or also with 1—2 (4) later-opening flowers each borne on
a short pedicel 1-4 mm long, the pedicels axillary in the uppermost
leaves (bracts) and bearing usually 2 sub-opposite or 1 subulate brac-
teole(s) about 2 mm long; calyx-lobes subulate, entire, densely hispidu-
lous, 4-5 mm long in anthesis, lengthening in fruit to 5-6 mm; corolla
pale blue to nearly white, glabrous except for a few small pointed tri-
chomes (similar to those of the herbage) on the outer face at the tip of
the corolla-lobe, funnelform, 9-10.5 mm long and about as broad dis-
tally, the lobes ovate-deltoid and acute, spreading and becoming re-
curved, 4.5—5 mm long, slightly shorter than the tube; anthers glabrous,
narrow-oblong, 2.5—3 mm long; filaments 2—2.5 mm long, the basal por-
tion ovate-deltoid and ciliate, ca. 1.5 mm long, slightly exceeding the
linear distal portion; style about as long as the corolla, 7-8 mm long in
anthesis, papillose on the upper one-third, the 3 stigmata each 1 mm
long, becoming strongly recurved after anthesis; hypanthium cup-shaped
(2—2.5 mm in diam.), scabrous-hipidulous throughout, 2—3 mm long and
usually slightly narrower, broadest above the middle; capsule erect,
broadly to narrowly cylindric or somewhat urceolate with broadened
base, 3-4.5 mm long and 3-4 mm broad, often longer than broad,
rounded and irregularly 3-lobed at base, opening just above the middle
by three valves; seed just less than 1 mm long, ovoid to ellipsoid with
a slight crest, somewhat 3-angled with the 2 broader sides flattened, the
third rounded, the seed coat smooth, shining, amber in color; chromo-
some number: n = 17.

Holotype. CALIFORNIA. Shasta Co.: Castle Crags State Park, along
trail to Castle Dome, ca. 4200 ft; forming mats in cracks of north- and

234 MADRONO [Vol. 20

northeast-facing granite cliffs, Heckard & Bacigalupi 1524, 14 June
1966 (JEPS).

Other specimens examined: CALIFORNIA. Shasta Co.: trail to sum-
mit of Castle Crags, almost at timber line, Hoffman 2644 (UC); 0.2 mile
south of Castle Dome, 4700 ft., Heckard 1525, 1731 (JEPS). Siskiyou
Co.: north-facing cliffs at Little Castle Lake, ca. 6000 ft., Roderick, 1
Sept. 1967 (JEPS).

The plants grow in granitic detritus and humus accumulated in the
crevices and cracks of steep and even vertical north- and northeast-
facing cliffs. Associated crevice-plants are Penstemon newberryi ssp.
berrvi, Heuchera merriamu, and Ivesia gordonii, while the most com-
mon and conspicuous trees and shrubs of the surrounding area are Pinus
ponderosa, Quercus chrysole pis, Lithocarpus densiflora var. echinoides,
and Arctostaphylos patula. A few individuals of Picea breweriana, a spe-
cies restricted to the mountains of northern California and southern Ore-
gon, are present. Although C. shetleri grows near and above the appar-
ent timberline of Castle Crags, it should be considered a part of the
Yellow-Pine Forest community of Munz and Keck (Munz and Keck,
1949; Munz, 1959) since timberline in the crags is probably largely con-
trolled by the granitic substrate and steepness of the topography.

The new species has a known range of less than 4 miles, but it should
be looked for on other unexplored granitic peaks in the general region.
The Emited distribution of C. sketleri is not unique in this genus. Shetler
(1963) points out that several North American campanulas are highly
localized.

Shetler (1963) divides the 16 Campanula species restricted to North
America into 5 species-groups based on habit and habitat. Campanula
shetleri has its closest affinity with group 2, the arctic-alpine endemics,
consisting of 5 species from western North America. Strictly speaking,
the relatively low elevation (4,000—6,000 ft.) occupied by C. shetleru
would not qualify it as an endemic of the arctic-alpine zone. The addi-
tion of C. shetleri to the group thus makes the group-name less appro-
priate. The species most similar to C. shetleri is another narrow endemic,
C. piperi of the Olympic Mountains in Washington, nearly 500 air-miles
distant. Campanula piperi occupies a rock-crevice habitat similar to
that of C. shketleri but in the arctic-alpine zone. In general, C. shetlert is
much smaller in almost all respects than C. piperi, a comparison which
includes all flower-parts such as anthers, stigmata, and hypanthium. Sev-
eral additional well-defined morphological differences between the two
species are listed in Table 1. Both species have the same chromosome
number of n == 17,

Two other species in the arctic-alpine group, C. wilkinsiana and C.
scabrella, are of some interest because, although rare in California, they
occur within a short distance of C. shetleri on neighboring mountains.

Campanula wilkinsiana is a rather narrow endemic of the Salmon-

1969 | HECKARD: CAMPANULA 23

TABLE 1. A COMPARISON OF THE PRINCIPAL MORPHOLOGICAL DIFFERENCES
BETWEEN CAMPANULA SHETLERI AND C., PIPERI

C. shetleri C. piperi

Height To 5 cm To 10 cm
Indument Scabrous-hispidulous through- Finely scabrous-hirtellous in
out, the trichomes up to 0.2 upper portions (calyx-lobes,
mm long. hypanthium, upper stem and

cauline leaves) ; trichomes less
than 0.1 mm long. Rosette
leaves glabrous.

Leaves Short-spatulate, to 1 cm long, Oblanceolate-spatulate, 1.5—3
dentate with 2 pairs of teeth. cm long, serrate-dentate with
3-6 pairs of teeth.

Calyx-lobes Narrowly triangular (subu- Triangular, 5-10 mm long, oc-
late), 4-6 mm long, entire. casionally with 1 or more

sharp teeth.

Corolla Pale blue to whitish, about Blue (rarely white), 12-16
i0 mm long, Icbes the same mm long, lobes almost twice
length as or slightly shorter as long as tube.

than tube.

Trinity Mts. and Mt. Shasta, the latter only 15 air-miles from Castle
Crags. Ecologically, it differs from C. shetleri in that it grows in spring
or streamside sites. Morphologically, C. wilkinsiana differs in being a
much larger, completely glabrous plant lacking a conspicuous rosette
and characterized by obovate to elliptic leaves which are serrate in their
upper halves. Moreover, it has deep blue corollas up to 15 mm long
terminating stems which are leafless in the upper portion.

The other neighbor, C. scabrella, a plant of western Montana to
Washington and southward, grows abundantly on Mt. Eddy just 10
air-miles north of Castle Crags. This species, which forms small mats on
talus slopes at elevations of 8—9,000 ft., is amply distinct from C. shet-
leri in its entire oblanceolate leaves, herbage with grayish puberulence,
and a capsule which opens near the summit.

Jepson Herbarium, University of California, Berkeley

LITERATURE CITED
Muvnz, P. A. 1959. A Califernia flora. Univ. Calif. Press, Berkeley.
. and D. D. Keck. 1949. California plant communities. Aliso 2:87-105.
SHETLER, S. G. 1963. A checklist and key to the species of Campanula native or
commonly naturalized in North America. Rhodora 65:319-337.

236 MADRONO [Vol. 20

NOTES AND NEWS

PROPOSED CALIFORNIA TREE ATLAS. For several years I have been com-
piling detailed distributional maps in collaboration with W. B. Critchfield. Eighty-
six tree species native to California, and adjacent fringes of Nevada, are included.
This U.S. Forest Service project is largely based on Vegetation Type Map Survey,
Soil Vegetation Survey, and Forest Survey sources. Much of the data has been
available for decades but never in a convenient form. For some species the avail-
able Forest Service data yield relatively complete maps. For others the coverage is
marginal, and we need help.

In 1970 working copies of the new compilations should be available for all
species. I hope to place copies for review in the larger botanical centers in Cali-
fornia. Several sets will be circulated to interested individuals. This note is a plea to
those kind hearted souls who have unpublished tree distributicn items stored away.
Perhaps some of you can contact me before publication ot the final atlas. Unpub-
licized tree records from any part of the state, even for the more obvious species,
would help. But data about specific range limits or isolated colonies in Lassen,
Modoc, Napa, Siskiyou, Trinity counties and all of the Mojave region would be
particularly helpful—Jameres R. Grirrrn, Hastings Reservation, Carmel Valley,
California 93924.

THE CLASSIFICATION SOCIETY. The annual meeting of The Classification
Society, North American Branch, will be held April 8-9, 1970 at Battelle Memorial
Institute, Columbus, Ohio. For further details, write the Program Chairman, Doc-
tor Joseph Kruskal, Bell Telephone Laboratories, Murray Hill, New Jersey 07994.

The Classification Society, founded in Great Britain in 1964, has as its main
purpose the promotion of cooperation and interchange of views and information
among those interested in the principles and practice of pattern recognition and
classification in any discipline that uses them. As a result, its membership includes
anthropologists, biologists, computer and information specialists, geologists, librari-
ans, linguists, psychologists, soil scientists and others.

The Society seeks to provide unique services to its members. These include sym-
posia on classification that are not discipline-constrained and a project under con-
sideration that will result in a bibliography of articles dealing with the theoretical
and applied aspects of classification. Supplements to the original bibliography would
be issued periodically.

Business of the Society is conducted by a Committee elected by the membership.
The Society recently organized into two branches, The European Branch and the
North American Branch. Other branches will be organized as the need arises. Cur-
rent membership numbers around 300, divided equally between the two branches.
Annual dues are U.S. $3.00 and entitle members to receive copies of the Bulletin
of the Society, which contains contributions of both a formal and informal nature.
Membership applications may be obtained from the Secretary, Doctor Theodore J.
Crovello, Department of Biology, The University of Notre Dame, Notre Dame,
Indiana 46556, or from Doctor J. Willmott, Department of Computation, Univer-
sity of York, Heslington, York, England.

1969 | REVIEWS Ry |

REVIEWS

Flora Europaea. Edited by T. G. Tutin, V. H. HeEywoop, N. A. Burcers, D. M.
Moore, D. H. VaLenTINE, S. M. Watters, and D. A. Wess. Vol. 2. Rosaceae to
Umbelliferae, xxvii + 455 pp., 5 maps. Cambridge University Press. 28 February
1969 [‘1968"']. $23.50.

The Editorial Organization of the Flora Europaea project produced Vol. 1,
Lyccpcdiaceae to Platanaceae, in 1964 after approximately eight years of existence
(reviewed, Madrono 18: 62-63. 1965). This second volume, about five years later,
continues the botanical synthesis in English for the multilingual region between the
Atlantic and the Ural Mountains. The project is expected to be completed in a
total of four volumes. Although this work is intended for use in Europe it is ap-
plicable, at least in part, because of the extensive distribution of many genera and
species, throughout the Northern Hemisphere and to whatever European plants are
cultivated cr naturalized. Appendices explain abbreviations of author’s names and
titles of references for Vol. 2. Each volume has its own fold out maps, English-
Latin vocabulary, and index and can be used independently.

Volume 2 treats 50 families including Rosaceae, Leguminosae, and Umbelliferae,
respectively with 35, 74, and 110 genera. The number of species keyed out for some
of the larger genera are 75 species for Rubus, 118 for Alchemilla, 133 for Astragalus,
99 fer Trifolium, 105 for Euphorbia, and 92 for Viola. Reconsideration ef some
generic points of view has returned Daszphora fruticosa to Potentilla, thus differing
from Flora USSR, and happily has reunited Cornus permitting respectful disregard
for Thelycrania and Chamaepericlymenum, thus differing from the Flora of the
British Isles. Inclusion of persistent introductions reveals, astonishingly, that Gun-
nera tinctoria is locally naturalized in western Europe. Eleven species of Eucalyptus
have been included although there is no mention that they reproduce.

The only deficiency worth commenting on concerns the failure to give generic
citations and to indicate the type species for the genera. This would amount to a
statement established from the original descripticn of the name of the species on
which the genus was founded. Admittedly this sometimes is very complicated but
authers having delt with a genus taxonomically surely are in a better position to
express an opinion on generic type species than normally is possible fcr the staff
members of the Index Nominum Genericorum project.

The clearly stated, indented keys with numbered couplets, the condensation of
vast numbers of synonyms, all clearly indexed, the annotations on variaticn and
problem situations appearing on every page, mark Flora Europaea as an important
taxonomic summation. Perhaps only one of many small steps ferward for Man-
kind of 1969, the continuation of this flora is a great step forward for botanists,
many of whem will turn to these vclumes for purposes other than to identify un-
known specimens of European plants—WaAatLtace R. Ernst, Smithsonian Institu-
tion, Washington, D.C.

Plant Taxonomy. By V. H. Heywoop. iv + 60 pp., illustrated. St. Martin’s
Press, New York. 1967.

This book is an outline of the current field of plant taxonomy and is intended
for beginning biology students. The author attempts not only to present the scien-
tific and theoretical basis for plant taxonomy, but also tries to give an overview
of mcedern research methods in this area of biology. Although the topics represent
a rather complete survey of plant taxonomy (e.g., populations and species, cyto-
taxonomy, biochemical systematics, numerical taxonomy, etc.), the discussion and

938 MADRONO [ Vol. 20

the accompanying examples of these topics are uneven in quality and often in-
complete. For example, the discussion of natural selection found on page 26 fails
to make any mention of the important contributions population genetics has made
in helping us to define and understand this most important biological phenome-
non. Also at times the author assumes of the beginning biology student a greater
biological background than is reasonable to expect and as a result the significance
of many conclusions that are drawn are cryptic to the student. In general, I feel
that the bock, because of its brevity, falls short of its expectations—Dennis R.
PaRNELL, California State College, Hayward.

Vascular Plants of the Pacific Northwest. By C. Leo Hircucocx, ARTHUR CRON-
Quist, Marton Ownsey, and J. W. Tyompson. Part 1, 914 pp., illustrated. Uni-
versity of Washington Press. 1969. $25.00.

Publication of this part of Vascular Plants of the Pacific Northwest marks com-
pletion of this outstanding contribution to floristic knowledge of North America.
Fourteen years have been required for the conclusion of the serial publication of
this flora, which was initiated in 1955. This span of time is remarkably short when
one considers its coverage and the fact that many less ambitious works have seri-
ously faltered or expired before consummation. This most recent part—actually
Part 1 of the series—includes vascular cryptogams, gymnosperms, and monocoty-
ledons. In addition, it contains an index of the plant families, genera, and common
names covered by parts 1-5 and the species covered in Part 1. Also included are
an un-illustrated glossary and an unexpected and somewhat immured vegetative
key to aquatic vascular plants (mostly at the family level) which precedes the
treatment of the monocots. There is a modest ‘Additions and Corrections” section
which lists recent records for the area (some of which are accompanied by illustra-
tions), nomenclatural changes, alterations of ranges, or other comments relvant to
portions already published. This section of the work is particularly interesting for
its documentation of weedy species that have been recorded recently in the Pacific
Northwest. The bulk of Part 1 is occupied by treatments of the Cyperaceae and
Gramineae, two families which have made my own taxonomic existence more com-
plex than I would like. However, the fine illustrations that accompany the de-
scriptions of each species in both families have convinced me that genera and
species do exist in both of them and, furthermore, that it is probably possible for
an informed amateur to determine the Northwestern members of these specialized
and enigmatic families with relative ease. The vegetative key to the grasses (and to
aquatic plants) will be of particular value to biologists concerned with range and
wildlife management.

Anyone traveling in Britain or Europe will probably see far more Pacific North-
western native plants in cultivation there than he will see grown in their natural
range. One feature of this flora which perhaps I have underemphasized in my re-
views of the earlier numbers of this series is the very valuable commentary on the
horticultural merits and demerits of the indigens of the region. These advisory
comments have been provided largely by C. L. Hitchcock, whose extensive experi-
ence with the cultivation of northwestern natives is well known to horticulturalists
in the Seattle area. Gardeners are counseled of the virtues of potential cultigens
such as Scoliopus, Camassia, and Allium spp.; warned against failure with some
of the attractive but difficult species of Erythronium, Fritillaria, and Calochortus;
cautioned against the aggressiveness of Maianthemum dilatatum; and admonished
in strong terms against picking Trillium or depleting the rapidly diminishing popu-
lations of Calypso and Cypripedium.

Experts in the families covered by Part 1 Anadis would find some cause for
criticism of “their” genera or families (e.g., why Libocedrus and not Calocedrus?),
but I encountered little with which I had serious disagreement. Although this part

1969 | REVIEWS 239

is the most expensive of the series, the cost per page is considerably less than that
of its predecessors! Having been weaned botanically in the Pacific Northwest and
in the institution which might be considered the home of this project I cannot
claim to be objective in my assessment of this flora. The superb standards of con-
ception and execution which have characterized this project since its inception have
persisted until its completion. The authors deserve our congratulations and warm
praise for providing such a durable and scholarly treatment of the vascular plants
of the Pacific Northwest.—RoBert Ornpurr, University of California, Berkeley.

Supplement to A California Flora. By Putte A. Munz. iv + 224 pp. University
of California Press, Berkeley and Los Angeles. 1968. 57.00.

The appearance of a 224-page supplement to a flora of California is an event
of interest to all botanists interested in the plants of North America. The size of
the supplement attests to the amount of work that has been done on the plants
of the state in the decade since the publication of the original work. On the other
hand, this very size likewise makes the use of the supplement inconvenient. One
wishes that a new edition of the Flora could have been prepared instead, but since
that was apparently not possible, the supplement is a welcome substitute.

Most of the material in the supplement has to do with changes proposed in
revisions and other monographic works that have appeared since 1959. Unfor-
tunately, as in the original Flora, bibliographical citations are abbreviated to the
point where they are of limited value. Thus the name of a worker may refer to a
publication, a personal communication, or even a specimen, and the status of the
date which sometimes follows the name is of comparably uncertain origin. There is
no printed bibliography, and the reader will often not be able to distinguish the
possibilities given above. On the other hand, the addition of a bibliography would
have made the Supplement even longer, and, for those with a thorough working
knowledge of the California flora, the brief references given here will be of some
use in indicating the sources of the statements given.

In addition, range extensions, new chromosome numbers, and other new infor-
mation is given for hundreds of species. The format is convenient and the informa-
tion presented is easily integrated with that in the flora, and the supplement itself
is nearly free of typographical errors. There is a useful index, and the sturdy, at-
tractive volume is well printed and bound.

In connection with the supplement, it is of interest to draw attention to two
articles that provided statistical analyses of the material in the original book:
Smith, Gladys L. and Anita M. Noldeke, “A statistical report on A California
Flora,” Leafl. West. Bot. 9: 117-123. 1960., and Noldeke, Anita M. and J. T.
Howell, “Endemism and A California Flora,” Leafl. West. Bot. 9: 124-127. 1960.
These papers reveal that 162 families, 1075 genera, 5675 species, 1586 additional
subspecies and varieties, and 443 taxa of indefinite status were reported in the
Flora, with the largest families being Compositae (822 species), Gramineae (449
species), and Leguminosae (372 species), and the largest genera being Carex (144
species), Astragalus (93 species), Phacelia (87 species), Lupinus (82 species), and
Eriogonum and Mimulus (77 species each).

Nearly 30 per cent of the native species were endemic to California, as com-
pared with about 40 per cent reported by W. L. Jepson in his (1925) Manual of
the Flowering Plants of California. The reduction appears to be due largely to the
successful abandonment of Jepson’s highly provincial view of the plants of Cali-
fornia, as well as to extensive and intensive exploration just beyond the borders of
the State, especially in Baja California and Oregon. Nevertheless, California still
has an extraordinarily high proportion of endemics for a continental area, and were
the proportion cf endemism computed for the entire “California floristic province,”
which excludes the desert areas of California but includes portions of the three
neighboring states, the proportion would be much higher.

240 MADRONO [Vol. 20

As the number of naturalized species represented in the flora of California creeps
inexorably upward, dozens of recently reported genera, including for example
Pteris, Cyrtomium, Viscum, Gunnera, Rhagadiolus, Boussingaultia, Halodule, and
A pera, are listed in this Supplement, as is the family Aponogetonaceae. It is becom-
ing increasingly obvious that if future works are to provide a baianced account of
the plants of the State, that we shall have to reexamine our standards for inclusion
or exclusion of weedy plants; old records of species that did not persist, for ex-
ample, should presumably not continue to be listed.

In the Supplement, the family Balsaminaceae is reported from California for
the first time, on the basis of a native and an introduced species; the same appears
to be true for Loganiaceae, as Buddleza utahensis Cov. was accidentally omitted
from the Flora itself. The Koeberliniaceae are likewise added to the flora of the
State on the basis of a recently published record of Koeberlinia spinosa Zucc. in the
Chocolate Mts. of Imperial County. Kobresia and Bensoniella (Bensonia) are re-
cently reported genera of native plants, and outstanding native species added to the
flora of the State during the past decade include Lycopodium inundatum L., Abies
amabilis (Dougl.) Forbes, Saxifraga caespitosa L., Rubus nivalis Dougl., and Juncus
marginatus Rostk. Thus as in recent decades, nearly all of the additions to the flora
of the State come from the geologically complex and floristically rich ranges of
northern California, and, to a lesser extent, from the Sierra Nevada. This strongly
suggests that the flora of central and southern California is relatively well known.

Floristic work among the plants of California, in part spurred by the appear-
ance of the Flora, has been extensive, with genera such as Streptanthus and Galium
continuing to receive a great deal of attention, and critical recently described species
being listed in such genera as Polystichum, Silene, Opuntia, Monardella, and Nema-
cladus. Recent generic segregates such as Calocedrus, Chrysolepis, and Munzotham-
nus are recognized in the Supplement. A few new combinations, new varieties, and
at least one new species (Layza ziegleri Munz) are presented in the work itself.

An entirely new treatment of the genus Eriogonum, based on notes by James L.
Reveal, is incorporated directly into the Supplement. This 40-page synopsis indi-
cates that 104 species are now known from the State, in cemparison with the 77
listed in the Flora. The new treatment is an excellent contribution which should
greatly aid students of the genus, and inccrporates much new information. It is
much more monographic in scope than most of the Flora, containing numerous
critical notes. As such, the work on Eriogonum stand in sharp contrast, fer example,
to the scattered notes on Arctostaphylos summarized here. In the latter group, a
variety of workers have continued to present new combinations and new taxa with-
out ever approaching the overall view of the group necessary to achieve taxonomic
synthesis. A useful taxonomic system for a complex group such as Arctostaphylos
will never be built up of such blocks, and indeed, the overall pattern of variation
tends to become more and mcre obscure as the new taxa are proliferated. It is
greatly to be hoped that some of the studies of this genus now under progress will
eventually provide a new synthesis, based on a sound understanding of the biology
of the plants, that will make possible an appreciation of this most critical and in-
teresting genus, whose history is inextricably bound up with that of the floristic
associations with which it occurs—PETER H. RAvEN, Department of Biological Sci-
ences, Stanford University.

A WEST AMERICAN JOURNAL OF BOTANY

The subscription price of Madrono is $6.00 per year ($4.00 for stu-
dents). If your institution does not now subscribe to Madrono, we would
be grateful if you would make the necessary request. Since there is no
extra charge for institutional subscriptions, an individual may hold mem-
bership in the California Botanical Society on the basis of his institu-
tion’s subscription. Address all orders to: Corresponding Secretary,
California Botanical Society, Department of Botany, University of Cali-
fornia, Berkeley, California 94720.

INFORMATION FOR CONTRIBUTORS

Shorter items, such as range extensions and other biological notes,
will be published in condensed form with a suitable title under the general
heading, “Notes and News.”

Institutional abbreviations in specimen citations should follow Lanjouw
and Stafleu’s list (Index Herbariorum, Pari 1. The Herbaria of the World.
Utrecht. Fifth Edition, 1964).

Abbreviations of botanical journals should follow those in Botanico-
Periodicum-Huntianum (Hunt Botanical Library, Carnegie-Mellon Uni-
versity, Pittsburgh, Pennsylvania, 1968).

Footnotes should be avoided whenever possible.

Membership in the California Botanical Society is normally considered
a requisite for publication in MApRONo.

STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION

(Act of Oct. 23, 1962; Section 4369, Title 39, United States Code)

Maprono, A West American Journal of Botany, is published quarterly at Berke-
ley, California.

The publisher is the California Botanical Society, Inc., Life Sciences Building, Uni-
versity of California, Berkeley, California 94720.

The editor is John H. Thomas, Division of hace Biology, Stanford Univer-
sity, Stanford, California 94305.

The owner is the California Botanical Society: Inc., Life Sciences Building, Uni-
versity of California, Berkeley, California 94720. There are no bond holders, mort-
gagees, or other security holders.

The average number of copies distributed of each issue during the preceding 12
months is 650; the numbers of copies of the single issue closest to the filing date is
650.

I certify that the statements made by me above are correct and complete.
Joun H. THomas, Editor

October 25, 1969

MADRONO

VOLUME 20, NUMBER 5 JANUARY 1970

Contents

A NEw SPECIES AND SOME NEw COMBINATIONS IN CALYLOPHUS
(ONAGRACEAE), Howard F. Towner and Peter H. Raven 241

Two NEw SPECIES AND SOME NOMENCLATURAL CHANGES IN
OENOTHERA SUBG. HARTMANNIA (ONAGRACEAE),

Peter H. Raven and Dennis R. Parnell 246
STUDIES ON PLANTS OF THE GALAPAGOS ISLANDS. I. NEW SPECIES

AND CoMBINATIONS, Ira L. Wiggins 250
A NEw CoMBINATION IN TRICHONEURA FROM THE GALAPAGOS

IsLanps, John R. Reeder and Charlotte G. Reeder 253
A New CoMBINATION IN CHAMAESYCE FROM THE GALAPAGOS

Istanps, Derek Burch 253
NEw COMBINATIONS IN THE CYPERACEAE OF THE GALAPAGOS

IsLanps, Tetsuo Koyama 253
A NEw VARIETY OF OPUNITA MEGASPERMA FROM THE GALAPAGOS

Istanps, J. Lundh 254
NEw CoMBINATIONS IN THE COMPOSITAE OF THE GALAPAGOS

IsLanps, Arthur Cronquist 255

New CoMBINATIONS AND TAXA IN THE CACTACEAE OF THE GALAPA-
cos Istanps, Edward F. Anderson and David L. Walkington 256

Notes ON GALAPAGOS EUPHORBIACEAE, Grady L. Webster 257
NOMENCLATURAL CHANGES AND NEW SUBSPECIES IN THE CENTRO-
SPERMAE OF THE GALAPAGOS IsLANDs, Uno Eliasson 264

A NEw SPECIES OF POLYGONUM (POLYGONACEAE), Jerrold Coolidge 266
CHROMOSOME NUMBERS AND A PROPOSAL FOR CLASSIFICATION IN

SISYRINCHIUM (IRIDACEAE), Theodore Mosquin 269
COMPARATIVE NATURAL History OF Two SYMPATRIC POPULATIONS

OF PHOLISTOMA (HyDROPHYLLACEAE), Karen B. Searcy 276
A PRELIMINARY REPORT OF THE MYXOMYCETES OF CRATER LAKE

NATIONAL Park, OreGon, Dwayne H. Curtis 278
A NEw VaRARIA FROM WESTERN NortTH AMERICA,

Robert L. Gilbertson 282
Reviews: Alfred M. Wiedemann, La Rea J. Dennis, and Frank H.

Smith, Plants of the Oregon Coastal Dunes (Robert Ornduff) ;
Frank W. Gould, Grass Systematics (Dennis Anderson) 287

A WEST AMERICAN JOURNAL OF BOTANY

PULISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

— 2 m& Tre
ant HSO Wig
wy S ; 4 bs

f
é

MADRONO
A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established
1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional
subscription price $12.00 per year. Published quarterly in January, April, July, and
October by the California Botanical Society, Inc., and issued from the office of
Madrono, Herbarium, Life Sciences Building, University of California, Berkeley,
California. Orders for subscriptions, changes in address, and undelivered copies
should be sent to the Corresponding Secretary, California Botanical Society, Depart-
ment of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS
CLASS OF:

1970—LyMaN BEnson, Pomona College, Claremont, California
Mitprep E. Martuias, University of California, Los Angeles

1971—Marion Ownsey, Washington State University, Pullman
Joun F. Davipson, University of Nebraska, Lincoln

1972—Ira L. Wiccrns, Stanford University, Stanford, California
REED C. Roxiiins, Harvard University, Cambridge, Massachusetts

1973—Wa LLAcE R. Ernst, Smithsonian Institution, Washington, D. C.
ROBERT OrNDUFF, University of California, Berkeley

1974—Kenton L. CHAMBERS, Oregon State University, Corvallis
EMLEN T. LitTEL, Simon Frazer University, Burnaby, British Columbia

1975—-ArTURO GOMEz PompPa, Universidad Nacional Autonoma de México
Duncan M. Porter, Missouri Botanical Garden, St. Louis

Editor — Jonn H. THoMAs
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCASKILL
P. O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Marion Cave, Department of Botany, University of California, Berk-
eley. First Vice-President: Arthur Nelson, Department of Ecology and Systematic
Biology, San Francisco State College: Second Vice-President, Charles T. Mason, Jr.,
Herbarium, College of Agriculture, University of Arizona, Tucson. Recording Sec-
retary: John West, Department of Botany, University of California, Berkeley.
Corresponding Secretary: John Strother, Department of Botany, University of
California, Berkeley. Treasurer: June McCaskill, Department of Botany, University
of California, Davis.

The Council of the California Botanical Society consists of the officers listed
above plus the immediate past President, Harry Thiers, Department of Ecology and
Systematic Biology, San Francisco State College; the Editor of Madrofo; and
three elected Council Members: Robert Ornduff, Department of Botany, University
of California, Berkeley; Malcolm Nobs, Carnegie Institution of Washington, Stan-
ford; and Elizabeth McClintock, Department of Botany, California Academy of
Sciences, San Francisco.

A NEW SPECIES AND SOME NEW COMBINATIONS IN
CALYLOPHUS (ONAGRACEAE)

Howarpb F. Towner and PETER H. RAVEN

The evolution and breeding systems of the genus Calvlophus have
been the subject of dissertation research by the senior author for the
past three years. Information from this work has led to a biosystematic
study and a taxonomic revision of the genus, which will be published
in the near future. The present paper is intended to validate our new
combinations and a new species in advance of their use in the forthcom-
ing Manual of the Vascular Plants of Texas by Donovan S. Correll and
Marshall C. Johnston. The research on Calylophus has been supported
by National Institutes of Health graduate research fellowship 2-FO1-
GM-32,708-02 to the senior author and by National Sience Foundation
research grant GB-7949X to the junior author.

Calvlophus encompasses two former subgenera of Oenothera which
were recognized by their peltate or discoid stigmas, and in our treat-
ment will consist of five species. The only monograph concerned with
those species has been that of Munz (1929). In it were recognized
four species of Oenothera subg. Salpingia and one of subg. “Calylophis.”
Recent unification of these taxa as the genus Calvlophus by Raven
(1964) created a small, cohesive group of species which showed more
affinity to such genera as Gaura and Clarkia than to other species for-
merly referred to Oenothera. This change was followed by Shinners in
his treatment of the Texas species of Calvlophus (1964). Shinners at-
tempted to draw new taxonomic lines between infraspecific groups, and
he reduced a number of taxa to synonomy or varietal status. Munz’s
contribution to the North American Flora (1965) retained the traditional
generic alignment of the Onagreae, with Calylophus included in Oen-
thera. Most of the forms in the subgenus Salpingia were accorded spe-
cific rank.

Our investigation has resulted in a view of the Calvlophus hartwe sii
group which agrees with Shinners’ treatment in regarding it as an as-
semblage of intergrading infraspecific taxa. Several minor differences
appear between the two discussions. First, we prefer to use the rank
subspecies as the only infraspecific taxon, for reasons presented by
Raven (1969). The entities within C. hartwegii intergrade, but retain
their identities over large geographical ranges. They are thus major
forms which are best dealt with by according them subspecific status.
Secondly, we recognize C. hartwegii fendleri, which constitutes a dis-
tinctive series of populations distributed over a definite eco -geograph-
ical range. The third difference, to be fully treated in the forthcoming
revision, consists of small changes in the boundaries between taxonomic

Maprono, Vel. 20, No. 5, pp. 241-288. September 9, 1970.

241

242 MADRONO [Vol. 20

entities. According to our study, there is no justification at present for
distinguishing between two major groupings within this complex, such
as O. greggu and O. hartwegi in the sense of Munz (1929). Intergrada-
tion occurs between nearly any two forms which come into contact. A
reticulate pattern of phenetic and crossing relationships is evident, so
the only logical approach is to place all of the forms in C. hartwegii.

The Calvlophus serrulatus group, like the C. hartwegi complex, has
never been well understood, either in a biological or taxonomic sense.
We have discovered that this assemblage includes one outcrossing self-
incompatible species, C. drummondianus, and two species which are
self-pollinating complex structural heterozygotes. The interpretation of
this group was particularly difficult in the past, since the three species
often exhibit parallel patterns of geographical variation. It is not sur-
prising, therefore, that this situation has led to widely differing taxo-
nomic treatments.

CALYLOPHUS Spach, Hist. Veg. Phan. 4: 349. 1835. Meriolix Raf.,
Amer. Monthly Mag. & Crit. Rev. 4: 192. 1819; nomen nudum. Raf.,
J. Phys. Chim. Hist. Nat. Arts 89: 259. 1819; nomen nudum. Walbp.,
Repert. 2: 79. 1843. Calylophus Spach, Hist. Veg. Phan. 4: 349. 1835.
Calyvlophis Spach, Nouv. Ann. Mus. Hist. Nat. IIT. 4: 337. 1835. Oeno-
thera subg. Calylophis (Spach) T. & G., Fl. North Amer. 1: 501. 1840.
Oenothera subg. Salpingia T. & G., Fl. North Amer. 1: 501. 1840. Type:
O. lavandulaefolia T. & G. = Calylophus hartwegii ssp. lavandulifolius
(T. & G.) Towner & Raven. Salpingia (T. & G.) Raimann, in Engler &
Prantl, Naturl. Pflanzenfam. III. 7: 217. 1893; non Mart. 1828. Gal-
pinsia Britt., Mem. Torrey Bot. Club 5: 236. 1894.

Perennial herbs, sometimes with slightly woody lower stems and
base. Flowers actinomorphic, borne in axils of upper leaves, opening
near sunset, in mid-afternoon, or near sunrise; hypanthial tube well-
developed; petals yellow, fading pink or orange in some species. Stigma
peltate to discoid or globose-peltate, sometimes shallowly 4-lobed. Gam-
etic chromosome numbers, n = 7, 14. Three of the five species are self-
incompatible; two are self-compatible.

Type. Calvlophus nuttallai Spach. C. serrulatus (Nutt.) Raven. Five
species will be recognized in this paper and in the forthcoming revision.
They occur throughout the Great Plains from southern Canada to Texas,
and from the Great Basin and Southwest to north-central Mexico. The
genus is vaguely divisable into groups corresponding to Oenothera subg.
Calylophus and Salpingia, but the characters used to distinguish them,
namely sepal midrib height and hypanthial tube length, vary within each
complex. This separation may be used with caution in keys, however no
sectional division has been made because of the variability and the small
size of the genus.

1970] TOWNER & RAVEN: CALYLOPHUS 243

CALYLOPHUS HARTWEGII (Benth.) Raven, Brittonia 16: 286. 1964.
Oenothera hartwegii Benth., Pl. Hartw. 5. 1839

CALYLOPUS HARTWEGII ssp. lavandulifolius (T. & G.) Towner &
Raven, comb. nov. Oenothera lavandulaefolia T. & G., Fl. North Amer.
1: 501. 1840.

CALYLOPHUS HARTWEGII ssp. fendleri (Gray) Towner & Raven,
comb. nov. Oenothera fendleri Gray, Mem. Amer. Acad. Arts II. 4: 45.
1849.

CALYLOPHUS HARTWEGII ssp. pubescens (Gray) Towner & Raven,
comb. nov. Oenothera greggit var. pubescens Gray, Pl. Wright. 1: 72.
S52.

CALYLOPHUS HARTWEGII ssp. filifolius (Eastw.) Towner & Raven,
comb. nov. Oenothera tubicula var. filifolia Eastw., Proc. Calif. Acad.
Ser Hil 1; 72.1897.

CALYLOPHUS HARTWEGII ssp. toumeyi (Small) Towner & Raven,
comb. nov. Galpinsia toumewi Small, Bull. Torrey Bot. Club 25: 317.
1898.

CALYLOPHUS HARTWEGII ssp. maccartii (Shinners) Towner & Raven,
comb. nov. Calvlophus hartwegu var. maccartu Shinners, Sida 1: 343.
1964.

CALYLOPHUS TUBICULA (Gray) Raven, Brittonia 16: 286. 1964.
Oenothera tubicula Gray, Pl. Wright. 1: 71. 1852.

CALYLOPHUS SERRULATUS (Nutt.) Raven, Brittonia 16: 286. 1964
(published in error as Calvlophus serrulata, since the gender of Calvo-
phus is masculine). Oenothera serrulata Nutt., Gen. North Amer. Pl. 1:
246. 1818.

CALYLOPHUS DRUMMONDIANUS Spach, Ann. Sci. Nat. Bot. II. 4: 272.
1835 (published as Calvlophis drummondiana).

CALYLOPHUS DRUMMONDIANUS ssp. berlandieri (Spach) Towner &
_ Raven, comb. nov. Calvlophis berlandiert Spach, Ann. Sci. Nat. Bot. IT.
4: 273. 1835.

Calylophus australis Towner & Raven, sp. nov. A Calvlopho serru-
_ lato similis, pubescentia sparsa trichomatibus crassis saepe incurvatis
_ differt; foliis oblanceolatis, interdum linearibus, longissimis 15-35 mm
_ longis, 1-4 mm latis, plerumque grosse serratis; petalibus 7-13 mm
_ longis; arenosum ‘‘Texas Gulf Coast” secus. Herba suberecta vel effusa,
15-5 dm. alta, simplex vel basi ramosa. Folia longissima pubescentia
sparsa appressa sursum deorsumque, linearia vel oblanceolata, sub-

244 MADRONO [ Vol. 20

integra vel grosse serrata, subsessilia, sursum deminuta solum exigue.
Tubus hypanthii infundibuliformis, basi tubularis, in sectione trans-
versali subquadratus, costibus 4 prominentibus, 6-12 mm longus, apice
4-9 mm diametro, extus subglaber vel sparse pubescens, intus glaber.
Sepala ovata, 4-7 mm longa, exigue vel manifeste costata, apicibus sub-
ulatis 0.2-1 mm longis. Petala obovata vel obcordata, saepa vadose in-
cisurata, 7-13 mm longa lataque. Stamina biseriata, filamenta epipeta-
lorum 3-6 mm longa, episepalorum ca. 2—4 mm longa; anthera 2—4 mm
longa. Stylus 8-15 mm longus, glaber; stigma discoideum, subquadrat-
um, infra subsulcatum, 1—2 diametro, antheris circumdatum. Capsula
cylindrica, sessilis, 12-28 mm longa, 1-2 mm crassa, sparse pubescens.
Semina brunnea, 0.7-1 mm longa, ovoidea, acute angulata, extremitate
unustantum lateribusque complanata, subgranulosa. Autogama. Numerus
chromosomaticus gameticus, 7 = 7.

Type. Texas. Cameron Co: On Texas route 4, 2.8 miles west of end of
road at Boca Chica. Large population along low ridge at roadside, ca.
100 yards from tidal sandflat. Towner 187, May 29, 1969 (DS 612434-
holotype, RSA, TEX, US). Chromosome determination of holotype:
n — 7 (ring of 14 at meiotic metaphase I).

Distribution. Occasional on sand and shell-hash soils along Texas
Gulf Coast, on shores of bays, and on offshore islands. Occurs from
Brazos and Galveston counties south to the Mexican border. Expected
from northern coast of Tamaulipas. Seventy-four specimens from the
the following herbaria have been examined during the course of this
study: ARIZ, DS, F, GH, LL, MO, NEB, NY, OKL, OKLA, PH, POM,
RSA, SMU, TEX, US, WTU.

Vouchers for chromosome counts. Texas. Aransas Co.: 1.3 miles west
of Copano Village, Towner 182 (DS), 1 plant with probable ring of 14
and 1 plant with 1,; and probable ring of 12; 5.8 miles southeast of
Aransas Pass on State Highway 361, Raven & Gregory 19393 (DS),
2 plants with ring of 14 grown from seed at Stanford. Cameron Co.:
Texas Route 4, 2.5 miles west of end of road at Boca Chica, Towner 188
(DS), 1 plant with probable ring of 14 and 1 plant with 1,;; and prob-
able ring of 12. Jackson Co.: Texas Highway 35, 11.2 miles west of
Palacios, Towner 175 (DS), one plant with 1,;, ring of 12, and 1, of
diminutive chromsomes. Matagorda Co.: 6.5 miles south of Matagorda
on road to coast (Farm Road 2031), Towner 174 (DS), probable ring
of 14. San Patricio Co.: 3.5 miles south of Ingleside on Farm Road 1069,
ca. 4 mile from Corpus Christi Bay, Towner 184 (DS), 2 plants with
ring of 14.

The new species, like Calvlophus serrulatus, differs from C. drum-
mondianus in its smaller flowers, self-compatibility, and pollen sterility
of 30-60‘. The small-flowered species are complex structural hetero-
zygotes, their 14 chromosomes normally forming a ring at diakinesis.
Some individuals of both species exhibit one pair and a ring of 12. Both

1970] TOWNER & RAVEN: CALYLOPHUS 24

C. australis and C. serrulatus seem to be derived from C. drummondt-
anus, which has normal chromosome pairing during meiosis. They alse
seem to have arisen independently from C. drummondianus, as suggested
by their phenetic affinities and geographical ranges.

Calylophus drummondianus ssp. berlandiert and C. australis are very
similar vegetatively, and their flower sizes occasionally overlap. Because
of this and their occasional sympatry, some herbarium material is diif-
cult to identify. Unopened buds may be removed from the sheets, and
the pollen can be stained and checked for fertility. This method can be
used to assign most doubtful specimens to one of the two taxa. Ber-
landier’s type, although from typical C. australis habitat, proved to have
highly fertile pollen. Thus it was assigned to C. drummondianus, and a
new type was required for the structural heterozygote.

Calylophus australis differs from C. serrulatus primarily in the short,
thick, sparse stem hairs, versus the fine, appressed, dense canescence of
the latter species. There are some plants from eastern Texas which have
the pubescence of C. australis, but have longer leaves and taller stature.
Calvlophus australis has short, usually coarsely serrate leaves, and is
sub-erect. Calvlophus serrulatus has a wide range of leaf dimensions, but
the margins are coarsely serrate only in specimens whose leaf size exceeds
that of C. australis. Calvlophus serrulatus occurs throughout the Great
Plains, occupying a wide range of soil types, while C. australis is en-
demic to sandy soil, generally along the southern coast of Texas. The
two species seem to come into contact only in inland eastern Texas. In
Brazos and Madison counties, the populations are australis-like, and in
neighboring counties to the north the plants combine the pubescence of
C. australis and the large leaves of eastern C. serrulatus. These inter-
mediate populations seem to be isolated slightly from the main distribu-
tions of the two species. They may be occupying an area of former
hybridization or they may be further autogamous derivatives of C.
drummondianus.

Department of Biological Sciences, Stanford University
LITERATURE CITED

Munz, P. A. 1929. Studies in Onagraceae IV. A revision of the subgenera Salpingia

and Calylophis of the genus Oenothera. Amer. J. Bot. 16: 702—715.
. 1965. Onagraceae. In North Amer. FI. Ser. 2, part 5.

Raven, P. H. 1964. The generic subdivision of Onagraeae, tribe Onagreae. Brittonia
16: 276-288.

————. 1969. A revision of the genus Camissonia (Onagraceae). Contr. U. S.
Natl. Herb. 37: 161-396.

SHINNERS, L. H. 1964. Calylophus (Oenothera in part: Onagraceae) in Texas. Sida
1: 337-345.

TWO NEW SPECIES AND SOME NOMENCLATURAL CHANGES
IN OENOTHERA SUBG. HARTMANNIA (ONAGRACEAE)

PETER H. RAVEN and DENNIS R. PARNELL

For the past three years, we have been engaged in a detailed bio-
systematic investigation of Oenothera subg. Hartmannia. At this time,
it appears desirable to record some of our findings which affect the
names in the group so that these names will be available for other pub-
lications. This work has been supported by a series of grants from the
National Science Foundation, most recently by GB-7879X.

Oenothera platanorum Raven & Parnell, sp. nov. Herba perennis
basi pauci- vel multiramosa, radice crassa, 0.5—5.6 dm alta, strigulosa.
Folia anguste elliptica vel elliptica, raro anguste ovata, subserrata vel
subintegria; folia basalia rosulata, 1-7 cm longa, 4-14 cm lata, interdum
sinuato-pinnatifida, subglabria vel strigulosa praesertim ad nervos cos-
tamque; folia caulina 1.2—5 cm longa, 3-11 mm lata; petiolum 3-32
mm longum. Inflorescentia erecta. Sepala sub anthesi connata, apicibus
sublatis ad 1.5 mm longis, 7.5—11.5 mm longa, 1.5—2 mm lata, strigulosa.
Petala rosea, 8-14 mm longa, 8—12.5 mm lata. Filamenta 4-9 mm longa;
antherae 2.5—3.5 mm longae. Lobi stigmatis 2-4 mm longi. Stylus 12—19
~ mm longus. Tubus hypanthii 9-14 mm longus, apice 1.5—3 mm diametro,
extus strigulosus. Capsula clavata vel anguste obovoidea, 9-14 mm longa,
3—4 mm crassa, 4-angulata, in quoque angulo vulvulaque costa promi-
nente, loculae subdistinctae; pedicellus 4-15 mm longus. Semina dilute
brunnea, 0.7-0.9 mm longa, 0.3—0.5 mm crassa, anguste obovoidea vel
interdum ovoidea, ad columnam persistentem centralem affixa. Numerus
chromosomaticus gameticus, n = 7.

Typé “Eéxas, Cochise Co: Near Fort Huachuca. Lemmon 2704, May
1846 (F 99335-holotype, F, G, GH, US).

Additional specimens examined. ARIZONA. Cochise Co.: Fort Hua-
chuca, base of Huachuca Mts., Mearns 1527 (US); near Fort Hua-
chucha, Wilcox 190 (US); Bear Creek Huachuca Mts., Goodding 272
(RSA); Garden Canyon, Huachuca Mts., Kearney & Peebles 14070
(GH, US); Huachuca Mts., Holkner 1662 (DS, US), Toumey in 1894
(GH, US); Hereford, Jones in 1947 (POM). Gila Co.: Rio San Carlos,
Mohr 269 (US). Pima Co.: Fort Lowell, Tucson, Thornber 457 (DS,
MO, US). Pinal Co.: near Sacaton, Harrison 1778 (US). Santa Cruz
Co.: Sycamore Canyon, Mason 1685 (MEX); near Canelo, Arnold in
1938 (DS, GH).

This proposed new species is most closely related to the widespread
rosea L’Her. ex Ait., to which it is superficially similar. In that species,
however, the hypanthial tube is only 5-7 mm long and the petals only
5—8 mm long and 4—6.5 mm wide. More importantly, O. rosea is a com-

246

1970] RAVEN & PARNELL: OENOTHERA 247

plex structural heterozygote, the chromosomes forming a ring of 14 at
meiotic metaphase I (Raven and Parnell, unpubl.), whereas O. plata-
norum, as determined by an examination of 3 plants grown from the
progeny of Parnell 1031, from Sycamore Canyon, Santa Cruz Co., Ari-
zona, forms 7 bivalents at meiotic metaphase I. Like other complex struc-
tural heterozygotes in Onagraceae, O. rosea has only 40-60 per cent
stainable pollen, O. platanorum normally more than 95 per cent. Although
it is at present known only from southern Arizona, O. platanorum will
doubtless eventually be discovered in adjacent northern Mexico, since
it occurs within a few miles of the international border.

Oenothera texensis Raven & Parnell, sp. nov. Herba perennis erecta
2.5—-5 dm alta, basi pauci- vel multiramosa, radice crassa, strgulosa
sparse hirsutaque. Folia elliptica vel anguste ovata, raro ovata, serrulata
vel sinuato-pinnatifida (praesertim basalia), subglabria vel sparse strigu-
losa praesertim ad nervos costamque, raro dense hirsuta, 2.5—4 cm longa,
8-18 mm lata; petiolum 4—21 mm longa. Inflorescentia erecta. Sepala
sub anthesi connata, apicibus subulatis ad 2 mm longis, 15—18 mm longa,
2-4 mm lata, strigulosa. Petala rosea, 12-21 mm longa, 10-20 mm lata.
Filamenta 9-13 mm longa; antherae 3.5-6 mm longae. Lobi stigmatus
3.5-6 mm longi. Tubus hypanthii 15-21 mm longus, apice 3-4 mm
diametro, extus dense strigulosa. Capsula obovoidea, 8.5—14 mm longa,
3.5—6 mm crassa, valde 4—angulata, in quoque angulo valvulaque costa
prominente, loculae subdistinctae; pedicellus 7-12 mm longus. Semina
delute brunnea, 0.8-1 mm longa, 0.2-0.3 mm crassa, obovoidea vel
ovoidea, ad columnam persistentem centralem affixa. Numerus chromo-
somaticus gameticus, n = 7,

Type. Texas. Jeff Davis Co: Upper Limpia Canyon near Mt. Liver-
more, Ferris & Duncan 2539 (DS 124606-holotype, MO), July 9-12,
1921.

Additional specimens examined. TEXAS. Jeff Davis Co.: Limpia
Canyon, Nealley 145 (F); Fort Davis, Young in 1918 (US); 5 miles
n w of McDonald Observatory, /unes & Moon 1141 (GH, TEX); Davis
Mts., Palmer 34376 (US).

MEXICO. Coahuila. Santa Rosa Mts., Marsh 1375 (TEX); Sierra
del Carmen, Canon de Sentenela on Hacienda Piedra Blanca, Wynd <
Mueller 513 (MICH,S); Muzquiz, Marsh 643 (TEX). Sinaloa. Ocura-
hui, Sierra Surotato, Gentry 6343 (MICH). Tamulipas. Summit of Cerro
Zamora, Sierra de San Carlos, Bartlett 13750 (MICH); Mesa de Tierra,
vicinity of San Jose, Bartlett 10452 (MICH).

Oenothera texensis differs markedly from both O. rosea and O. plata-
norum, its closest relatives, in its much larger flowers. Like O. plata-
norum, it regularly forms seven bivalents at meiotic metaphase I (deter-
mined in an examination of four plants from the progeny of Parnell
1029, from along stream bed, 6.2 miles north of city limit of Fort Davis,

248 MADRONO [Vol. 20

Jeff Davis Co., Texas); and also like that species, it regularly has 95
per cent or more stainable pollen.

OENOTHERA EPILOBIIFOLIA H.B.K. ssp. EPILOBIIFOLIA, Nov. Gen. &
Sp. 6: 92. 1823. O. multicaulis R. & P. var. tarquensis sensu Munz,
Amer. J. Bot. 19: 757. 1932; North Amer. FI. II. 5: 81. 1965, pro parte;
non O. tarquensis H.B.K., Nov. Gen. & Sp. 6: 91. 1823.

OENOTHERA EPILOBIFOLIA ssp. cuprea (Schlecht.) Raven & Parnell,
comb. nov. O. cuprea Schlecht., Linnaea 12: 269. 1838. O. multicaulis
R. & P. var. tarquensis sensu Munz, Amer. J. Bot. 19: 757. 1932; North
Amer. FI. 11. 5: 81. 1965, pro parte; non O. tarquensis H.B.K., Nov.
Gen. & Sp. 6: 91. 1823.

In this subspecies, which is found from central Mexico to Costa Rica
and again in southern Colombia, the yellow petals at anthesis have a
bright red blotch in the lower third to half; in ssp. epilobiifolia, they are
entirely yellow. Oenothera epilobiifolia ssp. epilobiifolia is found in cen-
tral and northern Colombia and adjacent Venezuela. In both subspecies,
the petals fade red after fertilization.

OENOTHERA MULTICAULIS R. & P., Fl. Peruv. 3: 80, t. 317. 1802.
O. tarquensis H. B. K., Nov. Gen. & Sp. 6: 91. 1823. O. multicaulis var.
tarquensis (H. B. K.) Munz & Johnston, Contr. Gray Herb. 75: 18. 1925.

Our unpublished investigations have shown that this species, in the
sense of Munz (Amer. J. Bot. 19: 755-765. 1932), consists of two bio-
logical entities. One of these, corresponding largely to his var. tvpzca, is
a complex structural heterozygote which regularly forms a ring of 14
chromsomes at meiotic metaphase I and is best known from high eleva-
tions in Peru and Bolivia. This relatively small-flowered species also
ranges north to Ecuador, however, and as shown by a comparison be-
tween recent collections from the Rio Tarqui (Valley of the Rio Tarqui,
a few km s. of Cuena, Giles 43a) and the type of O. tarquensis H. B. K.
(P), includes that entity. The oldest available name for the larger-
flowered, bivalent-forming species in then O. epilopufolia, but it cor-
responds largely to the entity that has been known as O. multicaulis var.
tarquensis (H. B. K.) Munz & Johnston. Like other complex structural
heterozygotes in Onagraceae, O. multicaulis forms only 40 to 60 per
cent stainable pollen, whereas in O. epilobufolia the plants regularly
have 95 per cent or more stainable pollen. A few popualtions in southern
Ecuador and Peru which are evidently not complex structural heterozy-
gotes are currently under investigation.

OENOTHERA KUNTHIANA (Spach) Munz, Am. J. Bot. 19: 759. 1932.
Hartmannia domingensis Urban & Ekman, Ark. Bot. 23A: 28. 1931.
Oenothera domingensis (Urban & Ekman) Munz, North Amer. FI. II.
32°82, 1965.

An examination of the holotype of Hartmannia domingensis (S) has

1970] RAVEN & PARNELL: OENOTHERA 249

shown it to be referable to this widely distributed complex structural
heterozygote (Raven and Parnell, unpubl.). It has hitherto been com-
pared with O. rosea, to which it is only distantly related, and this has
led to its continued recognition as a species.

OENOTHERA SPECIOSA Nutt., Jour. Acad. Nat. Sci. Philadelphia 2:
119. 1821. O. delessertiana Steud., Nom. Bot. ed. 2. 2: 206. 1841. O.
speciosa var. childsiu (Bailey) Munz, Leafl. W. Bot. 2: 87. 1935.

Although populations of this species from north Texas northward are
often diploid (n = 7), with white flowers that open at sunset, whereas
those from central Texas southward and also common in cultivation and
occasionally established elsewhere are usually tetraploid (n = 14),
with rose-purple flowers that open near sunrise, intensive studies in the
field have shown that these correlations do not always hold true (Raven
and Parnell, unpubl.). It seems best, therefore, to group all of these
plants in one species, without subdivision, and to describe the character-
istics of particular populations of interest rather than to accord them
distinctive Latin names.

SUMMARY

Critical remarks on Oenothera subg. Hartmannia are presented. Two
new species, O. platanorum and O. texensis, which for bivalents at
meiotic metaphase I, are segregated from the complex structural hetero-
zygote, O. rosea, which forms a ring of 14 chromosomes at meiotic meta-
phase I. The pair-forming O. epilobifolia (O. multicaulis var. tarquen-
sis Of most authors) is segregated from the complex structural hetero-
zygote O. multicaulis, whereas O. domingensis is synonymized with O.
kunthiana and O. delessertiana with O. speciosa. With these changes, we
currently recognize the following species of subg. Hartmannia as valid:
O. seifrizii Mung, O. epilobiifolia HBK., O. multicaulis R. & P., O. tet-
raptera Cav., O. kunthiana (Spach) Munz, O. deserticola (Loesener )
Munz, O. purpusu Munz, O. texensis Raven & Parnell, O. platanorum
Raven & Parnell, O. rosea L’Her. ex Ait., and O. speciosa Nutt. All of
these species are diploid (n= 7) except for the last, in which auto-
tetraploids (n = 14) occur commonly: and all of the diploids form biva-
lents at meiotic metaphase I except for O. kunthiana, O. multicaulis,
and O. rosea, which are complex structural heterozygotes. Oenothera
speciosa is self-incompatible, all the others are self-compatible, with
varying degree of out crossing.

Department of Biological Sciences, Stanford University
Department of Biological Science, California State College, Hayward

(Editorial note. The following series of papers dealing with the Galapagos
Islands are published together here in anticipation of the Flora of the Galapagos
Islands being edited by Ira L. Wiggins and Duncan M. Porter. — J. H. T.)

STUDIES ON PLANTS OF THE GALAPAGOS ISLANDS. I.
NEW SPECIES AND COBINATIONS

TRA L. WIGGINS

Galium galapagoense Wiggins, sp. nov. Herba gracillima pluribus
caulibus e basi communi tenuibus quadrangulis adscentibus vel scan-
dentibus usque 1 m longi sparse scaberulis. Folia plerumque 4-verticillata
3—7 cm distantia; lamina membranaceo-herbacea elliptica vel obovato-
elliptica apice rotundata breviter apiculata basi sub cuneata, utrinque
sparse pilosa margina scabra, 5—17 mm longa, 2—9 mm lata. Flores
minuti sessile in cymis axillaribus paucifloris dispositi; ovarium sub-
globosum unicate pilose 0.6—0.8 mm diamentiens; corolla rotata flaves-
cens in lacinias 4 ovatas acuminatus 1 mm longas divisa; stamina
paullum infra faucem inserta, filamentia brevissime, antherae parvae
rotundato-ellipsoidae vix 0.3 mm longae; stylus 1—1.4 mm longus in
ramulis 2 breves subdivergentus corolla laciniis circa dimidio breviores
divisus; stigmatibus capitatis coronatis. Fructus subglobosus 1.5—2
mm diametiens ruber breve papillatus.

Holotype. Ecuador. Isla Santa Cruz. Flanks of Cerro Copa, near
center of island, altitude about 570 m. Sigvard Horneman 2, Feb. 8,
1964 (DS).

This species is unlike any of the known species of Galium on the
mainland of South America, particularly in that the dichasium has each
terminal flower sessile between the subtending leaves, and with the slen-
der branches of the next order of the inflorescence arising in the axils
between the leaves and the terminal flower. This system of branching
and flowering repeats itself two to four times in each axillary dichasium.
The arrangement is totally unlike that in Relbunium, to which the
plant was referred by Stewart (Proc. Calif. Acad. Sci. 1V. 1: 146. 1911).
Occasionally one of the axillary branches in a dichasium fails to develop,
and then the morphologically terminal flower appears to be axillary to
a leaf borne on a straight branch.

The minute flowers are distinctive, also, for they are no more than
2—2.4 mm wide when fully open, and the cup is about 1 mm deep.

The leaves are distinctly 3-nerved, and generally are considerably
thinner than those of Galium ferrugineum Krause, to which it would
key in Macbride’s Flora of Peru (Field Mus. Nat. Hist., Bot. Ser. 13 (6):
260. 1936). Galium ferrugineum has 1-nerved leaves, with pinnately
arranged secondary veins, and the tissue is much more coriaceous than
that in G. galapense.

250

1970] WIGGINS: GALAPAGOS ISLANDS aie

Galium has not been reported, as such, from the Archipiélago de Colon
by earier workers.

Passiflora colinvauxii Wiggins, sp. nov. Herba scandens sparsim
puberula mox glabra, caule angulato; stipulae setaceae falcatae 2—4
mm longae; folia membrancea bilobata integra trinervia 3—5 cm longa
7—16 cm lata basi rotundata vel subtruncata, lobis oblongo-lanceolatis
divaricatis acutis vel obtusis lobo medio obsoleto vel brevissimo, petiolo
gracili eglanduloso; bracteae setaceae librae integrae falcatae; tubus
calycis cupulatus 6—7 mm latus glaber, sepalis late oblongis 6—7 mm
longis 3—4 mm latis apice rotundatis; petalae albae membranaceae
2—2.5 mm latae 3.5—6 mm longae anguste oblongae; corona biseriata,
filamentis purpureis vel apicem albis, exterioribus longioribus; operculum
membranaceum plicatum incurvatum breviter fimbriatum. Ovarium
ovoideum glabrum; fructus anguste ovoideus 3—4 cm longus 1.5—2 cm
latus glaber; semina ovoidea-lenticularia 2—2.5 mm longa ca 2 mm lata
minute corrugata fusca.

Holotype. Ecuador. Isla Santa Cruz. In treeless region en rounte to
El] Chato (west of village of Bella Vista). Paul A. Colinvaux 443, July
30, 1966 (DS).

The label on the holotype further states, “Creeping plant, covering
large areas of ground and climbing on trees. Flowers white, fruits green.”

This species is strikingly different from the other two passifloras known
to occur on the Galapagos Islands, both in the broad, lunate leaves and
in the narrowly ovoid fruits. The ocelae at the base of the leaf blade are
rather conspicuous, but the herbage is eglandular, and no glands occur
on the petioles.

Sicyocaulis Wiggins, gen. nov. Cucurbitaceae. Herba graciles scan-
dentes radice perennante. Folia integra vel plus minusve lobata; petiolus
gracilis quam lamina paullo brevior. Cirrhi bifidi vel rarissime simplices.
Flores monoici minuti lutei. Flores masculi pedicellati ad apices race-
morum dispositi; pedunculis gracilibus foliis duplo longioribus. Recepta-
culum complanatum latum. Sepala dentiformia minuta. Corolla profunde
5-partida segmentis ovatis vel lanceolatis integris. Stamina 4, filamentis
connatis basi tubo receptaculii inserta; antherae oblongae omnes bilocu-
lares in capitulum subglobosum conniventes. Pistillodeum nullum. Flores
feminei pedicellati solitarii vel 2—3 ad bases racemorum dispositi,
perianthium ut in mare. Staminodia nulla. Ovarium ovoideum unilocu-
laris, stylus gracilis in stigmata non profunde bifida divisus; ovulum
unum pendens. Fructus ovoideus rostratus ad basem parce minutusque
spinulosus. Semen solitarium oblongus compressum testa firma laevis
vel ad apicem minute tuberculata.

This plant keys out to the vicinity of Sicvos in Hutchinson (Gen. FI.
Pl. 2: 411. 1967) and at first glance resembles that genus. But the
flowers are much smaller than any I have seen in that genus, the pistil-

252 MADRONO [Vel. 20

late flowers are not gathered into heads of several flowers, each flower
being slenderly pedicellate. Further, the forward-pointing, few spines
at the base of the fruit are quite unlike the radiating, numerous bristles
common on the fruits of Sicvos.

Neither can it be placed in the genus Frantzia, for that genus has the
female flowers borne singly in the axils, or paired in that position with
a male flower. In contrast, our plant has the pistillate flowers borne on
slender pedicels at or near the base of the flowering part of the racemes,
these always being long-stalked and the flowers removced a considerable
distance from the stem on which the peduncle is borne.

Type species. Sicyocaulis pentagonus Wiggins.

Sicyocaulis pentagonus Wiggins, sp. nov. Plantae monoicae scandens
3—5 m altae; rami gracillimi angulatosulcati sparse puberula vel
subglabra internodia 1—2 dm longa; cirrhi bifidi graciles elongati
elabri ramis 10—25 cm longis; folia alterna, petiolis gracilibus 2—8
(—10) cm longis, laminae minute scaberulae, cordato quinquelobae
2—12 cm logae et latae, sinu basali 2—3 cm profundo; inflorescentiae
racemosae 1—5 cm longae elongatae; pedicelli 3—6 mm longi filiformes
ad apicem clavatos; sepals minute; lobi corollae oblongi 0.6—1 mm
longi 0.4—0.6 mm lati acuti vel minute apiculati flavascentes glabri;
columna staminum gracilis 1—1.4 mm alta antheris ca 0.6 mm longis
interne ad columnam adnatis externe libris; fructus ovoideus 10—15
mm longus 4—5 mm latus rostratus rostello 2.5—3 mm longo, fructu
supra basin sparce spinoso parallelo ad axem.

Holotype. Ecuador. Isla Santa Cruz. Along trail about 1 km south of
Bella Vista, along trail from Bahia Académia, altitude about 225 m.
I. L. Wiggins 18679, Feb. 21, 1964 (DS).

PLANTAGO PARALIAS Decne. var. pumila (Hook.f.) Wiggins, comb.
nov. P. tomentosa Lam. var. ?pumila Hook.f., Trans, Linn. Soc. Lond.
20: 194. 1847.

Hooker’s specimen was inadequate, consisting only of three or four
very depauperate plants that scarcely showed the characters needed to
make an identification. Its relationship, as disclosed by more and larger
plants collected in 1964, clearly show its relationship with Plantago
paralias. Known from Islas San Cristébal, Santa Cruz and Santa Maria.

VALLESIA GLABRA (Cav.) Link var. pubescens (Anderss.) Wiggins,
comb. nov. V. pubescens Anderss., Kongl. Svensk. Vet.-Akad. Handl.
ESobeelO>..1So5)

Differing from var. glabra only in having a fine, closely arranged, erect
indument of simple, non-glandular hairs on the twigs, petioles, inflores-
cences and under surfaces of leaf blades, and certainly not worthy of a
higher nomenclatoral rank.

Known only from the Galapagos Islands and there occurring on at

1970] BURCH: GALAPAGOS CHAMAESYCE 299

least six of the islands. Plants from Central America sometimes ap-
proach this, but have fewer hairs on the young vegetation.

Division of Systematic Biology, Stan‘ord University

ANEW COMBINATION IN TRICHONEURA FROM THE
GALAPAGOS ISLANDS

JoHN R. REEDER and CHARLOTTE G. REEDER

TRICHONEURA LINDLEYANA (Kunth) Ekman var. albemarlensis ( Rob-
ins. & Greenm.) Reeder & Reeder, comb. nov. Leptochloa albemarlensis
Robins. & Greenm., Amer. J. Sci. III. 50:145. 1895.

Known from Islas Genovesa, Isabela, Pinta, San Salvador, Santa
Cruz, and Santa Maria.

Department of Botany, University of Wyoming, Laramie

A NEW COMBINATION IN CHAMAESYCE FROM THE
GALAPAGOS ISLANDS

DEREK BURCH

CHAMAESYCE nummularia (Hook. f.) Burch var. glabra (Robins. &
Greenm.) Burch, comb. nov. Amer. J. Sci. III. 50:144. 1895. Euphorbia
nummularia var. glabra Robins. & Greenm., Amer. J. Sci. IIT. 50: 144.
1895.

Differs from var. nummularia only in being completely glabrous.

Known only from Isla Santa Maria. Recollected by Uno Eliasson in
1966 and 1967 and apparently well established at altitudes of 5-15 m
at Las Cuevas and at Black Beach.

University of South Florida, Tampa

NEW COMBINATIONS IN THE CYPERACEAE OF THE
GALAPAGOS ISLANDS

TETSUO KOYAMA

CYPERUS POLYSTACHYOS Rottboll ssp. holosericeus (Link) T. Ko-
yama, comb. nov. C. holosericeus Link, Hort. Berol. 1:317. 1827. C. mi-
crodontus Torr., Lyceum Nat. Hist. New York 3:255. 1836. C. gatesii
Torr., Lyceum Nat. Hist. New York 3:255. 1836. C. microdontus var.
texensis Torr., Lyceum Nat. Hist. New York 3:430. 1836. C. fugax
Liebm., Vidensk. Selsk. Skr. Kjoeb. ser. 5. 196. 1851. C. incons picuus
Liebm., Viednsk. Selsk. Skr. Kjoeb. ser. 5. 197. 1851. C. liebmannii
Steudel, Syn. Pl. Glumac. 2:7. 1854. C. texensis Steudel, Syn. Pl. Glumac.
2:9. 1854. C. polvstachyos var. leptostachyus Bockeler, Linnaea 35:478.
1868. C. polystachyos var. laxiflorus C. B. Clarke, in Urban, Symbol.
Antill. 2(1):17. 1900. C. polvstachvos var. leptostachyus {. incons picuus
Kukenth., in Engler, Pflanzenreich IV. 20:372. 1936. C. polystachyos

254 MADRONO [Vol. 20

var. leptostachyus f. fugax Kukenth., in Engler, Pflanzenreich IV. 20:372.
1936. C. polystachyos var. texensis Fernald, Rhodora 41:530. 1939.
In the Galapagos Islands, known only from Isla San Salvador.

CYPERUS VIRENS Michx. ssp. drummondii (Torr. & Hook.) T. Koya-
ma, comb. nov. C. drummondu Torr. & Hook., Lyceum Nat. Hist. New
York 3:437. 1836. C. virens var. drummondu Wikenth., in Engler,
Pflanzenrecih IV. 20. 181. 1936. C. surinamensis sensu Anderss., Kongl.
Vetensk. Acad. Handl. 1853:153. 1855; sensu Robins., Proc. Amer. Acad.
Arts 30:129. 1902; sensu Eliasson, Svensk. Bot. Tidskr. 59:478. 1965,
non Rottboll.

In the Galapagos Islands known from Islas San Cristobal and Santa
Cruz

RHYNCHOSPORA NERVOSA (Vahl) Bockeler ssp. ciliata (Vahl) T. Ko-
yoma, comb. nov. Dichomena ciliata Vahl, Enum Pl. 2:246. 1806. R.
cizata Kukenth., Bot. Jahrb. Syst. 56 (Beibl. 125):16. 1921. R. nervosa
var. ciliata Kukenth., Bot. Jahrb. Syst. 75:295. 1951.

Known in the Galapagos Islands from Islas Isabela, San Cristobal, and

Santa © Buz,
New York Botanical Garden, Bronx

A NEW VARIETY OF OPUNTIA MEGASPERMA
FROM THE GALAPAGOS ISLANDS

J. LUNDH

OPUNTIA MEGASPERMA Howell var. mesophytica J. Lundh, var. nov.
Plantae arborescentes vertice apertae; articuli caulium atrovirides;
spinae 2.5—3.9 cm longae aureofulvae albescentes apicem fuscae; flores
parvi 6—8 cm longi; fructus parvi 4—6 cm longi.

Plants more or less arborescent, with open crowns and slender, vertical
branches 2—6 m tall; trunk up to 40 cm in diameter, brownish; larger
terminal joints dark green becoming gray-green, obovate to elongate,
12-20 cm long, 5.5—9.5 cm wide, 0.8—1.7 cm thick; leaves 3-7 mm long;
areoles ovate, 1.5—2 mm in diameter, typically 1.8-2.2 cm apart; spines
often nearly absent, golden brown and banded, becoming bone white with
a rather dark tip, 1-17 per areole, 2.5-3.9 cm long, 0.5—1 mm in diam-
eter basally; flowers about 6 cm in diameter, 6—8 cm long; fruits 4—6 cm
long, 2.7-3.6 cm in diameter; seeds 7-10 mm long, 5-8 mm wide, 2—4
mm thick.

Holotype. Chatham Island (Isla San Cristobal), Galapagos Islands,
Ecuador, J. Lundh s.n.,in 1962 (AHFH).

Known only from the western end of Isla San Cristobal, where it has
been collected by Lundh and by E. F. Anderson.

NEW COMBINATIONS IN THE COMPOSITAE OF THE
GALAPAGOS ISLANDS

ARTHUR CRONQUIST

DARWINIOTHAMUS TENUIFOLIUS (Hook. f.) Harling var. glabriuscu-
lus (Stewart) Cronquist, comb. nov. Erigeron lancifolius var. glabrius-
culus Stewart, Proc. Calif. Acad. Sci. TV. 1:151. 1911. E. lancifolius
Hook. f., Trans. Linn. Soc. London 20:208. 1847. E. tenuifolius ssp.
lancifolius Solbrig, Contr. Gray Herb. 191:43. 1962. D. lancifolius Har-
ling. Acta Horti Berg. 20(3):115. 1962.

Known only from Isla Isabela.

DARWINIOTHAMNUS TENUIFOLIUS (Hook. f.) Harling var. glandulo-
sus (Harling) Cronquist, comb. nov. D. lancifolius ssp. glandulosus
Harling, Acta Horti Berg. 20(3):117. 1962.

Known only from Islas Fernandina and Isabela.

POROPHYLLUM RUDERALE (Jacq.) Cass. var. macrocephalum (DC.)
Cronquist, comb. nov. P. macrocephalum DC., Prodr. 5:648. 1836. P.
ruderale ssp. macrocephalum R.R. Jhtn., Univ. Kansas Sci. Bull. 48:233.
1969.

In the Galapagos, known from 11 islands. A widespread weed.

PSEUDOLEPHANTOPUS spiralis (Less.) Cronquist, comb. nov. Distrep-
tus spiralis Less., Linnaea 6:690. 1831. Spirochaeta funcku Turcz., Bull.
Soc. Imp. Naturalistes Moscou 24:167. 1851. Chaetospira funcku Blake,
J. Wash. Acad. Sci. 25:331. 1935. P. funcku Philipson, J. Bot. 76:301.
1938.

Theer is a small problem about the typification of Pseudele phanto pus
spiralis, inasmuch as the original publication of Distreptus spiralis Less.
gave the locality as Jamaica. The species is not otherwise known there,
although it is found occasionally in the Lesser Antilles and is wide-
spread on the South American mainland. However, the description is so
clearly that of the present species that it seems necessary to take up
Lessing’s epithet.

In the Galapagos Islands known from a single collection (Wiggins &
Porter 632, DS), taken at an altitude of about 570 m on Isla Santa Cruz.

Lecocarpus is a genus of three closely allied species, endemic to the
Galapagos Islands. Two of the species have usually been referred to the
related genus Acanthospermum, but the unity of the group is obvious
and has often been remarked. In the course of my work I learned that
Tod F. Stuessy, then a graduate student at the University of Texas had
independently arrived at similar conclusions in connection with his study
of Acanthospermum. Accordingly, the necessary new combinations are
attributed here to Cronquist and Stuessy. As this manuscript goes to
press (June 1970) we have learned that Uno Eliasson would have made
the combination L. lecocarpoides, had we not done so.

Des)

256 MADRONO [Vol. 20

LECOCARPUS lecocarpoides (Robins. & Greenmn.) Cronquist & Stues-
sv, comb. nov. Acanthospermum lecocar poides Robins. & Greenm., Amer.
J. Sci. TIT. 50. 141. 1895.

LECOCARPUS leptolobus (Blake) Cronquist & Steussy, comb. nov.,
Acanthospermum leptolobum Blake, J. Wash. Acad. Sci. 12:204. 1922.

New York Botanical Garden, Bronx

NEW COMBINATIONS AND TAXA IN THE CACTACEAE
OF THE GALAPAGOS ISLANDS
EDWARD F. ANDERSON and DAvip L. WALKINGTON

JASMINOCEREUS THOUARSII (Weber) Backeb. var. delicatus (Daw-
son) Anderson & Walkington, comb. nov. J. howellit Dawson, J. Cact.
Succ. Soc. Amer. 34:71. 1962. J. howellii var. delicatus Dawson, J. Cact.
Succ. Soc. Amer. 34:71. 1962.

Known from Islas Bartolomé, San Salvador, and Santa Cruz.

JASMINOCEREUS THOUARSII (Weber) Backeb. var. sclerocarpus (K.
Sch.) Anderson & Walkington, comb. nov. Cereus sclerocarpus K. Sch.
in Robins., Proc. Amer. Acad. Arts 38:179. 1902. J. sclerocarpus Bac-
keb., Jahrb. Deutsch. Kakteen-Ges. 2:24. 1944.

Known from Islas Fernandina and Isabela.

OPUNTIA ECHIOS Howell var. zacana (Howell) Anderson & Walking-
ton, comb. nov. O. zacana Howell, Proc. Calif. Acad. Sci. IV. 21:48.
1933. O. galapageia Hensl. var. zacana Backeb., Cactaceae 1:562. 1958.

Known only from tiny Isla Seymour, off the northeast segment of Isla
Santa Cruz.

OPUNTIA GALAPAGEIA Hensl. var profusa Anderson & Walkington, var.
nov. Habitus variabilis; folia parva, usque ad 4 mm longa; trichomata
numerosa; spinae non valde dimorphe nec pungentes; fructus profusi
17-25 mm longi, 22—27 mm diametro, spinas raro ferens.

Plants variable, mostly prostrate, or shrubby to arborescent, 1-3 m
tall; trunk, when present, flaky and reddish; larger terminal joints 21-38
cm long, 18-26 cm wide, 1.8—2.7 cm thick; leaves small, up to 4 mm
long; areoles 4—6 mm in diameter, typically 2.2-3.3 cm apart; spines
bristly, pungent only on new stem joints; fruits greenish, becoming yel-
low-green to brown, with glochids on some, without spines, nearly glo-
bose, 1.7—2.5 cm long, 2.2—2.7 cm in diameter, with a deep, small, green-
ish brown umbilicus, the fruits often profuse, up to 82 on a single joint;
seeds 2—3 mm long, 1.5—2 mm wide, | mm thick.

Holotype. Northwest corner of Isla Rabida, Galapagos Islands, Ecua-
dor, Anderson 2546 (RSA).

Occurring from near sea level to about 100 m elevation on Isla Rabida,
the only island from which it is known.

Whitman College, Walla Walla, Washington
California State College, Fullerton

NOTES ON GALAPAGOS EUPHORBIACEAE
GRADY L. WEBSTER
CROTON

As noted by a number of observers (especially Stewart, Proc. Cal.
Acad. Sci. IV. 1:206—209, 215-216. 1911), plants of Croton are an im-
portant component of the woody vegetation of the Galapagos at both
low and high elevations; and in the arid zones Croton may be the veg-
etational dominant. Although Andersson (Kongl. Svenska Freg. Eugenies
Resa, Bot. 105, 106. 1857) recognized six species of Galapagos Croton,
Mueller Argoviensis (DC., Prodr. 15(2):604—-606. 1866) amalgamated
all of the Galapagos taxa into the single species Croton scouleri, and
later workers such as Robinson and Stewart have followed Mueller’s
circumscription.

Svenson (Amer. J. Bot. 33:458-460. 1946) not only treated all the
Galapagos Crotons as a single species, but combined C. scowleri with the
South American species C. rivinifolius HBK. There is certainly a strik-
ing resemblance between these two species, and Svenson may be correct
in postulating that the Galapagos taxa were derived by long-distance
dispersal from Ecuadorian populations of C. rivinifolius. However, the
Galapagos plants differ from the mainland ones in a number of respects,
especially in the seeds, which are smooth to pitted rather than ribbed
as in C. rivinifolius and related species. Consequently, it seems best to
follow Mueller and retain C. scouleri as a distinct polytypic species
endemic to the Galapagos. The system of subspecific taxa presented here
to some extent resembles that of J. T. Howell, who examined and anno-
tated a large number of collections, but did not publish his conclusions.

The variation in Galapagos Croton is still not satisfactorily under-
stood despite the rather considerable number of collections made of
these common plants. Unfortunately, many collectors have not carefully
noted the habitat conditions or even the altitude at which specimens
were taken, so that it is difficult to establish the ecological status of the
various proposed taxa. Most of the characters of the taxa recognized
here appear to represent segments of altitudinal clines, and only future
work can establish whether or not sufficient discontinuities exist to
justify maintaining discrete taxa. Inter—island variation is on the whole
less well-defined than altitudinal intra—island variation, but one popu-
lation at least appears to have a distinct seed size. Nevertheless, even
seed size is not consistently diagnostic. It is possible that when the local
populations are more thoroughly collected any attempt to recognize
varieties within the species will have to be abandoned.

The Galapagos populations of Croton may be summarized as fol-
lows.

CROTON SCOULERI Hook. f., Trans. Linn. Soc. London 20:188. 1947.
Lectotype, Isla San Salvador (James), Scouler (K-n. v., A-isotype frag-
ment).

258 MADRONO [ Vol. 20

The populations of this species endemic to the Galapagos are here
classified into four rather ill-defined varieties.

CROTON SCOULERI var. SCOULERI. C. macraei Hook. f., Trans. Linn.
Soc. London 20: 188. 1847. Type, Isla Isabela (Albemarle), Macrae
(K-n. v.). C. albescens Anderss., Kong]. Vetensk. Acad. Handl. 1853:
242. 1855. Type, Isla Santa Maria (Charles), in ‘locis editioribus
umbrosis’, Andersson 208 (A-fragment of type from S.). C. incanus
Anders., Kongl. Vetensk. Acad. Handl. 1853:243. 1855 (non C. incanus
HBK.). Type, Isla San Salvador (James), in ‘locis siccis’, Andersson
(GH-isotype). C. scouleri var. glabriusculus Stewart, Proc. Calif. Acad.
Sci. IV. 1:89. 1911. Type, Isla Pinta (Abingdon), 1000-1650 ft., Stew-
art 1834 (CAS). C. scoulert var. castellanus Svenson, Amer. J. Bot.
22:239. 1935. Type, Isla Genovesa (Tower), Snodgrass & Heller (GH-n.
v.; no collection of Snodgrass and Heller from Genovesa was seen among
the Gray Herbarium specimens, and Svenson may have based the name
on the Genovesa collection of Baur).

Illustrations. Svenson, Amer. J. Bot. 22: pl. 4, figs. 1-3; pl. 9, figs
1,2. 1935.

The majority of lowland Croton populations belong to this highly di-
verse variety. The variation in leaf shape from narrowly linear to
elliptic is so striking that all previous authors have accepted the narrow-
leaved variant as either a species, C. macraet, or a variety. However, this
foliar variation appears to be at least partly related to an altitudinal
cline, linear-leaved plants in the lowlands being replaced by broad-
leaved ones at higher elevations. At some localities (Academy Bay, Isla
Santa Cruz) narrow- and broad-leaved forms evidently occur in the
same population. The transition between the extremes is so continuous
that it seems quite impractical to recognize the narrow-leaved plants as
a distinct variety. These stenophyllous variants must be classified only
at the rank of forma.

CROTON SCOULERI f. macraei (Hook. f.) Webster, comb. nov., based
on C. macraei Hook. f., cited above.

It is possible that the littoral populations with broad densely stellate
leaves, designated as var. castellanus by Svenson, may prove to merit
varietal recognition, particularly if they prove to be characterized by
large seed size (see discussion under var. darwinii). However, on the
basis of presently available specimens no useful diagnostic characters
can be found. The only other outstanding variant of var. scouleri is the
plant from Isla Pinta described as var. glabriusculus by Stewart. The
type specimen (Stewart 1834) is indeed unique in having stellate hairs
with one very long branch on the upper leaf surfaces. However, other
specimens from Isla Pinta do not show this character, so the taxon de-
scribed by Stewart does not appear to make up a discrete population.
It may be conveniently referred to at the rank of forma.

1970] WEBSTER: GALAPAGOS EUPHORBIACEAE 259

CROTON SCOULERI f. glabriusculus (Stewart) Webster, comb. nov.,
based on C. scouleri var. glabriusculus Stewart, cited above.

CROTON SCOULERI var. darwinii Webster, var. nov. Frutex vel arbus-
cula foliis rotundatis depresso-stellatis, seminibus 3.9—4.5 mm _ longis.

Type. Isla Darwin (Culpepper), J. R. Hendrickson H-7, Jan. 29, 1964
(DS—holotype).

This variety is apparently restricted to the two small northernmost
islands, Darwin and Wolf. Some forms of var. scouleri are similar in
leaf shape and pubescence, but these have smaller seeds (2.6-3.6 mm
long). One problematical collection from Isla Daphne (Pool 290, BKL)
has large seeds over 5 mm long but is vegetatively similar to plants de-
scribed from Genovesa and elsewhere by Svenson as var. castellanus.
Although Svenson’s variety has here been synonymized with var. scou-
leri, it might prove to be characterized by large seed size, and it is pos-
sible that var. darwinii would then fall into synonymy under var. castel-
lanus. However, until more complete collections are available, this pos-
sibility must remain only hypothetical. The plants from Isla Darwin
and Isla Wolf in any event have a characteristic aspect, and in the pres-
ent rudimentary state of our knowledge it seems most reasonable to treat
them as a distinct variety.

CROTON SCOULERI var. BREVIFOLIUS (Anderss.) Muell.-Arg., in DC.,
Prodr. 15(2): 605. 1866. C. brevifolius Anderss., Kong]. Vetensk. Acad.
Handl. 1853:105. 1855. Type, Isla Santa Maria (Charles), Andersson
206 (A-fragment of type from S).

As here circumscribed, this variety is restricted to higher altitudes on
Isla Santa Maria. Although Stewart (Proc. Calif. Acad. Sci. IV. 1: 88-89.
1911) recorded this variety from a large number of islands, most of the
collections cited are here referred to var. scouleri. As presently under-
stood, var. brevifolius may be distinguished from var. scouleri by the
loose tomentum on the undersides of the leaves, and from vars. darwintii
and grandifolius by its smaller seeds.

CROTON SCOULERI var. GRANDIFOLIUS Muell. Arg., in DC., Prodr.
15(2):605. 1866. Type, Isla San Salvador (James), Darwin (CGE-
isotype, K-holotype-—n. v.).

Illustration. Svenson, Amer. J. Bot. 22:272, pl. 4, fig. 5. 1935.

Most plants of Galapagos Croton with large leaves (at least 5 cm
long and 3 cm broad) may be assigned to this variety, which is recorded
from higher altitudes on Isla Isabela, Isla San Cristébal, Isla Santa Cruz,
and Isla San Salvador. In general, specimens of var. grandifolius are
easily distinguished from those of vars. darwinii and scouleri by the
larger leaves with flocculent tomentum beneath. It is much more difficult
to find good diagnostic characters to separate var. grandifolius from var.
brevifolius, although the leaves are usually smaller in the latter. Some
specimens from Isla Santa Maria (Howell 9312, Stewart 1837) overlap

260 MADRONO [ Vol. 20

var. grandifolius in size. The seeds of var. grandifolius on Isla Santa
Cruz appear to be larger, but the seeds of the large-leaved populations
on Isabela, San Cristobal, and San Salvador are still unknown. Until
fruiting collections are available, varietal assignment of the specimens
from those three islands must remain provisional.

ACALYPHA

In his synopsis of Galapagos Acalypha, Robinson (Proc. Amer. Acad.
Arts 38: 161-165. 1902) recognized no less than 13 species, including
four described by himself. Pax and Hoffmann (Pflanzenreich IV. 147
(XVI): 30-31, 132-134. 1924) accepted all 13 of Robinson’s species
and placed them into two widely separated taxa: ‘sect.’ Phleoideae in
‘ser.’ Polvgynae-Acrogynae, and ‘sect.’ Cus pidatae in ‘ser.’ Oligogynae.
This disposition, apparently made on the basis of position of spikes,
terminal or axillary, resulted in a completely artificial arrangement of
taxa and is wholly untenable, however the species may be circumscribed.

The treatments of Robinson and of Pax and Hoffmann contrast strongly
with the earlier one of Mueller Argoviensis, who treated all of the Gala-
pagos taxa of Acalypha known to him as seven varieties of the single spe-
cies A. parvula. A detailed analysis of the much larger number of speci-
mens now available shows that Mueller was justified in lumping together
under 4. parvula many of the populations of plants with glandular tri-
chomes. However, the populations of plants with more densely pubescent
leaves and stems lacking glandular trichomes do not appear to inter-
gerade with members of the A. parvula complex on the several islands
where they are sympatric. These non-glandular plants are therefore ac-
cepted as distinct species. The newly described A. wigginsii is to some
extent intermediate between A. parvula and A. sericea and is sympatric
with them on Isla Santa Cruz. The classification thus adopted here, with
five species and several varieties, represents a rather unstable com-
promise between the previous schemes. It, like its predecessors, may have
to be extensively remodeled when better population samples are avail-
able of these still rather poorly understood taxa.

Glandular trichomes absent or nearly so;
young stems densely villose-hirsute

ACALYPHA SERICEA Anderss., Kongl. Vetensk. Acad. Handl. 1853:238.
1855.

This is the most widespread and variable non-glandular species, cor-
responding to A. parvula in the glandular taxa. The species comprises
3 fairly well-marked varieties differing in mean size of leaves, staminate
spikes, and seeds.

ACALYPHA SERICEA Anderss. var. SERICEA. A. parvula y pubescens
Muell.-Arg., Linnaea 34:47. 1865; in. DC., Prodr. 15(2):878. 1866.
Type, Isla Isabela, ‘locis lapidosis regionis inferioris insulae Albemarle’,
Andersson (S-n. v.). The type locality was originally cited as Isla San

1970] WEBSTER: GALAPAGOS EUPHORBIACEAE 261

Cristobal (Chatham), but was corrected in Andersson’s later treatment
(Kongl. Svenska Freg. Eugenies Resa, Bot. 103. 1857).

Leaves 1-2 (-3) cm long, with mostly 10-15 teeth per side; stami-
nate spikes mostly 3-15 mm long; seeds 1.0—1.2 mm long. Recorded
from Marchena (Bindloe) and Pinta (Abingdon); some specimens from
Abingdon are unusually glandular and somewhat approach A. parvula.

ACALYPHA SERICEA Anderss. var. indefessus Webster, var. nov. Suf-
frutex caulibus villosulis eglandulosis, foliis 2.5—5.5 longis, dentibus lat-
eribus c. 15-25, spicis masculis plerumque 5—15 mm longis, seminibus
1.2-1.3 mm longis.

Type. Isla Santa Cruz (Indefatigable), along ‘new road’ from Bahia
Academy to Bella Vista, transition zone, Wiggins 18672, Feb. 9, 1964,
(DS-holotype). Other collections examined, all from the general locality,
include Fournier 243 (DAV), Taylor TT59 (CAS), and Wiggins 18492
(DS).

Endemic to Isla Santa Cruz, in relatively mesic vegetation between
100 and 200 m. The plants resemble var baurii from San Cristobal in
aspect, but have much shorter staminate spikes and styles; the larger
seeds and leaves (with more teeth per side) provide differential features
from var. sericea.

ACALYPHA SERICEA Anderss. var. baurrii (Robins. & Greenm.) Web-
ster, comb. nov. A. bauri Robins. & Greenm., Amer. J. Sci. 50:144. 1895.
Type, Isla San Cristobal (Chatam), southwest end, middle region,
Baur 285 (GH).

Leaves mostly 4-6 cm long, with 23-31 teeth per side; staminate
spikes 30-60 mm long; seeds 1.2 mm long. Known only from the type
collection on San Cristobal.

ACALYPHA FLACCIDA Hook. f., Trans. Linn. Soc. London 20:186. 1847.
A. parvula 7 flaccida Muell.-Arg., Linnaea 34:48. 1865; in DC., Prodr.
15(2): 878. 1866. Type, Isla San Salvador (James), Darwin (CGE).

Known only from the type collection; very similar to A. sericea in
appearance, but with much smaller, non-hispidulous pistillate bracts in
strictly axillary spikes.

ACALYPHA VELUTINA Hook. f., Trans. Linn. Soc. London 20:186. 1847.
A. parvula y pubescens & velutina Muell.-Arg., Linnaea 34:48. 1865; in
DC., Prodr. 15(2):878. 1866. Type, Isla Santa Maria (Charles), Darwin
(CGE). The variety recognized by Hooker, A. velutina 8 minor (Trans.
Linn. Soc. London 20:187. 1847), also based on a Darwin collection from
Charles (CGE), is only a small-leaved form of no taxonomic importance.

Very similar to some forms of A. sericea, but differing in the axillary
spikes and distinctly bullate-rugose leaves. Endemic to Isla Santa Maria.

Glandular trichomes present, at least on pistillate bracts

Acalypha wigginsii Webster, sp. nov. Annua erecta, caule dense gland-
uloso-tomentello; foliis ovatis basi cordatis obtuse dentatis 2—7 cm longis,

262 MADRONO [ Vol. 20

strigoso-hirsutis glandulosisve; spicis axillaribus, parte mascula 1.5—10
mm longa; calycis masculis glabris; involucris foemineis 2—3, 2—floris, ad
mediam 6—10-lobis, glandulosis, parce hirsutis; seminibus 1.3-1.5 mm
longis.

Type. Isla Santa Cruz (Indefatigable), north slope of Mt. Crocker, alt.
ca. 860 m, Wiggins & Porter 663, Feb. 18, 1967 (DS-holotype). Addi-
tional collections examined, Mt. Crocker, Wiggins & Porter 665 (DS);
Gebirge im Innern, Schimpff 96 (CAS, MO); among rocks in moist
zone, south slope of mountain, alt. 1000 ft., Svenson 96 (BRKL).

This species is known only from Isla Santa Cruz, where it appears
to be restricted to higher altitudes in the mountainous interior. The
completely glabrous staminate calyces and large stipules distinguish it
from the many taxa of the 4. parvula complex, while the glandular pu-
bescence sets it apart from the large-leaved species related to A. sericea.

ACALYPHA PARVULA Hook. f., Trans. Linn. Soc. London, 20:185. 1847.

All of the small-leaved populations of Galapagos Acalypha with gland-
lar pubescence are referred to this single protean species. As Robinson
noted, these plants include annuals and perrenials, erect and prostrate
forms, with striking differences in the spikes. His decision to recognize
nine species in this complex cannot be followed, because these various
marked characters vary in a largely uncorrelated manner from island
to island without showing any clear geographic separations. A number
of the taxa may be treated as varieties, in approximately the circum-
scriptions of Mueller, but it must be admitted that these are difficult to
distinguish in practice and may not be natural units. The four varieties
recognized here are extensively sympatric, and further study may show
that they are simply arbitrary assemblages of plants selected from a
mosaic of clinal and microgeographic variation.

ACALYPHA PARVULA Hook. f. var PARVULA. Type, Isla Isabela (Albe-
marle), Macrae (K-n. v.). A. cordifolia Hook. f., Trans. Linn. Soc. Lon-
don 20:187. Type, Isla Santa Maria (Charles), Darwin (CGE). A. dif-
fusa Anderss., Kong]. Vetensk. Acad. Handl. 1853:240. 1855. Type, Isla
Isabela (Albemarle), ‘in locis siccissimis’, Andersson (GH-isotype). A.
spicata Anderss., Kong]. Vetensk. Acad. Handl. 1853:239. 1854. Type,
Isla San Cristobal (Chatham), Andersson (GH-isotype). A. parvula B
cordifolia Muell.-Arg., Linnaea 34:47. 1865; in D.C., Prodr. 15(2):877.
1866. A. albemarlensis Robins., Proc. Amer. Acad. Arts 38:163. 1902.
Type, Isla Isabela (Albemarle), Tagus Cove, alt. 1220 m, Snodgrass &
Heller 885 (GH-holotype).

This variety is the most widespread and diverse taxon of Galapagos
Acalypha. It is recorded from Fernandina, Isabela, Pinzon, San Crist6-
bal, Santa Cruz, Santa Fé, and Santa Maria. Only on Espanola and San
Salvador does it appear to be lacking from the larger central islands. Vari-
ous forms of var. parvula are difficult to separate from other varieties of

1970] WEBSTER: GALAPAGOS EUPHORBIACEAE 263

A. parvula, and sometimes even from specimens of A. sericea. However,
in the majority of instances the populations included here may be rec-
ognized by the axillary pistillate spikes, distinctly glandular erect stems,
seeds 1.1 mm long or less, and leaves scarcely exceeding 2 cm long.

ACALYPHA PARVULA var. RENIFORMIS (Hook. f.) Muell.-Arg., Linnaea
34:48. 1865: in DC., Prodr. 15 (2):878. 1866. A. reniformis Hook. f.,
Trans. Linn. Soc. London 20:187. 1847. Type, Isla Santa Maria
(Charles), Darwin (CGE). A. adamsii Robins., Proc. Amer. Acad. Arts
38:161. 1902. Type, Isla San Cristobal (Chatham), southwest end, mid-
dle region, Baur 282, June 1891 (GH).

Stems often prostrate, usually not densely glandular; terminal spikes
usually present; pistillate bracts mostly subsessile; seeds small (0.9-1.1
mm long). Recorded from Espanola, Pinzon, Rabida, San Cristobal, San
Salvador, Santa Cruz, Santa Fé, and Santa Maria.

ACALYPHA PARVULA Hook. f. var. STROBILIFERA (Hook. f.) Muell.-
Arg., Linnaea 34:47. 1865; in DC., Prodr. 15(2):877. 1866. A. strobili-
fera Hook. f., Trans. Linn. Soc. London 20:187. 1847. Type, Isla San Cris-
tobal (Chatham), Darwin (CGE). A. parvula a procumbens Muell.-Arg.,
Linnaea 34:48. 1865; in DC., Prodr. 15(2):878. 1866.

Stems usually glandular, spikes terminal or axillary; pistillate bracts
usually pedunculate; seeds mostly large (1.2—1.5 mm long). Recorded
from Daphne, San Cristobal and Santa Cruz, common in the vicinity of
Academy Bay on the latter island.

ACALYPHA PARVULA Hook. f. var. chathamensis ( Robins.) Webster,
comb. nov. A. chathamensis Robins., Proc. Amer. Acad. Arts 38:163.
1902. Type, Isla San Cristobal (Chatham), Snodgrass & Heller 541
(GH-lectotype, DS-isotype).

Stems erect, glandular; leaves larger (2.5—4.5 cm long) than in most
other forms of A. parvula; spikes axillary; seeds small (1.0-1.1 mm
long). Endemic to San Cristébal, where it is known from only one other
collection (Snodgrass & Heller 540, DS, GH). In aspect, specimens of
var. chathamensis suggest A. sericea var. baurii, which also occurs on
San Cristobal; but the glandular and non-velutinous indumentum defi-
nitely places them within A. parvula.

I wish to thank the curators of the herbaria at Stanford, Harvard, the
Brooklyn Botanical Garden, and Cambridge for their generosity in loan-
ing critical specimens. The Darwin collections from the herbarium at
Cambridge (CGE) are duplicates of those at Kew examined by Hooker:
since the specimens at Kew were not seen, no attempt was made to
specify whether the collections at CGE are syntypes or isotypes.

Department of Botany, University of California, Davis

NOMENCLATURAL CHANGES AND NEW SUBSPECIES IN
THE CENTROSPERMAE OF THE GALAPAGOS ISLANDS

UNo ELIASSON

ALTERNANTHERA FILIFOLIA (Hook. f.) Howell ssp. glaucescens
(Hook. f.) Eliasson, comb. nov. Bucholtzia glaucescens Hook. f., Trans.
Linn. Soc. London 20:191. 1847. Telanthera glaucescens Mog. in D.C.,
Prodr. 13:369. 1849. T. strictiuscula Anderss., Kongl. Vetensk. Acad.
Handl. 1853:166. 1855. T. angustata Anderss., Kongl. Svenska Veten-
skapsakad. Handl. 1857:61. 1861. Achvranthes glaucescens Standley,
J. Wash. Acad. Sci. 5:74. 1915. A. strictiuscula Standley, J. Wash. Acad.
Sci. 5:75. 1915. Alternanthera glaucescens Howell, Proc. Calif. Acad. Sci.
IV. 21:104. 1933. A. glaucescens f{. strictiuscula Howell, Proc. Calif.
mead Ser lV 2 05.81933,

This subspecies apparently is restricted to Isla San Cristobal.

ALTERNANTHERA FILIFOLIA (Hook. f.) Howell ssp. microcephala
Eliasson, ssp. nov. A. glaucescens f. strictiuscula sensu Eliasson, Svensk.
Bot. Tidskr. 60: 143. 1966, not Howell, 1933.

Caules non multos ramos dimittentes, tenues, nonnumquam sub-
glauci; internodia 5—8 cm longa; folia inferiora linearia vel lineariolan-
ceolata, 3-4 cm longa, 2—7 mm lata, glabra; folia superiora filiformia
vel linearia, 2—4 cm longa, 0.5—-1 mm lata, glabra; capitula superior parte
plantae pauca, terminalia vel in brevibus pedunculis axillaribus posita,
rotundata, diametro 3—4 mm; flores laxe imbricati, apicibus rectis.

Holotype. Bahia Sullivan, 170 m, Isla San Salvador, Ecuador, /nga
cy Uno Eliasson 1393 (S).

Apparently restricted to Isla Bartolomé and to the adjacent areas
of Isla San Salvador.

ALTERNANTHERA FILIFOLIA (Hook. f.) Howell ssp. nudicaulis (Hook.
f.) Eliasson, comb. nov. Bucholtzia nudicaulis Hook. f., Trans. Linn.
Soc. Lond. 20:191. 1847. Telanthera nudicaulis Moq. in DC., Prodr.
13:369. 1849. Achyranthes nudicaulis Standley, J. Wash. Acad. Sci. 5:74.
1915. Alternanthera nudicaulis Christoph., Nyt Mag. Naturvidensk. 70:
(ice Roles

Known only from Isla Santa Maria.

ALTERNANTHERA FILIFOLIA (Hook. f.) Howell ssp. pintensis Eliasson,
ssp. nov.

Caules admodum ramosi, superiore saltem parte trichomatibus albis
subramosis praediti; internodia 3-5 (—7) cm longa; folia oblanceolata
vel oblongo-lanceolata, 2—3 (—6) cm longa, 4-7 (—15) mm lata, praeser-
tim infra trichomatibus albis stellatis praedita; capitula solitaria vel
2—7—glomerata, ovoidea vel cylindrica, 5-8 mm longa, 3-4 mm lata;

264

1970] ELIASSON: GALAPAGOS CENTROSPERMAE 265

flores admodum dense imbricati sed apicibus liberis rectis; flores tricho-
matibus densis subflavis simplicibus praediti.

Intermediate in characters between A. filifolia ssp. nudicaulis and A.
snodgrassit.

Holotype. At an altitude of about 200 m, Isla Pinta, Galapagos Islands,
Ecuador, /nga & Uno Eliasson 2151 (s).

Known only from Isla Pinta.

ALTERNANTHERA FILIFOLIA (Hook. f.) Howell ssp. rabidensis Fli-
asson, ssp. nov.

Caules et rami superiore parte trichomatibus simplicibus albis praediti;
internodia 2—5 cm longa; folia oblanceolata, 2—3 cm longa, 4-7 (—10)
mm lata, praesertim infra trichomatibus simplicibus albis praedita; capi-
tula solitaria vel 2-4—-glomerata, rotundata vel cylindrica, 4-10 mm
longa, 3-4 mm lata; flores laxe imbricati apicibus leviter incurvatis;
flores trichomatibus subflavis simplicibus praediti.

Holotype. On top of island, at an altitude of about 390 m. Isla Rabida,
Ecuador, /nga & Uno Eliasson 1414 (S).

Known only from Isla Rabida.

FROELICHIA NUDICAULIS Hook. f. ssp. lanigera (Anderss.) Eliasson,
comb. nov. F. lanigera Anderss., Kongl. Svenska Vetenskapsakad. Hand.
1857:63. 1861. F. lanata Anderss., Kongl. Svenska Vetenskapsakad.
Handl. 1857: pl. 3, fi. 1. 1861. F. scoparia Robins., Proc. Amer. Acad.
Arts 38:136. 1902. F. lanigera ssp. scoparia Howell, Proc. Calif. Acad.
Sen Wye Ze Ge 1938)

Occurring on fresh lava fields, especially at high altiudtes, on Islands
Fernandina and Isabela.

MOLLUGO FLAVESCENS Anderss. ssp. gracillima (Anderss.) Eliasson,
comb. nov. M. gracillima Anderss., Kong]. Vetensk. Akad. Handl. 1853:
226. 1855. M. gracilis Anderss., Kongl. Svenska Vetenskapsakad. Handl.
1857: pl. 15, fig. 3. 1861. M. gracillima Anderss. ssp. latifolia Howell,
Proc. Calif. Acad. Sci. IV. 21:17. 1933. M. flavescens Anderss. ssp. angus-
tifolia Howell, Proc. Calif. Acad. Sci. IV. 21:18. 1933. M. flavescens
Anderss. ssp. intermedia Howell, Proc. Calif. Acad. Sci. IV. 21.18. 1933.

MOLLUGO FLAVESCENS Anderss. ssp. insularis (Howell) Eliasson,
comb. nov. M. insularis Howell, Proc. Calif. Acad. Sci. IV. 21:19. 1933.
Known only from Islas San Crist6bal and Santa Maria.

MOLLUGO FLAVESCENS Anderss. ssp. striata (Howell) Eliasson, comb.
nov. M. striata Howell, Proc. Calif. Acad. Sci. IV. 21:19. 1933.

Known only from the type collection (Stewart 1477, CAS), from Isla
Wolf.

266 MADRONO [ Vol. 20

MOLLUGO FLORIANA (Robins.) Howell ssp. santacruziana (Chris-
toph.) Eliasson, comb. nov. M. snodgrassi Robins. var. santacruciana
Christoph., Nyt Mag. Naturvidensk. 70:75. 1931.

Known only from the type collection, taken at “Academy Bay,” Isla
Santa Cruz (Rorud 1230).

ANREDERA ramosa (Mogq.) Eliasson, comb. nov. Tandonia ramosa
Mogq., in D.C. Prodr. 13:227. 1849. Boussingaultia ramosa Hemsley,
Biol. Centr.-Amer. 3:27. 1882. B. baselloides in Galapagos literature,
not H.B.K., 1825.

University of Goteborg, Sweden

A NEW SPECIES OF POLYGONUM (POLYGONACEAE)
JERROLD COOLIDGE

During the course of study in preparing a revision Section Avicularia
of the genus Polygonum, (Coolidge, 1964) a group of specimens were
found which possessed charateristics distinct enough to warrant descrip-
tion as a new species.

Polygonum triandrous Coolidge, sp. nov. Herba annualis; erecta; 1
vel 4 dm longa; folia lineari-lanceolata vel linearia, 2 vel 4 cm longa;
inflorenscentia axillaris; 2 vel 3 flores in axilla foliorum; flores erectae, 2
vel 3.5 cm longae, virides marfinibus roseis vel albis; stamina 3; ache-
nium ovoideum, 3 mm longum, atrum, glabrum.

Annual; erect, glabrous except glaucescent or scurfy at nodes; stem
1 to 4 dm long, terete, branched from the base or throughout; leaves
linear-lanceolate or linear, 2 to 4 cm long, 3 to 5 mm wide, revolute or
flat, light green, glaucous on upper surface, acute, midvein prominent,
articulation to ocrea conspicuous; ocrea 2-parted when young, lacerate
with age, silvery with reddish-brown base, 4 to 5 mm long; pedicels
stout, 2 mm long; inflorescence axillary, 2 to 3 flowers in the axils
throughout the length of stem; articulated with the flower at the flower
base, flower and fruit erect, 3 to 3.5 mm long, green with pink or white
margins; calyx segments ovate, obtuse, 5 -parted to near the base;
stamens 3, 1.5 mm long, anthers white, filaments dilated gradually; style
0.1 to 0.2 mm long, 3-cleft to near the base; achene ovoid, 3 mm long,
black, smooth and shining (fig. 1).

Type. Idaho: Blaine Co., along trail to Hyndman Peak, W. H. Baker
11005 (ID-holotype, ARIZ, MONTU, NY, OSC, RM, UC, WS, WTU).

Numerous collections of P. triandrous from herbaria throughout the
western states have been examined. In most cases, it has been identified
as P. sawatchense Small and does appear to be most closely allied with

1970] COOLIDGE: POLYGONUM

i)
a

Fic. 1. Photograph of type of Polygonum triandrous.

this species. It differs in having narrower leaves below, the upper leaves
of a more bracteate nature and the most apparent difference, the pres-
ence of only three stamens. Polygonum triandrous may be distinguished
from other related species by the following key:

268 MADRONO LVol. 20

Fic. 2. Distribution of Polygonum triandrous in western North America.

Leaf size somewhat reduced upward, lanceolate, liner
Leaves linear, linear-lanceolate

Stamens3 . . . . . . . P. triandrous Coolidge

Stamens S$ = << .« . | «= & .). | P.tenve Niche

Leaves lanceolate . . . . . . .P. sawatchense Small
Leaf size about the same throughout, ovate, obovate

Flowers fewinaxils,remote . . . . . P. minimum Wats.

Flowers several in axils, congested . . . P.cascadense Baker

The known range includes areas within the Colorado Plateau, New
Mexican Highlands, Rocgy Mountains, Columbia Plateau, Great Basin
and the Sierra Nevada (fig. 2). The distributional pattern of this species

1970] MOSQUIN: SISYRINCHIUM 269

would indicate a probable origin in central Arizona, radiating into all
western states except perhaps, Washington and Montana.

Department of Botany, University of Idaho, Moscow

LITERATURE CITED
Coorince, J. O. 1964. A revision of the genus Polygonum section Avicularia in the
western United States. M. A. Thesis (unpublished), University of Idaho.

CHROMOSOME NUMBERS AND A PROPOSAL FOR
CLASSIFICATION IN SISYRINCHIUM (IRIDACEAE)

THEODORE MOSQUIN
INTRODUCTION

The classification of North American Sisvrinchium has been highly
unsatisfactory for many decades. For example, it is difficult with the
aid of standard floras such as Abrams (1923), Fernald (1950), Gleason
(1952), and Munz (1959) to identify many collections of this genus
from regions covered by these floras. One reason for this difficulty is
that plants of this genus are notoriously lacking in qualitative differ-
ences such as are necessary to distinguish species. A second reason is
that regional floras which deal with different or confluent parts of the
continent have continued to follow traditional and local classifications.
The existing disagreements concerning the occurence and the nature
of phenotypic discontinuities illustrates the need for a reappraisal of
variability not only by the use of techniques of modern taxonomy but
by examination of variation on a continent-wide basis. The present
paper brings together some field, herbarium, published, and laboratory
observations on correlations between morphology, chromosome num-
bers, ecology, and geography of plants of this genus from populations
throughout much of the North American distribution area.

The Sisvrinchium populations considered in this paper comprise the
widely distributed, small-flowered perennials in which the anther fila-
ments are united in a tube. Excluded are the annuals of Texas and
adjacent regions which have been discussed recently by Shinners (1962).
Also excluded are the large-flowered perennials of Mexico and the Carib-
bean Islands as well as the large-flowered and very distinctive S. doug-
lasii Dietr. of western North America. So defined, the plants commented
on in this paper range from Greenland to Alaska and south to Florida,
Texas, and California and may extend into Mexico.

Directly relevant to their classification and an outstanding feature
of the Szsvrinchium populations considered here is that a high degree
of self-pollination appears to be a characteristic feature (Knuth, 1909;
Ingram, 1967; Table 1). Table 1 shows the results of an experiment de-
signed to determine the potential for automatic self-pollination in a
tetraploid poupulation growing under natural conditions near Banff,

270 MADRONO [Vol. 20

TABLE 1, SEED SET RESULTING FRoM ARTIFICIAL BAGGING EXPERIMENTS
IN WILD PLANTS OF SISYRINCHIUM BERMUDIANA

Data from a 12-ploid population about 6 miles west of Banff, Alberta, Canada
(vouchers Mosquin & Seaborn 7048 and 7164, DAO).

Number of Number of capsules Number of Number of seeds in
individual on inflorescence seeds per control capsules
plant capsule (adjacent plants)

7048-1 3 35 2028 16
7048-2 2 33 16 38
7048-3 2 63:6 20
7048-4 1 32 30
7048-5 S Sip ae) 24
7048-6 1 1 28

Alberta. The pollination bags used in this experiment were white and
highly porous. They were made from synthetic material especially de-
signed for tree-breeding work in the field and do not deteriorate under
usual field conditions. The plants were bagged on June 25 and the data
collected on July 29, 1968. The bagged plants of this population, iso-
lated from insects, produced nearly 45% seed set as compared with the
controls. There can be no doubt, therefore, not only that these plants
were self-compatible but that they are highly homozygous. A consid-
eration of the morphological effects of self-pollination (Stebbins, 1957)
reads like a description of the variation pattern of wild populations
considered here. Thus local populations are highly uniform morpho-
logically, while conspicuous differences often occur between geograph-
ically isolated colonies. Intermediates are common between character
extremes. Morphologically very similar plants occur in widely separated
places, for example on the American northwest coast and in the Appa-
lachian region, and again in Greenland and the western United States.

Another feature of the Sisyrinchium populations considered here is
polyploidy, an important evolutionary mechanism. To date the num-
bers 2n — 16, 32, 64, 82, 84, 88, 90, and 96 have been reported among
these plants (Bowden, 1945; Love and Love, 1958; 1961; Lewis and
Oliver, 1961; Oliver and Lewis, 1962; Clapman, et al.; 1962; Bocher,
1966; Oliver, 1966; Ingram, 1967; table 2). It is possible, therefore,
that in some places chromosome numbers might provide the necessary
key for discovering corresponding morphological breaks.

In closely related species, differences in chromosome numbers are
frequently reflected in differences in pollen grain size. This is, however,
not so in Szsyrinchium, where I have found that tetraploids may have
pollen as large as the 12-ploids. It is of interest that Bocher (1966) in

1970] MOSQUIN: SISYRINCHIUM 7A

TABLE 2. CHROMOSOME NUMBERS.

Sisyrinchium arenicola Bicknell. 2n = 32: NORTH CAROLINA, Moore Co., 2.7
mi S of Pine Bluff, Mosquin & Mosquin 5935 (COLO, DAO).

S. bellum Wats. n = 16: CALIFORNIA, San Diego Co., Santa Yysabel, Mosquin
& Snow 3974 (COLO,, DAO); several mi W of San Pasqual, Mosquin & Snow
3975 (COLO, DAO). 2n = ca 90 CALIFORNIA, Inyo Co., 6.7 mi W of the Lee
Vining Junction along Tioga Pass Hwy, Mosquin 4780 (COLO, DAO, DS, UAC).

S. bermudiana L, 2n = 32: ALBERTA, about 6 miles west of Banff, Mosquin
& Seaborn 7164. CALIFORNIA, Plumas Co., about 2 mi SE of Graegle, Mosquin
& Gillett 5305 (COLO, DAO). n = 16: NEVADA, Lyon Co., road to Virginia City
near Hwy 50, Gillett & Moulds 12700 (COLO, DAO). 2n = ca 64: MONTANA,
Meagher Co., Ringling, Mosquin & Gillett 5226 (DAO); Missoula Co., 13.9 mi W
of Lolo, Mosquin & Gillett 5262 (DAO). TEXAS, Van Zandt Co., 12.7 mi E of
Terrell, Mosquin & Mosquin 5469 (DAO, DS); Galveston Co., Galveston Island,
near SW tip of the Island, about 22 mi SW of Galveston, Mosquin & Mosquin
5529 (DAO). 2n = 96: ALBERTA, 7.5 mi S of the Trans Canada Hwy, along road
to Kananaskis Lakes, Mosquin & Benn 5185 (COLO, DAO, UAC). BRITISH
COLUMBIA, 7.7 mi E of Galloway (no voucher). CALIFORNIA, Sierra Co.,
along road from State Hwy 89 to Independence Lake (DAO). MONTANA, Cas-
cade Co., 15 mi E of Great Falls, Mosquin & Gillett 5215 (DAO); Jefferson Co.,
1.5 mi E of Pipestone Pass, Mosquin & Gillett 5261 (COLO, DAO, UAC). NEW
BRUNSWICK, Restigouche Co., 20 mi NE of Kedgwick, Mosquin & Spicer 6374
(DAO). ONTARIO, Norfolk Co., Turkey Point, Bowden 138—55 (DAO); Carleton
Co., 3 mi SW of North Gower, Mosquin & Frankton 6519 (DAO). QUEBEC, Bon-
aventure Co., 1 mi W of Nouvelle, Mosquin & Spicer 6366 (DAO); Gaspe Co., at
lodge about 2 mi N of Mt. Albert, Mosquin & Spicer 5997 (DAO); 9 mi W ot
Petite Vallee, Mosquin & Spicer 6346 (COLO, DAO); Rimouski Co., about 6 mi
SW of Ste. Flavia, Mosquin & Spicer 5997 (COLO, DAO); Stanstead Co., about
5 mi due E of Fitch Bay, Mosquin & Spicer 6338 (DAO). SOUTH DAKOTA,
Pennington Co., 1.9 mi NE of Hill City, Mosquin & Mulligan 5148 (COLO, DAO).

comparing tetraploids and 12-ploids found that seed diameter in the
former was 0.7-0.8 mm, while in the latter it ranged from 1.1 to 1.4 mm,
but the sample size was small. Seed diameter in the 12-ploid plants
from Banff referred to in Table 1 ranged from 0.7 to 1.1 mm.

Yet another feature of the genus considered here, and one that has
contributed to the creation of many names of dubious value, is the
wide ecological diversity of the wild populations. This diversity is evi-
dent from the habitat descriptions in various floras and from first-hand
field observations. Wild populations occur in montane meadows, sage
deserts, prairies, and seashores, and often along roads as weeds.

TAXONOMY

Among the many species names currently in use for the American
populations of Sisyrinchium, the most widely employed are perhaps
S. angustifolium Mill. and S. montanum Greene. The former is applied
to eastern North American populations from eastern Canada to Florida
and Texas (Fernald, 1950; Shinners, 1963); the latter most often to
western populations from the Northwest Territories and British Colum-

272 MADRONO [ Vol. 20

bia south to Colorado, but also eastward to New York, eastern Canada,
and Greenland (Fernald, 1950; Bocher, 1966). The most important
character by which these two species are purported to differ is branch-
ing, S. angustifolium being branched and S. montanum unbranched. It
is very common, however, to find both branched and unbranched plants
in many populations, although one type is usually much more frequent
than the other. It would be much more useful, therefore, to consider
these two species to be conspecific, as Rydberg (1932) thought. An-
other widely used name is S. idahoense Bicknell. Yet plants of this
species from Idaho, Oregon, and California do not differ in any single
trait or combination of traits from the S. montanum—sS. angustifolium
populations considered above. Similarly some other names which are
currently used in floras appear to apply to populations which have
morphological traits well within the range of variability of the popu-
lations considered above. Thus species like S. campestre Bicknell, S.
albidum Raf., S. graminoides Bicknell (see Shinners, 1962, for dis-
cussion of this name), S. dangloisai Greene, S. sagittiferum Bicknell,
S. littorale Greene, S. sarmentosum Suksdorf, and S. halophilum Greene,
as far as I can judge from descriptions in floras and also from com-
parisons of herbarium specimens, are very likely best treated within
a single widespread species. The correct name for this widespread spe-
cies appears to be S. bermudiana L. (Shinners, 1962).

Several additional species have been described which, in contrast to
those mentioned above, are, at least in their morphological character-
istics, modally distinct from S. bermudiana. These are S. atlanticum
Bicknell, S. mucronatum Greene, S. arenicola Bicknell, S. capillare
Bicknell, and in the west perhaps S. bellum Wats., although many pop-
ulations of the last species, particularly from the Sierra Nevada of
California, would readily pass for collections from the eastern United
States (see Munz, 1959, for brief discussion of S. bellum.)

CORRELATIONS AND DISCUSSION

Correlations between morphological characters on the one hand and
ecology, chromosome number, distribution, and breeding habit on the
other provide the basis for contemplating the details of a classification
that not only would be readily usable but would closely reflect our
present knowledge of genetic relationship. Perhaps the most important
correlation between these characteristics in Sisyrinchium, and one that
is of vital significance to the classification of this genus, is a negative
one, namely, morphological differentiation is, for the most part, not
associated with chromosomal differentiation. Hence chromosome num-
bers will not play an important role in helping to construct a useful
classification for this genus.

Another fact is that ecological similarities or differences very fre-
quently are not accompanied by corresponding morphological patterns.
For example, a population from a marshy habitat in sage desert near

1970] MOSQUIN: SISYRINCHIUM 213

Virginia City, Nevada (Gillett & Moulds 12700, DAO), is virtually
identical in all features of external morphology with plants collected
on dry grassy stream banks in montane yellow pine forest of the Sierra
Nevada (Mosquin & Gillett 5307, DAO). In this example the former
colony is tetraploid while the latter is 12-ploid. In yet another case,
the octoploid populations of Montana do not appear to differ ecolog-
ically from the geographically adjacent 12-ploid colonies. Neither do
these two chromosome races in Montana differ morphologically in any
perceivable way. As a last example, plants of Greenland (Bocher, 1966)
are essentially identical with many populations in the western United
States; the Greenland plants are tetraploid, while at least three chro-
mosome races (tetraploid, octoploid, and 12-ploid) are found in the
western United States. Other examples could be cited. That such rela-
tionships may also occur in other species of Sisyrinchium is suggested
from reports of the numbers n = 8, 16, and 48 and 2n = 96 from S.
atlanticum Bicknell (Love and Love, 1958; Oliver and Lewis, 1962;
Oliver, 1966).

The geographical distribution of the chromosome races of S. ber-
mudiana is of interest. The tetraploids occur in southern portions of
the United States, in the Great Plains, in western Alberta, in California,
and, surprisingly, in Greenland. Octoploids are known only from Texas
and Montana, while 12-ploid populations are very widespread in the
northern regions and also occur in the Queen Charlotte Islands, Brit-
ish Columbia (Taylor and Mulligan, 1968). Figure 1 gives the distri-
bution of the chromsome races. Evolution by aneuploid decrease from
2n = 96 is clearly occurring in the eastern United States, with numbers
as low as 2n = 82 recorded (Ingram, 1967). The count of 2n = 90
(Bowden, 1945; and present paper) from the Sierra Nevada suggests
that an aneuploid reduction series from 2n = 96 may also be present
in the western United States.

The geographical origin of the Greenland tetraploids poses a special
problem. It would be useful to determine the chromosome number of
populations in Newfoundland and particularly in the vicinity of Goose
Bay, Labrador, since presumably the Greenland tetraploids originated
from some locality in northeastern North America.

Some additional reports of chromosome numbers in S. bermudiana
have been published by Love and Love (1958) under different species
names. These are given in their paper as having been determined from
“many plants from the southern and eastern parts of the province of
Quebec” and from “northern Virginia” (2n = 96 as S. angustifolium),
from “the Canadian prairies” and from ‘“‘a few places in Ontario and
Quebec” (2n = 32 as S. montanum), and from “a couple of localities
in southern Wisconsin and Ontario” (2n = 32 as S. albidum). These
report could not be included in Fig. 1 because the exact localities
and voucher information were not given in the paper. Love and Love
also report having determined the number 2n = 96 from plants of

274 MADRONO [ Vol. 20

LEGEND
@ TETRAPLOID (2n = 32)
MOCTOPLOID (2n= 64)
ATWELVE-PLOID (2n = 96)

Fic. 1. Distribution of Szsyrinchium bermudiana (dotted area) showing loca-
tions of populations with chromosome counts. Chromosome counts previously re-
ported under the names S. albidum (Bowden, 1945; Oliver and Lewis, 1962),
S. angustifolium (Bowden, 1945), S. campestre (Oliver and Lewis, 1962; Oliver,
1966), S. groenlandicum (Bocher, 1966), S. langloiszi (Oliver and Lewis, 1962),
and S. montanum (Ingram, 1967) are included. The counts for S. bermudizana
given by Ingram (1967) are also included.

S. bermudiana from “several localities in Britain and Scandinavia,” as
well as the number 2n = 64 from one Irish population which they de-
scribe as a new species (Love and Love, 1961). The correct number of
the Irish population has now been shown to be 2n = 88 (Ingram, 1967).
The morphology and chromosome number of the Irish plants fall well

1970] MOSQUIN: SISYRINCHIUM 218

into the range of variability of S. bermudiana even when grown under
similar conditions in the greenhouse (Ingram, 1967). In a detailed study
of the Ireland plants Ingram concluded that they should be most use-
fully treated as S. bermudiana. There seems little doubt that adopting
a wide species concept for the North American Sisyrinchiums would
provide biologists with a maximally useful classification.

ACKNOWLEDGMENTS

I am indebted to the Research Branch of the Canada Department of
Forestry for providing facilities for conducting field studies at Banff,
Alberta, where the pollination experiment on Sisyrinchium was carried
out. This paper is Contribution No. 605 of the Plant Research Institute.

Plant Research Institute, Canada Department of Agriculture, Ottawa

LITERATURE CITED

Aprams, L. 1923. An illustrated flora of the Pacific States. Vol. 1. Stanford Univ.
Press.

Bocuer, T. W. 1966. Experimental and cytological studies of plant species. X. Sisy-
rinchium with special reference to the Greenland representative. Bot. Tidsskr.
61:273-290.

Bowpen, W. M. 1945. A list of chromosome numbers in higher plants. I. Acantha-
ceae. Amer. J. Bot. 32:81-92.

CrapHaM, A. R., T. G. Turin, and E. F. Warpurc. 1962. Flora of the British Isles.
2nd edition. Cambridge Univ. Press.

FERNALD, M. L. 1950. Gray’s Manual of Botany. 8th edition. American Book Co.,
NGYe

Gurason, H. A. 1952. The new Britton and Brown illustrated flera of the North-
eastern United States and adjacent Canada. Vol. 1. New York Bot. Gard.

INGRAM, R. 1967. On the identity of the Irish populations of Sisyrinchium. Wat-
sonia 6:283-289.

KnutuH, P. 1909. Handbook of flower pollination. Vol. 3. Clarendon Press, Oxford.

Lewis, W. H., and R. L. Oriver. 1961. Meiotic chromosomes in six Texas and
Mexican Nemastylis and Sisyrinchium (Iridaceae). Southw. Naturalist 6:45—46.

Love, A., and D. Love. 1958. The American element in the flora of the British Isles.
Bot. Not. 111:376-388.

. 1961. Some nomenclatural changes in the European flora. Bot. Not.
114:33-47.

Muwnz, P. A. 1959. A California Flora. Univ. Calif. Press, Berkeley.

Oriver, R. L. 1966. Chromosome numbers of phanerogams I. Ann. Missouri Bot.
Gard. 53: 100-103.

.. and W. H. Lewis. 1962. Chremoscme numbers of Sisyrinchium (TIrida-
ceae) in eastern North America. Sida 1:43—48.

Rypperc, P. A. 1932. Flora of the prairies and plains of Central North America.
Scientific Press Printing Co., Pennsylvania.

SHINNERS, L. H. 1962. Annual Sisyrinchiums (Iridaceae) in the United States. Sida
1:32-42.

STEBBINS, G. L. 1957. Self-fertilization and population variability in the higher
plants. Amer. Naturalist 91:337-354.

Taytor, R. L., and G. A. Mutrican. 1968. Flora of the Queen Charlotte Islands,
Part 2. Cytological aspects of the vascular plants. Canad. Dept. of Agric.
Monogr. 4.

COMPARATIVE NATURAL HISTORY OF TWO SYMPATRIC
POPULATIONS OF PHOLISTOMA (HYDROPHYLLACEAE)

KAREN B. SEARCY

The generalization that similar species are allopatric has stood the test
of numerous systematic studies. It is usually assumed that it is compe-
tition which makes the ranges of these species mutually exclusive. Occa-
sionally populations of two similar species can be found growing together
at the same site. Such sympatric occurrences provide an ideal opportu-
nity to test the assumption and to look for influences of one population
on another. In addition, sympatric occurrences may help us recognize
features of each population which provide for their continued reproduc-
tive independence.

The object of the study was to investigate a site where two closely
related species of Pholistoma were found growing together. Constance
(1939) has shown by careful morphological study that P. racemosum
(Nutt.) Constance and P. auritum (Lindl.) Lilja. are far more similar
to each other than to any other taxa, and has segregated the two out of
the genus Nemophila and placed them in Pholistoma. The other taxa in
Pholistoma, P. membranaceum (Benth.) segregated out of the genus
Ellisia, and P. auritum var. arizonicum (M. E. Jones) were not consid-
ered in this study.

Pholistoma auritum var. auritum and P. racemosum are allopatric
throughout most of their range. Pholistoma auritum, which has larger
purple flowers, is found in the coast ranges of California from San Diego
north to Lake Co., in the Sierra Nevada foothills from Calaveras Co., to
Kern Co., and on Santa Catalina, San Clemente, and Santa Cruz islands.
Pholistoma racemosum, with small white flowers, is found in northern
Baja California, San Diego Co., and on the off-shore islands of Baja
California and California. The one know site where they do occur to-
gether (Raven, 1963) is at the base of a large rock outcrop in Little
Sycamore Canyon near the west end of the Santa Monica Mountains,
Ventura Co. The site is moist, and is characterized by morning and eve-
ning fog tending to make a more equable climate than other such wood-
lands in the Santa Monica Mountains.

The two species differ greatly in their breeding systems. Greenhouse
culture showed P. racemosum to be autogamous while P. auritum is self-
compatible but does not set any seed unless artificially self-pollinated.
Pholistoma auritum is kept from self-pollinating by two mechanisms:
protandry, and the fact that the inflorescence rarely has more than one
flower shedding pollen at a time. The next flower on the inflorescence
usually does not open until 1-3 days after the previous flower has
opened. Although the flowers of P. auritum have no perceptible odor, they
have 5 yellow nectaries which were conspicuous when the flower first
opened but faded within one day.

276

1970} SEARCY: PHOLISTOMA Od

Field studies confirmed that P. auritum was predominantly outcrossed
and P. racemosum inbred. Pholistoma auritum was regularly visited by
pollinators, usually between 0900 and 1100, which was shortly after the
flowers opened and while pollen was being shed. The bees removed al-
most all of the pollen before noon, at which time the stigmas elongated
and became receptive. Observation of pollinators was made during the
time of maximum flowering from 0700 to 1400 on 16 April 1966 and
from 1630 to 1800 on 13 April 1966. No pollinators were observed visit-
ing P. racemosum during the time of maximum flowering. Observations
were made between 1300 and 1900 on 30 March 1966 and 0530 to 1300
on 2 April 1966. In addition, P. racemosum lacked any regular time of
opening, although it tended to open in the late afternoon.

No hybrids were found between P. auritum and P. racemosum al-
though individuals of the two species were growing just a few feet from
each other. The absence of hybrids may be partly due to a difference in
flowering time. Pholistoma racemosum bloomed earlier than P. auritum
during 1966. The first flower of P. racemosum opened about 15 March
1966; maximum flowering was two weeks later, capsules were formed
and the plants were beginning to dry about 16 April 1966. Pholistoma
auritum opened for the first time on 2 April 1966; maximum flowering
occurred two weeks later, the capsules matured and the plants were
dying by 11 May 1966.

The two species also differed in microhabitat. Pholistoma racemosum
occupied areas receiving much less direct sunlight than the areas in
which P. auritum was found. The densest clusters of P. racemosum oc-
curred on the N slope of the rock outcrop where it received direct light
for only about 20 minutes each day. The next densest clusters were on
the N sides of rocks and trees on the NW slope of the rock outcrop, and
received about 2 hours of sunlight during each day. In contrast, P. auri-
tum was found on the SW slopes leading away from the rock outcrop,
and received direct sun most of the day. Early in the year, P. auritum
also occurred on the NW slopes, but almost all of these plants died be-
fore flowering. In addition, the sites occupied by P. racemosum were
characterized by shallow soil, little leaf litter and few other plants,
whereas sites occupied by P. auritum were characterized by deep soil,
considerable litter, and much other vegetation.

An examination of herbarium specimens of P. racemosum and P.
auritum var. auritum showed that the plants in the study area do not
differ morphologically from those in other parts of the ranges. Flowering
dates also seem to be the same and the populations do not differ chromo-
somally. Cave and Constance (1942:1957;1959) have reported both
species to be n= 9. The chromosome numbers of plants from both
species from the study site are also n = 9. (P. auritum, Bartholomew
023, LA; P.racemosum, Bartholomew 015, LA).

A study of the two sympatric populations of Pholistoma has led me

278 MADRONO [ Vol. 20

to the following conclusions. The two species growing together in Little
Sycamore Canyon occupy sufficiently different microhabitats so that
they are not in direct competition. Their breeding systems are sufficient-
ly different so that they do not even compete for the same pollinator.
The two sympatric populations show no differences from other popula-
tions of their species, so that neither character displacement nor intro-
gression have occurred. Thus, the normally allopatric distribution cannot
be attributed to competitive exclusion, rather the populations must be
though of as ecologically quite different, and it is this difference which
causes their difference in geographical distribution.

Department of Botanical Sciences, University of California, Los Angeles

LITERATURE CITED

Cave, M., and L. Constance. 1942. Chromosome numbers in the Hydrophyllaceae.
Univ. Calif. Publ. Bot. 18: 205-216; 1947. III. 18:449-465; 1959. V. 30:233-
257:

ConstTANCE, L., 1939. The genus Phclistoma Lilja. Bull. Torrey Club 66:341-352.

RAVEN, P. H., 1963. A flora of San Clemente Island, California. Aliso 5:289-347.

A PRELIMINARY REPORT OF THE MYXOMYCETES OF
CRATER LAKE NATIONAL PARK, OREGON

DWAYNE H. CurtTIs

Crater Lake National Park is located in southeastern Oregon where
volcanic activity and glaciers shaped the surrounding mountains and
valleys. During the winter months the park is noted for an abundance
of snow accumulation, often exceeding 50 feet of measured depth from
November to May. The average annual precipitation is about 70 inches.
In contrast, the summer is quite dry since very little rain falls during
the months of July and August. The flora in the park must withstand
the extreme weather conditions in order to survive. It is common to
observe living trees bent from the shifting snow. In many places the
forest floor is covered with broken limbs and fallen trees.

Slime molds or Myxomycetes are characteristically associated with
moist areas on decaying organic matter such as duff, wood, bark, and
fallen twigs. An ideal habitat for slime molds is formed on the fallen
logs and forest litter dampened by the large amount of water from the
melting snow.

The collections for this report were obtained during the summers of
1966 and 1967. The 43 species, listed here, were collected in the field
on some form of decaying wood or duff at altitudes from 4,000 to 7,500
feet. At least one collection of each species has been deposited in the
University of Iowa Herbarium, Iowa City, Iowa and where possible, du-

1970] CURTIS: MYXOMYCETES 279

plicate specimens have been given to the Crater Lake National Park
Herbarium, Crater Lake, Oregon. The numbers given for the collections
are my own and they indicate only those specimens given to the Univer-
sity of lowa Herbarium.

(CERATIOMYXACEAE

Ceratiomyxa fructiculosa (Mull.) Macbr. On decayed wood, Kerr Val-
ley, 6,500 feet, 45, July 28, 1966.

LICEACEAE

Lisea minima Fries. One collection on decayed wood, Sleepy Hollow
Creek area, 6,500 feet, 16, June 18, 1966.

L. pusilla Schrad. One collection on decayed coniferous wood, 0.2 miles
north of Park Headquarters, 6,500 feet, 6, June 15, 1966. This exceed-
ingly tiny species was recently reported by Kowalski (1966) from Cali-
fornia. Previously, it had only been found as far west as Iowa, and is
considered rare.

RETICULARIACEAE

Lycogala epidendrum (L.) Fries. On decayed wood, Grouse Hill, 7,000
feet, 632, Jume 29, 1967.

L. flavofuscum (Ehrenb.) Rost. I obtained only one aethalium on the
side of a dead, barkless stump about 4 feet above the ground on the
east side of Kerr Valley, 6,500 feet, 52, July 28, 1966.

Enteridium olivaceum Ehrenb. On decayed wood, Vidae Falls area,
6,500 feet, 859, July 2, 1967.

(CCRIBRARIACEAE

Cribraria argillacea (Pers.) Pers. Inside a decaying log, 2 miles south
of Park Headquarters, 6,400 feet, 1007, July 8, 1967.

C. rufa (Roth) Rost. On decayed wood, Kerr Valley, 6,800 feet,
1130, July 29, 1967.

Lindbladia effusa (Ehrenb.) Rost. One large collection on the side of
a decaying log, 0.5 miles west of Rim Village, 6,800 feet, 7083, July 16,
1967.

DIANEMACEAE

Dianema andersoni Morgan. One collection on decayed wood, about
2 miles north of Park Headquarters, 6,800 feet, 7067, July 14, 1967.
This Myxomycete has been reported from Washington, British Colum-
bia, and more recently from California (Kowalski and Curtis, 1968). It
is considered rare.

D. corticatum Lister. On decayed wood buried in duff, 2 miles north
of Park Headquarters, 6,800 feet, 1052, July 14, 1967. Numerous col-
lections of this slime mold were obtained at elevations from 5,000 to
7,000 feet.

Prototrichia metallica ( Berk.) Massee. On decayed wood, Grouse Hill,

280 MADRONO [ Vol. 20

7,000 feet, 829, June 29, 1967. This Myxomycete is very common
throughout the park.
TRICHIACEAE

Arcyria versicolor Phill. On bark, 0.5 miles west of Rim Village, 6,800
feet, 1114, July 18, 1967. Several specimens were collected at elevations
from 6,000 to 7,000 feet.

Hemitrichia karstenii (Rost.) Lister. On the bark of a small fallen
twig, 4 miles north of the south park boundry, 5,000 feet, 779, June 17,
1967.

H. montana (Morgan) Macbr. On decayed wood, 2 miles south of
Park Headquarters on Munson Ridge, 6,500 feet, 67, August 4, 1966.
This Myxomycete is very common in the park at elevations from 6,000
to 7,000 feet.

Oligonema schweinitzu (Berk.) G. W. Martin. On decayed wood,
Goodbye Creek area, 6,000 feet, 748, June 14, 1967.

Trichia affinis De Bary. On decayed wood, 1 mile northeast of Park
Headquarters, 6,650 feet, 45, July 18, 1966.

T. contorta (Ditmar) Rost. On fallen twigs, 0.5 miles west of Rim
Village, 6,800 feet, 1077, July 16, 1967.

T. favoginea (Batsch) Pers. On decayed wood, Lightning Springs area,
6,800 feet, 77, August 16, 1966.

T. lutescens Lister. One collection on duff, Sleepy Hollow Springs area,
6,600 feet, 794, June 25, 1967.

T. pusilla (Hedw.) G. W. Martin. On decayed wood, Sleepy Hollow
Springs area, 6,500 feet, 570, June 28, 1967.

T. varia (Pers.) Pers. One collection on decayed wood, Annie Creek
near park boundary, 4,400 feet, 20, June 18, 1966.

STEMONITACEAE

Barbevella minutissima Meylan. One collection of three sporangia on
decayed wood, 0.5 miles south of Park Headquarters on Munson Ridge,
6,600 feet, 782, June 22, 1967. I (Curtis, 1968) recently reported the
occurrence of this exceedingly rare Myxomycete from the park. Previ-
ously, it had only been reported from Switzerland, Poland, and Japan.

C. fusiforme Kowalski. On decayed wood, 1 mile south of Vidae Falls,
6,700 feet, 883, July 3, 1967. This Myxomycete is very common in the
park at elevations from 5,000 to 7,000 feet.

C. nigra (Pers.) Schroet. On decayed wood, Grouse Hill, 7,000 feet,
830, June 29, 1967.

C. pacifica (Macbr.) Peck & Gilbert. On bark and twigs, 0.3 miles
kest of Goodbye Creek and the Park Road, 6,000 feet, 897, June 15,
1967.

C. suksdorfu Ellis & Ev. On bark, Munson Point, 7,000 feet, 757, July
4, 1967. Numerous collections were obtained throughout the park at
elevations from 5,700 to 7,100 feet.

1970] CURTIS: MYXOMYCETES 281

C. tyvphoides (Bull.) Rost. One collection on decayed wood, Goodbye
Springs area, 6,300 feet, 73, September 7, 1966.

Enerthenema melanospermum Macbr. & Mart. On decayed wood,
Grouse Hill, 7,000 feet, 848, June 29, 1967.

Lam proderma arcyrioides (Sommerf.) Rost. On decayed twigs, Sleepy
Holly Springs area, 6,600 feet, 7075, July 9, 1967. A slime mold com-
monly found throughout the park.

L. bias perosporum Kowalski. One collection on the side of a decaying
log, 0.5 miles west of Rim Village, 6,800 feet, 7068, July 14, 1967. This
rare Myxomycete was recently described by Kowalski (1968). It has
only been reported from California, Kentucky, and Oregon.

L. carestiae (Ces. & De-Not.) Meylan. On decayed twigs, Grouse Hill,
7,000 feet, 842, June 29, 1967.

L. sautert Rost. On bark, Sleepy Hollow Springs area, 6,600 feet,
S04, June 25, 1967.

PHYSARACEAE

Fuligo septica (L.) Webber. On a decaying log, White Horse Creek
area, 5,700 feet, 55, August 3, 1966.

Physarum albescens Macbr. On decayed twigs, 1.5 miles southeast of
Park Headquarters, 6,600 feet, 940, July 6, 1967. Numerous specimens
of this Myxomycete were collected in the park.

P. auripigmentum G. W. Martin. Beneath layers of decayed wood on
a fallen log, 2 miles south of Park Headquarters, 6,400 feet, 1006, July
8, 1967.

P. decipiens Curt. On decayed twigs, Sleepy Hollow Springs area,
6,600 feet, 7012, July 9, 1967.

DIDYMIACEAE

Diderma de planatum Fries. On decayed wood, 0.5 miles west of Good-
bye Bridge, 6,000 feet, 762, June 17, 1967.

D. nigrum Kowalski. One collection on coniferous twigs, 2 miles south
of Park Headquarters, 6,200 feet, 7041, July 10, 1967. This rare Myxo-
mycete was recently described by Kowalski (1968).

D. niveum (Rost.) Macbr. On bark and fallen twigs, 2 miles west of
Annie Creek Entrance Station, 6,100 feet, 812, June 28, 1967. This slime
mold occurs throughout the park at elevations from 5,000 to 7,000 feet.

D. subcaeruleum Kowalski. On coniferous twigs, 2 miles south of Park
Headquarters, 6,400 feet, 956, July 8, 1967. Kowalski (1968) recently
described this Myxomycete from the park.

Lepidoderma carestianum (Rab.) Rost. One collection on a decayed
twig, 0.5 miles west of Rim Village, 6,800 feet, 7109, July 18, 1967.

L. chailletu Rost. On fallen twigs, Grouse Hill, 7,000 feet, 846, June
29, 1967.

ACKNOWLEDGEMENTS

IT am indebied to Donald T. Kowalski for verifying the determina-
tions and for his assistance throughout the course of this investigation.

282 MADRONO [ Vol. 20

This study was supported in part by the Chico State College Founda-

tion, Grant GU—2095.
Department of Biology, Chico State College, Chico, California

LITERATURE CITED

Curtis, D. H. 1968. Barbeyella minutissima Meylan, A new record for the west-
ern hemisphere. Mycologia 60:708-710.
KowatskI, D. T. 1966. New records of Myxomycetes from California I. Madrofio
18:140-142.
. 1968. Three new species of Diderma. Mycologia 60:595-603.
—. 1968. Observations on the genus Lamproderma Rost. Mycologia 60:756—
768.

. and D. H. Curtis. 1968. New records of Myxomycetes from Califor-
nia III. Madrono 19:246-249.

A NEW VARARIA FROM WESTERN NORTH AMERICA

ROBERT L. GILBERTSON

Collecting in Alberta in 1964 and 1966 and in Arizona in 1967 has
yielded a number of interesting wood-rotting fungi not previously re-
ported from western North America. One of these is a striking species of
Vararia P. Karst. (Basidiomycetes-Thelephoraceae s.l.) described as
new in this paper.

Vararia athabascensis Gilbertson, sp. nov. Fructification effusa, och-
racea vel incarnata, 30-350 » crassa; hyphae nodoso-septatae, 2-3.5 pu
diam; dichohyphidia abundanta, 1—3 » diam, tunicus densus, dextrinoi-
deus; gloeocystidia abundanta, tenitunicata, sinuoso-constricta vel cylin-
dracea, 3-10 » diam; basidia cylindracea-clavata, 30-40 x 6-6.5 p,
4-sterigmatibus; basidiosporae tenuitunicatae, laeves, hyalinae, subclav-
atae, subarcutatae, non-amyloideae, 11-16 X 3-5 p.

Type. Canada, Alberta. Along Athabasca River, Jasper National Park,
on Pinus contorta Dougl., Gilbertson 4752, July 21, 1964 (BPI-holo-
type).

Basidiocarps annual, effused in small patches up to 10 cm long, 30-
350 » thick, not readily separable; margin not differentiated, abrupt to
thinning out; hymenial surface Pale Ochraceous-Buff to Pinkish-Buff
when fresh and on drying, cracking on drying, finely tomentose under a
30x lens; subiculum concolorous with hymenial surface, soft, easily sec-
tioned, uniform in color and consistency.

Sections not darkening in KOH solution, darkening in Melzer’s rea-
gent; generative hyphae of subiculum difficult to discern, thin-walled,
nodose-septate, 2-3.5 w in diam (fig. la) giving rise to the dichohyphidia
and gloeocystidia which are the conspicuous elements of the subiculum;
eloeocystidia abundant, imbedded in subiculum and projecting from
hymenial region, staining deeply in phloxine and also strongly positive
in sulfobenzaldehyde reagent, spherical to elongated, up to 10 p» in diam,

1970 | GILBERTSON: VARARIA 283

Fic. 1. Microscopic characters of basidiocarps of V. athabascensis: a, thin-walled,
nodose-septate subicular hyphae; b, irregularly shaped, imbedded gloeocystidia ;
c, elongated, mammillate gloeocystidia from hymenial region; d, dichohyphidia;
e, basidia; f, basidiospores (type).

but usually 3-5 » in diam, often mammillate (fig. 1b, c); dichohyphidia
abundant, strongly dextrinoid in Melzer’s reagent, especially in the
hymenial region, thick-walled, aseptate, main branches up to 3 » in diam,
some branches long, slender, and unbranched (fig. 1d); hymenial struc-
tures forming a typical catahymenium, with basidia developing in the
mass of dichohyphidia, apparently from spherical probasidia, becoming
elongated, cylindric to narrowly clavate with slight constrictions, swollen
at the base, 4-sterigmate, 6—6.5 » in diam, 30-40 pw long (fig. le); basi-
diospores hyaline, smooth, nonamyloid, attenuated at the basal end and
appearing tear-shaped, 11-16 » long, 3—5 » wide at the distal end (fig.
Het).

Specimens examined. ALBERTA: Grizzly Creek, near Athabasca
River, on P. contorta, Gilbertson 6504, 6515, July 30, 1966. ARIZONA:
Mt. Lemmon, Santa Catalina Mtns., Coronado Nat. Forest, Pima Co.,
on Populus tremuloides Michx. (quaking aspen), Gilbertson 7135, Aug.
16, 1967.

Other species of Vararia known from the Rocky Mountains are V. in-
vestiens (Schw.) Karst., V. racemosa (Burt) Rogers and Jackson, and
V. granulosa (Fries) M. Laur. The growth habit of V. athabascensis

284 MADRONO [Vol. 20

is similar to that of V. investiens, both fruiting on small twigs and
branches in the litter as well as on larger branches and logs on the
ground. Vararia investiens differs in having more slender and abundant
dichohyphidia, spores that taper toward both ends, and lacks the sulfo-
benzaldehyde-positive gloeocystidia. Vararia granulosa differs in its
amyloid spores, less conspicuous gloeocystidia, and larger dichohyphidia.
Vararia racemosa has smal! clusters of densely branched dichohyphidia,
cylindric spores, and broad, mucronate gloeocystidia that are not posi-
tive in sulfobenzaldaldehyde reagent. Welden (1965) and Gilbertson
(1965) give complete descriptions and illustrations of these three species.
Another species, Scytinostroma praestans (Jacks.) Donk, has spores and
cystidia very similar to those of V. athabascensis, but has dendrohyphi-
dia typical of the genus Scytinostroma (Jackson, 1948). Scytinostroma
praestans also is present in the lodgepole pine stands along the Athabasca
River (Gilbertson 6511) and resembles V. athabascensis very closely
macroscopically. The presence of V. athabascensis in widely separated
stations in Alberta and Arizona indicates its probable occurrence in the
Rocky Mountains between those areas.

The decay associated with basidiocarps of V. athabascensis is of the
white rot type with a pale orange discoloration in the early stages. The
positive oxidase reaction of cultures on gallic and tannic acid media and
with gum guaiac support of the field observations of a white rot.

Cultures were obtained from freshly collected basidiocarps by sus-
pending small pieces over the slant surface of 2% Difco malt extract
agar medium in a culture tube. Spore prints were transferred to sterile
tubes as soon as they become discernible. Cultures from which descrip-
tive data were taken were grown on 2% Difco malt extract agar medium
in the dark at 25 C. Gallic and tannic acid media (Davidson, et al,
1938) and gum guaiac solution (Nobles, 1958) were used to test for the
presence of extracellular oxidases. Cultures examined: Gilbertson 6504
and 6515 previously listed.

Growth characters: Growth slow, radial growth 40-50 mm in 17
days; mat white, appressed, with short, radially appressed fibrils, rather
uniform over entire surface in 17 days, with some faint radial zones of
sparser aerial mycelium; margin not differentiated, even to slightly
bayed; cottony aerial mycelium developing around and over original
inoculum after 17 days, this mycelium with a faint pinkish tint; no dis-
tinctive odor; no reverse discoloration on malt agar medium; oxidase
reactions strongly positive with gum guaiac solution and on both gallic
and tannic acid media within 48 hours, no growth on either acid medium.

Microscopic characters: hyphae of advancing zone staining with phlo-
xine, thin-walled, with conspicuous clamp connections, 2.5—6 » in diam
(fig. 2a), giving rise to branches which may become contorted or much-
branched (fig. 2b), frequently branching just behind the transverse sep-
tum of the clamp, commonly with constrictions, these non-staining (fig.

1970] GILBERTSON: VARARIA 285

Fic. 2. Microscepic characters cf cultures of V. athabascensis: a, hyphae from
advancing zone; b, much-branched hypha from advancing zone; c, hyphae from
advancing zone with constrictions; d, large, unbranched hyphae from submerged
mycelium; e, much-branched, slender hypha from submerged mycelium; f, gloeo-
cystidia from submerged mycelium; g, crystals associated with submerged myceli-
um; h, thin-walled hyphae from aerial mycelium; i, slender, thick-walled fiber
hypha trom aerial mycelium; j, gloeocystidia from aerial mycelium; k, vesicular
bedies found in submerged and aerial mycelium after 6 weeks (Gilbertson 6504).

286 MADRONO [ Vol. 20

2c); cordons formed from several intertwined hyphae also frequent;
hyphae of submerged mycelium thin-walled, staining, with conspicuous
clamp connections and also some secondary septa, some extending long
distances with little or no branching, 2.5—6 » in diam (fig. 2d), others
branching frequently, often just behind the transverse septum of a clamp,
eventually giving rise to profusely branched complexes of very slender
and flexuous hyphae tapering down to less than 1 mm in diam (fig. 2e) ;
some hyphae giving rise to gloeocystidia similar to those seen in the
basidiocarp, these often mammillate, highly refractive and varying great-
ly in size (fig. 2f); crystals small plate-like hexagons, rhomboids, or
druse-like clusters (fig. 2g); aerial mycelium with thin-walled hyphae,
these with conspicuous clamp connections and some secondary septa
(fig. 2h), giving rise to fiber hyphae not seen in submerged mycelium
or advancing zone, these hyaline, thick-walled, non-staining in phloxine,
aseptate, 1-2 » in diam, tapering to a very slender tip (fig. 21), genera-
tive hyphae also giving rise to gloeocystidial hyphae and gloeocystidia
as seen in submerged mycelium, these highly refractive in KOH and
phloxine and positive in sulfobenzaldehyde reagent (fig. 2}), generative
hyphae also giving rise to much-branched hyphae as in other areas;
after 6 weeks large, globose or elongated, thin-walled to moderately
thick-walled, hyaline vesicular bodies develop in submerged and aerial
mycelium, these up to 15 » in diam (fig. 2k).

According to the key system proposed by Nobles (1965) the key
pattern for cultures of V. athabascensis would be 2.3.8.15.36.38.44.55.
Using the system proposed by Davidson, et al. (1942), one finds the key
pattern to be E-P-S-1-11-16.

Cultures of V. athabascensis in petri plates eventually develop areas
of brownish mycelium around the periphery of the plates. This brownish
mycelium contains abundant slender fiber hyphae as described above in
addition to other types of hyphae typical of the aerial mycelium. No
dichohyphidia were found in this brownish mycelium and no dextrinoid
reactions were observed. No fruiting structures developed in any of the
plates or tubes under study. Older cultures also showed secondary septa
to be frequent in thin-walled aerial hyphae. The distinguishing charac-
ters of V. athabascensis in culture are the cottony pinkish aerial myce-
lium and the conspicuous gloeocystidia.

Duplicates from all collections cited have been deposited in the Na-
tional Fungus Collections, Beltsville, Md., and the Canadian National
Herbarium, Ottawa. Capitalized color names are based on Ridgway
(1912).

ACKNOWLEDGMENTS

The Alberta collections of V. athabascensis cited in this paper were
made on field trips with the mycology class of the University of Mon-
tana Biological Station, while the author was serving as visiting Asso-
ciate Professor of Botany there during the summers of 1964 and 1966.

1970] REVIEWS 287

This is University of Arizona Agricultural Experiment Station Journal
Paper No. 1350.

Department of Plant Pathology, University of Arizona, Tucson

LITERATURE CITED

Davipson, R. W., W. A. CAMPBELL, and D. J. BiatspeLyi. 1938. Differentiation of
wood-decaying fungi by their reactions on gallic or tannic acid medium. J.
Agric. Res. 57:683-695.

Davipson, R. W., W. A. CAMPBELL, and D. B. VAUGHN. 1942. Fungi causing decay
in living oaks in the eastern United States and their cultural identification.
Tech. Bull. U.S. D. A. 785.

GitBerTson, R. L. 1965. Some species of Vararia from temperate North America.
Pap. Michigan Acad. Sci. 50:161-184.
Jackson, H. S. 1948. Studies of Canadian Thelephoraceae IJ. Some new species of

Corticium. Canad. J. Res., Sect. C, Bot. Sci. 26:143-157.

Nostes, M. K. 1958. A rapid test for extracellular oxidase in cultures of wood-
inhabiting Hymenomycetes. Canad. J. Bot. 36:91-99.

1965. Identification of wood-inhabiting Hymenomycetes. Canad. J.
Bot. 43:1097-1139.

Ripcway, R. 1912. Color standards and color nomenclature. Published by the
author.

WELpEN, A. L. 1965. West Indian species of Vararia with notes on extralimital
species. Mycologia 57:502-520.

REVIEW

Plants of the Oregon Coastal Dunes. By At¥reD M. WIEDEMANN, LA REA J.
Dennis, and FRANK H. Smiru. 117 pp., illus. O.S.U. Book Stores, Inc., Corvallis,
Oregon, 1969. $1.95.

In recent years there has been an increasing variety of inexpensive popular books
that deal with the plants in various regions or habitats of the Pacific coast states.
The present book is concerned with plants of the Oregon coastal sand dunes and
with some of the climatic and geological features of these dunes. It is directed at
“the visitor to the sand dunes, regardless of his background.” The coastal sand
dunes of Oregon are perhaps the best developed of those of any of the Pacific
coast states and support an interesting flora that attracts the attention of ordinary
vacationers as well as more experienced natural historians. The first portion of this
attractive little book is concerned with the physical setting of coastal dunes and
their vegetational history. Subsequent chapters deal with plant communities and
succession and the use of plants for stabilizing dunes. The final chapters present a
key to about 90 characteristic dune plants and descriptions and black and white
photographic illustrations of half of these plants. The quality of the photographs
generally is good, although some of them—such as those of Rumex maritima var.
fueginus, Cakile edentula var. californica, and Lonicera involucrata—are not as
informative as they might be. The level of accuracy is high and the format is a
pleasing one, although I suspect that the level of presentation is somewhat too high
for the average citizen.

This book might be considered superfluous to P.A. Munz’s Shore Wildflowers
of California, Oregon and Washington issued by the University of California Press
in 1964. However, I think that because of its extensive discussion of the ecology of
coastal dunes and its low price, the Wiedemann, Dennis, and Smith volume should
be considered as a complementary volume—if not a replacement in Oregon—for the
Munz book.—Rosert OrnpburFfr, University of California, Berkeley.

288 MADRONO [Vol. 20

Grass Systematics. By FRANK W. GouLp. xi + 382 pp., illustrated. McGraw-
Hill Book Co., New York. 1968. $14.50.

This well designed book brings grass taxonomy up to date. In a systematic and
careful approach Professor Gould has covered all of the classical aspects of grass
taxonomy, and has superimposed upon them the more recent developments in the
field. Thus the introductory three chapters, and part of the fourth, cover micro-
scopic characters such as epidermal and cytological studies, data from biochemical
studies, physiological, ecological and genetic aspects in addition to the expected
discussion of gross vegetative and spikelet morphology. A good balance between
coverage and depth has been achieved; the reader is given basic information on a
topic and liberal references to individual papers should he wish further informa-
tion. The illustrations in this section are well chosen and well executed.

The discussion of characters is concluded in chapter four with the presentation
of a classification system, essentially following that of Stebbins and Crampton
(Rec. Adv. Bot. 1: 133-145. 1961). To those systematists raised on the manuals
and revisions of Hitchcock and Chase this system is radically different, both in the
number of subfamilies as well as their generic content. Essentially many genera
formerly included in the Festucoideae have been shifted to new subfamilies which
reflect more accurately the correlation of new characters with the old. Especially
clever are the maps showing relative geographic representation of these subfamilies.

Chapter five includes keys to and descriptions of the genera of grasses in the
United States. The generic key is basically artificial. While I have not had the op-
portunity to use the key at length, a few chosen genera keyed out with no difficulty.

Most of the genera treated will be familiar but a few unfamiliar names appear,
mostly following usage now common in other countries, especially European, or as
the result of relatively recent monographs and revisions. Among the “new” names
not often seen previously in American agrostological literature are the following
examples. The annual species of Festuca, often treated in U.S. manuals as the
section Vulpza of that genus, are given generic status. Elymus caput-medusae be-
comes Taeniatherum caput-medusae. Trichachne is placed in Digitaria. Paspalidium
is recognized, containing two U.S. species placed in Panicum by Hitchcock. The
four U.S. species in Panicum subgen. Paurochaetium are in Setaria, following a
recent monograph of the latter. Andropogon, Bothriochloa and Schizachrium are
recognized instead of the single genus Andropogon following current thinking of
workers in that group. Neeragrostis is recognized, containing the former Eragrostis
reptans as its single species. Erioneuron, contains five species, all placed in Tridens
by Hitchcock. Allolepis contains one of the species formerly included in Distichlis,
D. texana. Uniola has lost some of its species to Chasmanthium, again reflecting a
recent monograph. Ventenata dubia, reported for the U.S. since Hitchcock’s Manual
is included. No new combinations are made. Representative species of many genera
are illustrated. Most of the drawings are good, some are excellent, some over-
shaded or over-reduced. The placement of some drawings leaves as much as one-
half of the page blank contributing to a “clean” appearance, but wasting space.

The final chapter is devoted to a discussion of grassland associations in North
America, with the recognition of seven associations: true prairie, coastal prairie,
mixed prairie, fescue prairie, palouse prairie, pacific prairie and desert plains
grassland.

A short appendix on the preparation of specimens concludes the basic part of
the book.

Considering its size, but not its value, the book is overpriced, but perhaps we
have been spoiled by the past availability of Hitchcock’s Manual at such a low
price.—DENNIS ANDERSON, Division of Biological Sciences, Humboldt State Col-
lege, Arcata, California.

A WEST AMERICAN JOURNAL OF BOTANY

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The subscription price of Madrofo is $8.00 per year ($4.00 for stu-
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Institutional abbreviations in specimen citations should follow Lanjouw
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Abbreviations of botanical journals should follow those in Botanico-
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Footnotes should be avoided whenever possible.

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ay @

ih
we

ADRONO

VOLUME 20, NUMBER 6 APRIL, 1970

Contents

THE FLORA AND PLANT COMMUNITIES OF
BopEca Heap, CaAtirorniA, M. G. Barbour 289

THE CONSPECIFICITY OF HETEROSIPHONIA ASYMMETRIA
AND H. DENSIUSCULA AND THEIR LIFE HISTORIES

IN CuLtTuRE, John A. West 313
A NEw PROSTRATE VARIETY OF ERIOGONUM APRICUM

(POLYGONACEAE), Rodney Myatt 320
CLARKIA JOLONENSIS (ONAGRACEAE), A NEw

SPECIES FROM THE INNER COAST

RANGES OF CaLirorniA, Dennis R. Parnell 321
CONCERNING THE VALIDITY OF LAMPRODERMA

ECHINOSPORUM, Donald T. Kowalski 323

PERENNATION IN ASTRAGALUS LENTIGINOSUS AND
TRIDENS PULCHELLUS IN RELATION TO
RAINFALL, Janice C. Beatley 326

Reviews: Otto T. Soisric, Principles and Methods
of Plant Biosystematics (John L. Strother) ;
Lyman Benson, The Native Cacti of California
(Ralph N. Philbrick); V. H. HeEywoop (editor),
Modern Methods in Plant Taxonomy (Dennis R.
Parnell); Tuomas M. C. Taytor, Pacific North-
west Ferns and Their Allies (Alan R. Smith) 332

NoTEs AND News: NEw PuBLICATIONS 336

A WEST AMERICAN JOURNAL OF BOTANY

‘BLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

. gn’ HWSO Ty 4 Atyy ae

( NOV 1° 1970
\ /

‘ . Pg
his tne?

MADRONO
A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established
1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional
subscription price $12.00 per year. Published quarterly in January, April, July, and
October by the California Botanical Society, Inc., and issued from the office of
Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley,
California. Orders for subscriptions, changes in address, and undelivered copies
should be sent to the Corresponding Secretary, California Botanical Society, Depart-
ment of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS
CLASS OF:

1970—Lyman BENson, Pomona College, Claremont, California
Mitprep E. Marutas, University of California, Los Angeles

1971—Marion OwnBEY, Washington State University, Pullman
JouNn F. Davinson, University of Nebraska, Lincoln

1972—Ira L. Wiccrns, Stanford University, Stanford, California
ReEeEp C. Rotiins, Harvard University, Cambridge, Massachusetts

1973—WaALLACE R. Ernst, Smithsonian Institution, Washington, D.C.
ROBERT ORNDUFF, University of California, Berkeley

1974—-KEenTON L. CHAMBERS, Oregon State University, Corvallis
EMLEN T. LitTEL, Simon Frazer University, Burnaby, British Columbia

1975—Arturo Gomez Pompa, Universidad Nacional Autonoma de México
Duncan M. Porter, Missouri Botanical Garden, St. Louis

Editor — Jonw H. THomas
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCASKILL |
P.O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Marion Cave, Department of Botany, University of California, Berke-
ley. First Vice-President: Arthur Nelson, Department of Ecology and Systematic
Biology, San Francisco State College. Second Vice-President: Charles T. Mason, Jr.,
Herbarium, College of Agriculture, University of Arizona, Tucson. Recording Sec-
retary: John West, Department of Botany, University of California, Berkeley.
Corresponding Secretary: John Strother, Department of Botany, University of
California, Berkeley. Treasurer: June McCaskill, Department of Botany, University
of California, Davis.

The Council of the California Botanical Society consists of the officers listed
above plus the immediate past President, Harry Thiers, Department of Ecology and
Systematic Biology, San Francisco State College; the Editor of Madrofio; and
three elected Council Members: Robert Ornduff, Department of Botany, University
of California, Berkeley; Malcolm Nobs, Carnegie Institution of Washington, Stan-
ford; and Elizabeth McClintock, Department of Botany, California aaa of
Sciences, San Francisco.

THE FLORA AND PLANT COMMUNITIES
OF BODEGA HEAD, CALIFORNIA

M. G. BARBOUR

The distribution, time of flowering, habit (annual-biennial-perennial) ,
and history (native or introduced) of 215 vascular plant species in a
2.5 mi? coastal peninsula are noted. The flora is divided into six com-
munities: dune, grassland, ocean-facing bluff, saline-wet, fresh-wet, and
disturbed. Those species which occur in two or more very different com-
munities are discussed. Recent history of Bodega Head is summarized,
and aspects of the climate and soil are presented.

INTRODUCTION

Bodega Head, California (38°20’N, 123°04’W) is a coastal penin-
sula which lies about 65 miles north of San Francisco. The 2.5 mi” area
of this study (fig. 1) is limited by State Highway 1 on the northeast,
Salmon Creek on the north, Bodega Harbor on the southeast, and Pacific
Ocean on the west and south. The Bodega Marine Laboratory and
Refuge, owned by the Regents of the University of California, occupies
326 acres near the middle of the peninsula. The facility is used for teach-
ing and research by faculty and students of the Berkeley, Davis, San
Francisco, and Santa Cruz campuses.

According to the Geologic Map of California (1963), the southern
fourth of the peninsula is of Mesozoic granitic rock (tonalite and dio-
rite), the central half is dune sand, and the northern fourth is of Pleis-
tocene marine and marine terrace deposits. The southern fourth (to
which the term “‘head”’ is often restricted) is hilly and edged with steep
cliffs leading to a rocky shoreline. The highest hill reaches 266 feet. The
central dunes are low except for a pronounced foredune (39 feet or less)
and hinddune (145 feet or less). Beachgrass (Ammophila arenaria) has
been planted on the dunes at several times over the past 45 years, and
in addition native plants such as Lupinus arboreus are common on the
hinddune; but sand movement is still considerable and land accretion
along the harbor has resulted under prevailing winds from the north-
west. The Ammophila plants were brought from Golden Gate Park in
San Francisco, and originally came from European stock collected by
Adolf Sutro. About 45 acres of this central section (along the harbor
side) are in a low, fresh-water marsh. The northeastern fourth consists
of gentle hills cut by several gulleys running roughly east-west.

Climatic information for the peninsula itself is extremely sketchy but
is perhaps sufficient to distinguish it from the climate at the nearest
official Weather Bureua station at Fort Ross, 17 miles north. Weather
instruments at the Marine Laboratory include a recording anemometer

Maprono, Vol. 20, No. 6, pp. 289-336. October 29, 1970.

289

290 MADRONO [Vol. 20

SCALE MON
0 (miles) /2

PACIFIC |
OCEAN |

BODEGA BAY

BODEGA

4 Ke HARBOR
Se, so
BML
MARINE LAB oo. REFUGE

4 rn COVE

ne ie

Fic. 1. Map of Bodega Head, boundary of the 326 acre Bodega Marine Labora-
tory (BML) Refuge is shown by dashed lines.

1970] BARBOUR: BOGEDA HEAD 291

30 feet above the ground, a thermograph in a standard shelter 4.5 feet
above the ground, a pyrheliometer, a standard rain gauge, and a max-min
thermometer (Taylor type) hung vertically on a shaded stake such that
the bottom of the U-tube is 2cm above the ground. These instruments
are all located in grassland on the hilly, granitic southern part of the
peninsula. The max-min thermometers are read daily. Rainfall records
have also been collected since July, 1958 by Otto Henninger in the town
of Bodega Bay, less than a mile across the harbor from the center of
the peninsula. A summary of the wind records, from Deceember, 1966
to October, 1968 was kindly provided by J. W. Johnson, Professor of
Hydraulic Engineering at Berkeley.

As at Fort Ross, the warmest month seems to be August or September.
Mean daily maximum (4.5 feet) for August, 1968 was 70°F, mean daily
minimum was 60°; mean daily maximum for January, 1968 was 51°,
mean daily minimum was 37°. Mean max-mins at Fort Ross (over a
period of years) are 68—61° in September and 56—42° in January. Mean
max-mins 2 cm above the ground were 73—53 in August, 1968 and 60—43
in January. Annual rainfall is extremely erratic, ranging from 18 inches
in July, 1958—June, 1959 to 49 inches in July, 1966—June, 1967. Average
rainfall for the 10-year period was 30 inches, and all but 1 inch falls in
the period October-April. In contrast, average annual rainfall at Fort
Ross is 40 inches. Wind is very predominantly out of the northwest. Of
49 equal-length data collecting periods, the prevailing wind was out of
the northwest 63% of the time, out of the southwest 20%, and out of
the southeast 17%. Wind speed is approximately equal throughout the
year, with average wind speed at 10-12 mph. Storms are most frequent
in November, January, and February and during storms billows of foam
may be blown across the dunes and grassland. Fog is especially common
during late spring and early summer. Insolation ranges from 0.15 kcal/
cm?/day in winter to 0.50 kcal/cm?/day in spring.

As an indication of temperatures just above and below the soil in
contrast to air temperature, Table 1 lists air, leaf, and soil temperature
(taken with thermistor leads and a telethermometer) on the south- and
north-facing sides of a small sand hummock in the dune area. The plant
was Ambrosia chamissonis and its leaves were 1 cm from the sand
surface.

Soils are predominantly sandy, ranging from pure sand in the central
dunes to sandy loam in the northeast and southern parts of the penin-
sula. Marked gradients of topsoil salinity exist in the dunes and grass-
land as one moves inland (east) from the shore. At selected points, about
500 g of soil from the top six inches were collected, dried, put through
a Z mm sieve, and a 1:1 (soil: water) extract made. Conductivity of the
extract was measured and converted to ppm total salt (Jackson, 1958).
Samples were collected in grassland from the lip of an ocean-facing bluff
and at 10-m intervals inland to a distance of 50 m. Table 2 presents the

292 MADRONO [Vol. 20

TABLE 1. AIR AND SOIL TEMPERATURES ON NORTH- AND SOUTH-FACING SLOPES
OF A SAND DuNE. OCTOBER 26, 1968, 2 P.M., CLEAR Day. THE LEAF Is OF AMBROSIA
CHAMISSONIS AT 14 INCH ABOVE THE DUNE SURFACE.

Temperature (°F)

Position (inches) South-facing North-facing
+48 65 65
+ 2 76 67
—\y 107 81
— 3 105 69
— 6 66 66
leaf iZ UP

TABLE 2. TOPSOIL SALINITY IN GRASSLAND AS A FUNCTION OF DISTANCE FROM THE
1 Gia

Distance from lip Soluble salts
(meters) (ppm)

6) 1,920
10 960
20 (ey Ih
30 634
40 454
50 314

results. Samples were collected in dunes from the strand where Cakile
maritima was growing and at 50-m intervals to a distance of 750 m,
where Lupinus arboreus was established on the hinddune. Table 3 pre-
sents the results.

In addition, the salinity of soil at the base of ocean-facing bluffs
(which supported dense stands of Scirpus koilolepis, Distichlis spicata
var. stolonifera, and Jaumia carnosa) and of sandy flats at the harbor
edge (which supported Salicornia virginica, Distichlis spicata var. stolont-
fera and Scirpus americanus) was determined by the same method. Aver-
age of two bluff bottom samples was 1,600 ppm; average of five sand
flat samples was 4,100 ppm.

The pH of the 1:1 extract was measured with a Beckman portable pH
meter. Soil pH did not fluctuate with distance from shore, but did differ
between dune (average = 8.3) ad grassland (average = 7.2). Bluff bot-
tom and sand flat pH were similar and equal to that of grassland.

RECENT HISTORY

This short summary is taken principally from Kinnard (1966), Hoover
and Rensch (1948), Hunt and Sanchez (1929), and personal communi-
cations from Cadet Hand.

The Spanish explorer Jaun Francisco de la Bodega y Cuadra discov-
ered Bodega Bay and Bodega Harbor on October 3rd, 1775, but the
Russians were the first white men to settle the immediate area. Ivan
Kuskov, in 1809, built a settlement at the southeast side of Bodega

1970] BARBOUR: BOGEDA HEAD 293

TABLE 3. ToPpsoIL SALINITY IN DUNES AS A FUNCTION OF DISTANCE FROM THE
STRAND.

Distance from strand Soluble salts

(meters) (ppm) Comments
@) 460 strand with Cakile

50 185 foredune
100 122 foredune
150 45 low dunes
200 51 ie
250 32
300 38 oe
350 40
400 42 s
450 36 f
500 45 gh
550 38 s
600 82 Se
650 45 hinddune with Lupinus
700 83 hinddune with Lupinus
750 oo hinddune with Lupinus

Head, near the inlet, at a place now known as Campbell Cove. Kuskov
noted it was a wind-blown site and lacked trees, but did have a spring.
A larger settlement was built at Fort Ross in 1811. Although the area
between Modega Head and Fort Ross was devoid of timber, the Rus-
sians found it adequate for grazing and vegetable growing. Cattle, sheep,
horses, and pigs, bought from the Spanish nearby, heavily grazed the
area, and potatoes became the favorite crop. By 1830, Fort Ross had
become established as a shipyard, and the settlement on the Head became
the principal import-export port for food, bricks, and general supplies. A
brickyard, built near the Head, exported 10,000 bricks in 1830. The
Bay became known to Americans as a good place for ships to take on
water.

Within another decade, however, the Russians came to look at their
California settlements as a liability because the agriculture was no longer
sufficient to sustain the colony without imports, the otter and seal had
been hunted nearly to extinction, and it seemed impossible to negotiate
with Mexico for more land. The Emperor of Russia gave permission to
move to Sitka, and a sale for all movable property was concluded with
Captain John A. Sutter. Heavily in debt, Sutter never made payment,
and the discouraged Russians left in 1841. Voznesenski’s 1841 collection
of the flora near Bodega Head has been discussed by Howell (1937).

The Mexican government claimed the land and divided it into large
ranchros. Rancho Bodega, some 35,000 acres which ran from the Russian
River south to Estero Americano River (present boundary of Marin and
Sonoma Counties), was granted to an American named Stephen Smith
in 1844. He set up a sawmill in a redwood area to the northeast of
Salmon Creek.

204 MADRONO [Vol. 20

TABLE 4. COMPARISON OF CoMMUNITY NAMES IN THIS STUDY WITH THOSE DE-
SCRIBED BY Muwz (1959).

Bodega Head Munz
dune coastal strand
grassland coastal prairie
ocean-facing bluff coastal strand + northern coastal scrub
saline-wet coastal salt marsh
fresh-wet freshwater marsh + northern coastal scrub
disturbed no equivalent

The goldrush and statehood swelled the population. Rancho Bodega
became broken up into several holdings on the peninsula alone, and the
Gaffney family came to own over 400 acres in the center. Mrs. Rose
Gaffney, who came to Bodega Head in 1913, still lives in Salmon Creek.
She recalls that much of the present dune area was pasture in the early
1900’s, but that sand continually encroached from the west. Raising
potatoes and grazing dairy herds continued to be prime land uses until
the 1930’s, when many marginal dairy herds were exchanged for sheep.
Mrs. Gaffney claims that sheep were grazed on her property for only
two months in 1942, but that cattle and horses were regularly present.
She remarked that the show of early-summer flowers now (since the
property has been a preserve) is much more spectacular than at any
previous time.

In 1959, a Chancellor’s Committee (Berkeley) For The Selection of
a Marine Station Site recommended Bodega Head, and in 1962 the Uni-
versity of California acquired most of the Gaffney property and estab-
lished it as a reserve. Over 200 acres at the tip of Bodega Head was
purchased by Pacific Gas and Electric Company in 1960, with an eye to
establishing an atomic-powered steam generator for production of elec-
tricity. Proximity of the site of the San Andreas fault, however, led to
abandonment of the idea and the land is currently being leased to
Sonoma Co. as a recreation area. Most of the dune area was purchased
by the State of California in 1962 and incorporated into Sonoma Coast
State Beach. The Laboratory was constructed in 1966 with the help
of a National Science Foundation grant, and the facility is currently
funded by the University of California.

PLANT COMMUNITIES

The flora and community descriptions were compiled in the course of
monthly visits to the area over a period of more than 2 years. Un-
doubtedly there are species and varieties which I have missed, and
undoubtedly my choice of communities could be refined with further
field work. However, at this point in time, there appear to be six commu-
nities on the peninsula: dune, grassland, ocean-facing bluff, saline-wet,
fresh-wet, and disturbed. In comparing my species lists for these com-
munities to similar communities described by Munz (1959), it became

1970] BARBOUR: BOGEDA HEAD 295

TaBLe 5. Cover ALONG A 500-M X 1%3-m Strip TRANSECT OF DUNE (RUNNING
FROM STRAND TO HINDDUNE) IN JUNE, 1968.

Species Cover (%)

bare 49
Ammo phila arenaria 30
dead herbage (mostly of

Ammophila and Mesembryanthemum) 18
Mesembryanthemum chilense 1
Lupinus arboreus il
others 1

apparent that the two lists of communities were not quite the same.
Table 4 summarizes the relationships between my choice of communities
and their closest equivalents in Munz.

Dune. This community covers the largest area (39% of the penin-
sula). It is characterized by only a few common species, and a change
in species with increasing distance from shore. On the strand itself is only
one species, Cakile maritima, and it occurs in scattered clumps. Ammo-
phila arenaria, planted in rows parallel to the shore, dominates the
ground from the foredune back. It is especially dense on the foredune.
Behind the foredune, Mesembryanthemum chilense, Lotus heermannit,
and Camissonia cheiranthifolia are occasional; Cakile is absent. Finally,
about 700 m from the strand, the hinddune is reached and Lupinus
arboreus becomes common. On dunes dominated by Lupinus, and pre-
sumably older, many other species are common: Abronia latifolia, A go-
seris apargioides, Baccharis pilularis ssp consanguinea, Aplopappus ert-
coides, Elymus vancouverensis, Ambrosia chamissonis, and Poa douglasit.

In June, 1968, I noted plant cover along a 500-m *& 14-m strip transect
which ran from strand to hinddune. Table 5 summarizes plant cover
along the transect.

Grassland. Grassland covers almost as large an area (36% of the
peninsula) as dune. In contrast, it is marked by a great diversity of
species and a changing seasonal aspect. Very generally, the grassland is
dominated by annual and perennial herbs and annual grasses. Lupinus
arboreus dominates patches of grassland, but is nearly absent over much
of it (but small seedlings are occasional to common). Many Lupinus
shrubs—sometimes singly, sometimes in clusters, sometimes large, some-
times small—exhibit wilted foliage. With time the leaves turn gray and
then fall, leaving a skeleton of apparently dead branches. If recently
afflicted plants are uprooted, it is seen that the roots are nearly hollowed
out by the burrowing activity of a small larva, identified as Hepialus
behrensi Stretch by W. H. Lange, Professor of Entomology at Davis.
An additional pest of Lupinus, prevalent in summer, is the caterpillar
of the salt marsh moth, Estigmene acraea, identified by Paul Hurd, Pro-
fessor of Entomology at Berkeley.

296 MADRONO [Vol. 20

TABLE 6. Cover ALONG A 200-M X 4%%-M STRIP TRANSECT OF GRASSLAND (RUN-
NING FROM Lip OF OCEAN-FACING BLUFF INLAND) IN JUNE, 1968.

Species Cover (%)

dead herbage (mostly of Lupinus and

scattered litter) 33
annual grasses (many dying) 32
perennial and annual herbs 30
perennial grasses (mainly

Bromus carinatus) 2
bare 2
Lupinus arboreus 1

The spring grassland is a wet green color, dominated by Montia per-
foliata, Stachys rigida, Luzula subsessilis, with occasional color from
Arabis blepharophylla, Amsinckia menziesiu, Iris douglasiana, Nemo phila
menziesii, and Ranunculus californicus.

The early summer grassland is a carpet of yellow petals, principally
of Lasthenia chrysostoma, Eschscholzia californica, Layia platyglossa,
and Platystemon californicus. Less abundant herbs includes Marah faba-
ceus, Phacelia distans, Silybum marianum, Sisyrinchium bellum, and
Sonchus spp. By June and July the grasses dominate the community.
The most common species are Aira caryophylla, Bromus arvensis, B.
diandrus, and Lolium multiflorum, all introduced. The most common
native grass, the perennial Bromus carinatus, is much less common than
the others.

In June, 1968, I noted plant cover along a 200-m *& 14-m strip tran-
sect which ran across typical grassland from the lip of an ocean-facing
bluff inland. Table 6 shows annual and perennial cover along the transect.

The late summer grassland is dull brown in color from dead annual
grasses. Eschscholzia californica and Lupinus arboreus continue to flower,
and Achillea borealis ssp. arenicola and Cirsium vulgare are common.
Holcus lanatus forms dense stands in low spots, and its purple-tinged
florets add a little more color.

Throughout the year, several species are restricted to the rocky crests
of grassland hills: Avabis blepharophylla, Brodiaea pulchella, Chloro-
galum pomeridianum, Iris douglasiana, Luzula subsessilis, and Rhus
diversiloba.

Ocean-facing bluff. Only 7% of the area is dominated by this commu-
nity, which occurs at the lip and rocky shelf at the base of ocean-facing
bluffs. When the bluff is cut with a gulley so that the face is no longer
vertical, the same community occurs down the face. Characteristic
species include the low shrubs Artemisia pycnocephala and Eriophyllum
staechadifolium, and the perennial herbs Armeria maritima var. califor-
nica, Mesembryanthemum chilense, Plantago maritima var. californica,
and Spergularia macrotheca. The annual bulrush, Scirpus koilolepis, is

1970] BARBOUR: BOGEDA HEAD 297

restricted to the basal shelf and often occurs with Distichlis spicata and
Jaumia carnosa. If a seep runs down to the shelf, species of grassland or
disturbed communities may be present: Anagallis arvensis, Polypogon
mons peliensis, Sonchus spp.

Saline-wet. This community dominates the saline, sandy flats at the
edge of Bodega Harbor—about 1% of the area of this study. The species
are principally low, rhizomatous perennials: Distichlis spicata, Frankesia
grandifolia, Jaumia carnosa, Salicornia virginica, Scirpus americanus.
Further from shore, and less common, are Atriplex patula ssp. hastata
and Holcus lanatus.

The water table (brackish water) lies 15 cm or less beneath the sur-
face. Algal mats often coat the ground surface. In October the upper
shoots of Salicornia turn bright red and make this ordinarily dull-
colored, monotonous community more lively.

A prominent species found here but not along saline cliff basis is
Scirpus americanus, and species common to the cliff bases but not found
here are Mesembryanthemum chilense, Scirpus koilolepis, and Spergu-
laria macrotheca.

Fresh-wet. Members of this community range in habitat from fresh-
water marshes to soil near seeps to moist, shaded hillsides and gulleys to
depressions which exhibit standing water only during the wet season.
Fresh-water marshes are dominated by Scripus microcarpus and Spar-
ganium eurycarpum in the center, Juncus balticus, Juncus leseurii,
Oenanthe sarmentosa, and Potentilla egediu var. grandis near the edge.
Surprisingly, Oenanthe and Potentilla also occur at the edge of some
saline flats.

Depressions which are seasonally wet support Cotula coronopifolia,
Scripus microcarpus and Typha angustifolia. Cotula does also occur in
the center of fresh-water marshes, but is not very prevalent there.

Shaded banks, gulleys, and seeps support a great variety of species,
principally Anaphalis margaritacea, Calamogrostis nutkaensis, Castilleja
wrightit, Conium maculatum, Equisetum telmateia var. braunu, Hera-
cleum lanatum, Mimulus guttatus ssp. litoralis, Nasturtium officinale,
Polystichum munitum, Rubus procerus, Salix laevigata, Salix lasiole pis,
Stellaria media, and Vicia gigantea. There are also a great number of
uncommon species, which will not be listed here, except for the rarest:
Fritillaria recurva, only one specimen seen during the entire two years.

This diverse fresh-wet community occupies about 5% of the study
area.

Disturbed. “Disturbed” must be a relative term here, for the entire
area has been severely disturbed by grazing, farming, and human activ-
ity over the past 150 years. However, plants which are placed in this
community occupy sites continuously being traveled over such as road-
sides and footpaths. In this light, the disturbed community occupies
12% of the total area.

298 MADRONO [Vol. 20

TABLE 7. SPECIES WITH LONG FLOWERING PERIODS.

Species Flower ing period
Abronia latifolia April-October
Brassica campestris March-October
Cakile maritima March-October
Castilleja wrightu April-October
Cotula coronopifolia March-October
Erodium cicutarium March-October
Eschsholzia californica March-October
Hypochoeris radicata April-October
Mimulus guttatus ssp. litoralis April-October
Nemophila menziesii March-October
Mesembryanthemum chilense March-September
Camissonia cheiranthifolia April-October
Sonchus spp. April-September

Along grassland paths are Lasthenia minor, Calandrinia ciliata, Car-
dionema ramosissimum, Hypochoeris radicata, Orthocarpus erianthus
(in patches), Plantago lanceolata, Phacelia distans, and Spergularia
rubra.

Roadsides show tremendous fluctuations in seasonal aspect. In spring,
Brassica campestris and Raphanus sativus dominate; in early summer,
Brassica nigra, Cotula coronptfolia, Lotus corniculatus, and Polypogon
mons peliensis dominate; and in late summer and fall Baccharis pilularis
ssp. consanguinea, Conyza canadensis, Epilobium adenocaulon var. occi-
dentale, Foeniculum vulgare, Melitotus albus, Rumex crispus, and Rubus
vitifolius dominate.

MISCELLANEOUS NOTES

The flora of Bodega Head consists of at least 215 species, representing
157 genera and 56 families. Introduced species make up 36%. Seven
introduced species were probably only noted where planted and should
not technically be included in the flora: Acacia longifolia, Ceanothus
thyrsiflorus var. repens, Cupressus macrocarpa, Eucalyptus globulus,
Myoporum laetum, Pinus muricata and P. radiata.

Some 14 species exhibited very erratic flowering times and were almost
equally in flower for a period of 7-8 months. Table 7 lists these species
and their flowering period. Only three of these species showed a major
peak in flowering within that long period: Abronia latifolia (July-
August), Brassica campestris (March-April), and Nemophila menziesii
(March-April). Cakile, Cotula, Mesembryanthemum, and Sonchus had a
few flowers even in December.

Another 11 species were found in two or more quite different habitats;
their distributions are summarized in Table 8.

Voucher specimens of all species included in the check list have
been deposited in the herbarium of the University of California, Davis
(DAV).

1970] BARBOUR: BOGEDA HEAD 299

TABLE 8. SPECIES WITH UNUSUAL DISTRIBUTIONS.

Species Distribution
Anagallis arvensis roadsides, base of ocean bluff in seep
Cotula coronopifolia roadsides, wet ditches, fresht-water marsh
Dudleya farinosa lip of ocean-facing bluff and lip of gulley
Grindelia stricta ssp. venulosa roadsides, ocean-facing bluff
Holcuslanatus edge of saline flat, wet area of grassland
Mesembryanthemum chilense dunes, lip of ocean-facing bluff
Oenanthe sarmentosa edge of fresh-water and salt-water marshes
Potentialla egedii var. grandis edge of fresh-water and salt-water marshes
Polypogon mons peliensis roadside, base of ocean-facing bluff (in

seep), wet, shaded stream bank
Sonchus spp. grassland, base of ocean-facing bluff in seep
Solanum nodiflorum stabilized dunes, wet, shaded stream bank
ACKNOWLEDGMENTS

Travel expenses and supplies were paid for by a campus research
grant issued through the Opportunity Fund. Weather data for Bodega
Head were collected by Ken Van der Laan, Otto Henniger, and J. W.
Johnson. Some of the salinity measurements were made by Frank Drys-
dale. John Tucker advised in the establishment of a reference herbarium
at the Laboratory. Some important historical information came through
the cooperation of Mrs. Rose Gaffney and Cadet Hand. I would especial-
ly like to thank the Director of Bodega Marine Laboratory, Cadet Hand,
for his general help to me during the course of this study.

CHECK LIST
Calamophyta

Equisetaceae

Equisetum arvense L. Common horsetail. Perennial, native. Occasional
to common in seasonally wet, but disturbed areas along roads and on
steep, wet hillsides. Only vegetative shoots seen.

Equisetum telmateia Ehr. var. braunii Milde. Giant horsetail. Peren-
nial, native. Common to abundant on steep, wet hillsxides. Fertile shoots
produced in March.

Pterophyta

Aspidiaceae
Athyrium felix-femina (L.) Roth var. sitchense Rupr. Lady fern. Per-
ennial, native. Rare to occasional in shaded gulley near stream.
Polystichum munitum (Kaulf.) Presl. Sword fern. Perennial, native.
Common in large clumps on steep, wet hillsxides.

300 MADRONO [Vol. 20

Polypodiaceae
Poly podium scoulert Hook. & Gray. Polybody. Perennial, native. Occa-
sional on grassland hill-tops next to rocks.

Pteridiaceae
Pteridium .aquilinum (L.) Kuhn var. lanuginosum (Bong.) Fern
Bracken. Perennial, native. Occasional in grassland, more common when
Lupinus arboreus is present.

Coniferophyta

Cupressaceae
Cupressus macrocarpa Hartw. ex Gordon. Monterey cypress. Peren-
nial, planted or escaped. Rare along roads, in dunes, near Marine
Laboratory.

Pinaceae
Pinus muricata D. Don. Bishop pine. Perennial, probably planted.
Rare along road near Salmon Creek.
P. radiata D. Don. Monterey pine. Perennial, probably planted. Occa-
sional along roads.

Anthophyta—DICOTYLEDONEAE

Aizoaceae
Mesembryanthemum chilense Mol. Sea-fig. Perennial, introduced.
Common along ocean-facing cliffs (especially at the lip) and on dunes.
Flowering sporadic, March-September.
M. edule L. Hottentot-fig. Perennial, Introduced. Common in dunes,
planted along roadbanks. Flowering March-July.

Anacardiaceae
Rhus diversiloba T. & G. Poison oak. Perennial, native. Occasional in
erassland and along shaded streambank. Only vegetative shoots seen.

Apocynaceae
Vinca major L. Periwinkle. Perennial, introduced. Possibly planted in
dense strips along one road and at edge of a fresh-water marsh; other-
wise rare. Flowering in March and October, possibly in months between.

Berberidaceae
Berberis pinnata Lag. Barberry. Perennial, native. Common on one
erassland hill, otherwise rare. Only vegetative shoots seen.

Boraginaceae
Amsinckia menziesii (Lehm.) Nels. & Macbr. Fiddleneck. Annual, na-
tive. Common in grassland; variable in size and leaf shape. Flowering
April-June.
A. spectabilis F. & M. Fiddleneck. Annual, native. Common on estab-
lished dunes (with Lupinus arboreus); prostrate and sparsely hispid.
Flowering June-July.

1970] BARBOUR: BOGEDA HEAD 301

Cry ptantha leiocarpa (F.&M.) Greene. Annual, native. Occasional in
dunes. Flowering June.

Plagiobothrys tenellus (Nutt.) Gray. Annual, native. Occasional to
common in disturbed areas. Flowering April-May.

Caryophyllaceae

Cardionema ramosissimum (Weinm.) Nels. & Macbr. Perennial, native.
Occasional to common as prostrate mats in grassland footpaths. June-
July.

Silene gallica L. Windmill pink. Annual, introduced. Occasional in
erassland. May-June.

Spergula arvensis L. Spurrey. Annual, introduced. Occasional in dis-
turbed areas. March-July.

Spergularia macrotheca (Hornem.) Heynh. Sand-spurrey. Perennial,
native. Occasional to comon on shelf at base of ocean-facing cliff, also
occasional along path in grassland; prosetrate. May-August.

S. rubra (L.) J. & C. Presl. Sand-spurrey. Annual, native. Common
in disturbed areas along roads. March-July.

Stellaria media (L.) Cyr. Chickweed. Annual, introduced. Common in
grassland, abundant along shaded stream. March-April.

Chenopodiaceae

Atriplex patula L. ssp. hastata (L.) Hall & Clem. Annual, introduced.
Occasional at outer edge of saline flats near shore. August.

A. patula L. ssp. obtusa (Cham.) Hall & Clem. Annual, introduced.
Occasional to rare on saline flat near ocean. October.

Chenopodium album L. Pigweed. Perennial, introduced. Rare in sea-
sonally wet area along road. July.

C. ambrosoides L. var. vagans (Standl.) Howell. Mexican tea. Annual-
perennial, native. Rare in disturbed areas. August-September.

C. californicum (Wats.) Wats. Pigweed. Perennial, native. Occasional
in grassland, more common in disturbed areas. April-June.

Salicornia virginica L. Pickleweed. Perennial, native. Abundant in
saline flats near shore, common along shelf at base of ocean-facing cliff.
October; inflorescences and upper stems becoming reddish at that time.

Compositae

Achillea borealis Bong. ssp. arenicola (Hel.) Keck. Yarrow. Perennial,
native. Common in grassland, established dunes (with Lupinus arboreus),
and occasional in disturbed areas along roads. May-October.

Agoseris apargioides (Less.) Greene ssp. maritima (Sheld.) Jones.
Beach dandelion. Common on established dunes. Flowering sporadic,
April-October. Perennial, native.

Anaphalis margaritacea (L.) B. & H. Pearly everlasting. Perennial,
native. Common to occasional on steep, shaded, wet hillsides. July-
October.

Haplopappus ericoides (Less.) H. & A. Perennial, native. Occasional

302 MADRONO [Vol. 20

on established dunes (with Lupinus arboreus). August-October.

Artemisia douglasiana Bess. in Hook. Sagebrush. Perennial, native.
Semi-prostrate; occurs in dense clusters on established dunes, in grass-
land, and along roads; generally rare. August-October.

A. pycnocephala DC. Sagebrush. Perennial, native. Occasional along
lip of ocean-facing bluff. July-September.

Aster chilensis Nees. Perennial, native. Occasional in grassland. Sep-
tember.

Baccharis pilularis DC. ssp. consanguinea (DC). Wolf. Coyote bush.
Perennial, native. Rare in dunes, occasional in disturbed areas along
roads. September-October.

Carduus pycnocephalus L. Italian thistle. Annual, introduced. Occa-
sional in disturbed areas. May-June.

Centaurea solstitialis L. Star thistle. Annual, introduced. Rare along
roads. September.

Chrysanthemum segetum L. Corn chrysanthemum. Annual, introduced.
Rare in disturbed areas along roads. June-July.

Cichorium intybus L. Chicory. Perennial, introduced. Rare in dis-
turbed areas along roads. August.

Cirsium andrewsti (Gray) Jeps. Thistle. Perennial, native. Flowering
stalk short (hardly higher than basal leaves); occasional in grassland.
May-July.

C. occidentale (Nutt.) Jeps. Thistle. Perennial, native. Tall (to 2 m)
and clumped, herbage covered with dense arachnoid pubescence; occa-
sional on established dunes. May-July.

C. vulgare (Savi) Ten. Bull thistle. The most common of the three
thistles; common in grassland and in shaded, wet hillside. July-October.

Conyza canadensis (L.) Crong. Horseweed. Annual, introduced. Com-
mon to abundant in disturbed areas along roads. September-October.

Cotula coronopifolia L. Brass buttons. Perennial, introduced. Com-
mon in fresh-water marshes and near roads. Flowering sporadic. March-
October.

Erechtites arguta (A. Rich.) DC. Fireweed. Annual, introduced. Oc-
casional along roads. July-August.

E. prenanthoides (A. Rich.) DC. Fireweed. Annual, introduced. Occa-
sional at edge of fresh-water marsh, common along steep, wet hillsides,
sometimes near roads. July-September.

Erigeron glaucus Ker. Seaside daisy. Perennial, native. Occasional on
established dunes and cliff edges, rare in grassland. June-September.

Eriophyllum lanatum (Pursh) Forbes var. arachnoideum (Fisch. &
Ave-Lall.) Jeps. Perennial, native. Rare to occasional in dunes. May.

E. staechadifolium Lag. Perennial, native. Common along lip of ocean-
facing bluff. July-September.

Ambrosia chamissonis (Less.) Greene. Perennial, native. Common on
dunes. Two forms occur intermixed: one form with broadly lobed leaves,

1970] BARBOUR: BOGEDA HEAD 303

the other with smaller, pinnate to bipinnate lobes. The two leaf forms
do not occur on the same plant, but the two plants may grow side-by-
side. July-August.

Gnaphalium chilense Spreng. Cud-weed. Annual-biennial, native. Oc-
casional in grassland and along roads. May-October.

G. chilense Spreng. var. confertifolium Greene. Cud-weed. Annual-
biennial, native. Less common than the species; grassland and along
roads; June.

G. purpureum Locc. Cud-weed. Annual-biennial, native. Rare along
road. May.

Grindelia stricta DC. ssp. venulosa (Jeps.) Keck. Gum-weed. Peren-
nial, native. Occasional along road and ocean-facing bluff. May-Sep-
tember.

Hypochoeris radicata L. Hairy cat’s ear. Perennial, introduced. The
most common member of Compositae; variable in flower size and color
(yellow to gold-orange) ; sometimes with swollen stems. Common in dis-
turbed areas, especially along grassland footpaths. Sporadic flowering,
April to October.

Jaumia carnosa (Less.) Gray. Perennial, native. Occasional to com-
mon in saline flats near coast and at base of ocean-facing bluff on shelf.
September-October.

Lasthenia chrysostoma (F.&M.) Greene. Goldfields. Annual, native.
Common to abundant in grassland and in paths through it. May-June.

L. minor (DC.) Ferris. Annual, native. Common in footpaths, less
common in grassland. March-April.

Lavia platvglossa (F. & M.) Gray. Tidy tips. Occasional to common
in grassland and along footpaths through it. May-July.

Madia sativa Mol. Coast tarweed. Annual, native. Common in grass-
land, occasional in disturbed areas along roads. May-October.

Silybum marianum (L.) Gaertn. Milk-thistle. Annual-biennial, intro-
duced. Occasional in grassland. April-June.

Solidago californica Nutt. California goldenrod. Perennial, native.
Rare along shaded stream bank. September.

Sonchus asper L. Sow-thistle. Annual, introduced. Common to abun-
dant in grassland, occasional along roads and on shelf at base of ocean-
facing cliff. Flowering sporadic. April-September.

S. oleraceus L. Sow-thistle. Annual, introduced. Distribution and flow-
ering as with S. asper.

W yethia angustifolia (DC.) Nutt. Perennial, native. Rare along roads.
April.

Convolvulaceae

Convolvulus occidentalis Gray var. saxicola (Eastw.) Howell. Morn-

ing glory. Perennial, native. Occasional in grassland. April-May.

Crassulaceae
Dudlevya farinosa (Lindl.) Britt. & Rose. Live-forever. Perennial, na-

304 MADRONO [Vol. 20

tive. Occasional on ocean-facing bluff, on rocky hill tops, and at lip of
wet, shaded gulley. July-August.

Cruciferae

Arabis blepharophylla H. & A. Rock-cress. Perennial, native. Common
near summits of grassland hills. March-April.

Barbarea orthoceras Ledeb. Winter-cress. Biennial-perennial, native.
Rare in disturbed part of grassland. March-April.

Brassica campestris L. Field mustard. Annual, introduced. Common
along roads, occasional in seasonally wet sites. Principal flowering time
March-April, a few plants in flower in October, possibly some flowered
during intervening months.

B. nigra (L.) Koch. Black mustard. Annual, introduced. Common
along roads. Principal flowering time May-August, some through October.

Cakile maritima Scop. Sea rocket. Annual, introduced. Common fac-
ing ocean on outer-most dunes or on strand, once seen inland near road
in seasonally wet area; occurs in clumps which seem to build mounds of
sand. Although C. edentula var. californica has been reported for the
area, I have yet to see it. Flowering sporadic, March-October.

Cardamine oligosperma Nutt. Bitter-cress. Annual-biennial, native.
Common in grassland in patches, the siliques popping open and spraying
out seeds as one walks through in April. Flowering in March.

Nasturtium officinale R. Br. Water-cress. Perennial, introduced. Com-
mon to abundant on very wet shaded streambank and in the stream
itself. June-August.

Raphanus sativus L. Wild radish. Annual-biennial, introduced. Com-
mon along roads, occasional in grassland; leaves may be smooth or
hispid; petals white, yellow, or blue. Principal flowering time March-
July, a few in flower in September-October.

Rorippa curvisiliqua (Hook.) Bessey. Yellow-cress. Annual-biennial,
native. Rare in disturbed area of grassland. June.

Cucurbitaceae
Marah fabaceus (Naud.) Dunn. Manroot. Perennial, native. Common
in grassland dominated by Lupinus arboreus. March-June.

Cuscutaceae
Cuscuta salina Engelm. Dodder. Perennial, native. Occasional on
Salicornia in salt marsh. June.
C. subinclusa Dur. & Hilg. Dodder. Perennial, native. Occasional on
Jaumia in salt marsh. June.

Frankeniaceae
Frankemia grandifolia C. &S. Perennial, native. Occasional to common
at edge of salt flats near ocean. August.

Geraniaceae
Erodium cicutarium (L.) L’Her. Red-stem filaree. Annual, introduced.

1970] BARBOUR: BOGEDA HEAD 305

Common in disturbed areas of grassland. Flowering sporadic, March-
October.

E. moschatum (L.) L’Her. White-stem filaree. Annual, introduced.
Also in disturbed areas, but less common than £. cicutarium. March-
May.

Hydrophyllaceae

Nemo phila menziesii H. & A. Baby-blue-eyes. Annual, native. Common
in grassland; variable in petal color (white to dark blue). Principal flow-
ering time March, some flowering to August.

Phacelia californica Cham. Perennial, native. Occasional along lip of
ocean-facing bluffs. Not seen flowering.

P. distans Benth. Wild heliotrope. Annual, native. Common in
disturbed areas of grassland, less common in grassland. April-August.

Labiatae

Mentha pulegium L. Pennyroyal. Perennial, introduced. Occasional on
established dunes. September.

Stachys rigida Nutt. ex Benth. ssp. quercetorum (Heller) Epl. Hedge-
nettle. Perennial, native. Common to abundant in grassland, less notice-
able in late summer and fall. Principal flowering in April-May, some
flowering to October.

Leguminosae

Acacia longifolia Willd. Perennial, introduced. Probably planted, rare
along seasonally wet roadside. April.

Cytisus mons pessulanus L. French broom. Perennial, introduced. Rare
along roadsi des. September.

Lotus corniculatus L. Bird’s foot trefoil. Perennial, introduced. Com-
mon to abundant along roadsides, creating solid strips of yellow when
in flower. June-August.

L. heermanii (Dur. & Hilg.) Greene var. ertophorus (Greene) Ottley.
Bird’s foot trefoil. Perennial, native. Occasional in dunes; petals red to
yellow. May-June.

L. subpinnatus Lag. Bird’s foot trefoil. Annual, native. The least at-
tractive and least common on the three trefoils; rare along roads. August.

Lupinus arboreus Sims. Lupine. Perennial, native. Common in grass-
land (abundant in patches), on stabilized dunes at some distance from
shore, and occasional on ocean-facing bluffs. April-August; flowers vari-
able in color even on same shrub (white, yellow, blue).

L. bicolor Lindl. ssp. umbellatus (Greene) Dunn. Lupine. Annual,
native. Occasional along roads. April-July.

Medicago polymorpha L. var. vulgaris (Benth.) Shinners. Bur-medick.
Annual, introduced. Occasional to common along roads. April.

Melitotus albus Desr. White sweet-clover. Annual-biennial, introduced.
Occasional to common along roads. August-September.

M. indicus (L.) All. Yellow sweet-clover. Annual-biennial, introduced.
Occasional along roads. April-July.

306 MADRONO [Vol. 20

Trifolium barbigerum Torr. Clover. Annual, native. Occasional in dis-
turbed areas. April.

T. fucatum Lindl. Clover. Annual, native. Rare along lip of ocean-
facing bluff. May.

T. repens L. White clover. Perennial, introduced. Occasional to com-
mon along roads. June.

T. wormskioldu Lehm. Clover. Perennial, native. Rare to occasional in
disturbed areas of grassland. May-July.

Vicia americana Muhl. ssp. oregana (Nutt.) Abrams. Vetch. Peren-
nial, native. Occasional to rare in grassland and disturbed areas of
grassland. April.

V. californica Greene. Vetch. Perenial, native. Occasional to common
in grassland and disturbed areas of grassland. May.

V. gigantea Hook. Vetch. Perennial, native. Common on steep, wet
hillsides. March-July.

Malvaceae
Lavatera arborea L. Tree-mallow. Perennial, introduced. Rare along
roadsides. July.
Sidalcea malviflora (DC.) Gray ex Benth. ssp. laciniata Hitchck.
Checker. Perennial, native. Rare along roadsides. April.

M yoporaceae
Mvyoporum laetum Forst. Perennial, introduced. Planted near Marine
Laboratory. Flowering sporadic, March-July.

Myricaceae
Myrica californica C. & S. Wax myrtle. Perennial, native. Rare on
steep, wet hillsides. February.

Myrtaceae
Eucalyptus globulus Labil. Tasmanian blue-gum. Perennial, intro-
duced. Planted or possibly escaped in wet, shaded gulley. April-August.

Nyctaginaceae
Abronia latifolia Esch. Sand verbena. Perennial, native. Common on
established dunes (with Lupinus arboreus), occasional along roads in
grassland. Flowering sporadic, April-October.

Onagraceae

Epilobium adenocaulon Hausskn. var. occidentale Trel. Willow-herb.
Perennial, native. Occasional along roads. August-September.

E. watsonu Barbey var. franciscanum (Barbey) Jeps. Willow-herb.
Perennial, native. Rare along roads. August-September.

Camissonia cheiranthifolia (Hornem. ex Spreng.) Raimann. Evening
primrose. Common on established and shifting dunes. Flowering spo-
radic, April-October.

1970] BARBOUR: BOGEDA HEAD 307

Oxalidaceae
Oxalis corniculata L. Wood-sorrel. Pearennial, introduced. Occasional
in grassland. September.
O. pes-caprae L. Wood-sorrel. Perennial, introduced. Rare in season-
ally wet area near road. March.

Papaveraceae

Eschscholzia californica Cham. California poppy. Perennial, native.
One of the most comon species in grassland and on established dunes
(with Lupinus arboreus). Flowering sporadic, March-October.

Platystemon californicus Benth. Cream cups. Annual, native. Com-
mon to abundant in grassland; some petals all white, others with yellow
tpis; variable in size of plant; together with Lasthenia chrysostoma and
Eschscholzia californica, forms much of the spring color show. April-July.

Plantaginaceae
Plantago lanceolata L. Plantain. Perennial, introduced. Occasional to
common in disturbed areas. April-July.
P. maritima L. var. californica (Fern.) Pilg. Plantain. Perennial, native.
Common at base of ocean-facing bluffs. May-July.

Plumbaginaceae
Armeria maritima (Mill.) Willd. var. californica ( Boiss.) Lawr. Thrift.
Perennial, native. Abundant along lip of ocean-facing bluffs, common at
their base. April-July.

Polemonicaceae

Gilia capitata Sims. var. chamissonis (Greene) Grant. Annual, native.
Rare on established dunes (with Lupinus arboreus). May.

Navarretia squarrosa (Eschs.) H. & A. Skunkweed. Annual, native.
Occasional on established dunes (with Lupinus arboreus); giving off a
strong skunklike odor easily detected when walking near the plants.
June-July.

Polygonaceae

Chorizanthe cuspidata Wats. var. villosa (Eastw.) Munz. Annual,
native. Occasional along footpath in grassland. June.

Eriogonum latifolium Sm. Wild buckwheat. Perennial, native. Occa-
sional in grassland at some distance from shore, and along lip of ocean-
facing bluffs. July-September.

Polygonum patulum Biebst. Knotweed. Annual, introduced. Rare along
roads. May.

Pterostegia drymarioides F. & M. Annual, native. Rare in grassland.
April.

Rumex acetosella L. Sheep sorrel. Perennial, introduced. Common in
grassland and disturbed areas of grassland. March-July.

R. crispus L. Curly dock. Annual, introduced. Occasional along roads.
May-July.

308 MADRONO [Vol. 20

R. pulcher L. Fiddle dock. Perennial, introduced. Occasional along
roads. June-July.

Portulacaceae

Calandrinia ciliata (R.& P.) DC. var. menziesii (Hook.) Macbr. Red
maids. Annual, native. Common in disturbed areas; variable in morphol-
ogy. March-April.

Montia perfoliata (Donn) Howell. Miner’s lettuce. Annual, native.
Common to abundant in grassland; together with Stachys rigida ssp.
quercetorum, forming much of forb growth in grassland in very early
spring. March-April.

Primulaceae

Anagallis arvensis L. Scarlet pimpernel. Annual, introduced. Occasional
to common in grassland, in disturbed areas, and rare at base of ocean-
facing bluff. Flowering sporadic, March-September.

Ranunculaceae
Ranunculus californicus Benth. var. cuneatus Greene. Buttercup. Per-
ennial, antive. Common in grassland and in paths of grassland; petals
all yellow or white-tipped. March-April.

Rhamnaceae

Ceanothus thyrsiflorus Eschs. var. repens McMinn. Blue-blossom.
Perennial, native. Probably planted, near Marine Laboratory; only veg-
etative shoots seen (young plants).

Rhamunus californica Eschs. ssp. tomentella ( Benth.) Wolf. Buckthorn.
Perennial, native. Rare to occasional on steep, wet hillsides. Only vege-
tative shoots seen.

Rosaceae

Fragaria chiloensis (L.) Duchn. Beach strawberry. Perennial, intro-
duced. Despite the name, only seen in grassland; rare. March.

Horkelia marinensis (Elmer) Crum ex Keck. Perennial, native. Rare
about rocks in grassland.

Potentilla egedii Wormsk. var. grandis (Rydb.) Howell. Cinquefoil.
Perennial, native. Abundant in seasonally wet (fresh water) swale
through grassland, also in narrow strip at edge of saline flat near ocean.
Flowering April-July principally, a few flowering to September.

Rosa eglanteria L. Eglantine. Perennial, introduced. Occasional in
erassland away from coast; viciously armed. Only vegetative shoots seen.

Rubus procerus P.J. Muell. Himalaya berry. Perennial, introduced.
Forming tall thickets along shaded stream banks, also as a climber over
shrubs in grassland away from the coast. Occasional. June.

R. spectabilis Pursh. var. franciscanus (Rydb.) Howell. Salmon berry.
Perennial, native. Rare to occasional on steep, wet hillsides. March.

R. ursinus C. & S. California blackberry. Perennial, native. Rare as
climber over shrubs away from coast. April.

R. vitifolius C. & S. California blackberry. Occasional in grassland

1970] BARBOUR: BOGEDA HEAD 309

away from coast and along roads. Perennial, native.

Rubiaceae
Galium asperrimum Gray. Perennial, native, April.

Salicaceae
Salix laevigata Bebb. Willow. Perennial, native. Common in wet,
shaded gulleys. Only vegetative shoots seen.
S. lasiolepis Benth. Arroyo willow. Perennial, native. Common in wet,
shaded gulleys. Only vegetative shoots seen.
S. lasiolepis Benth. var. bigelovii (Torr.) Bebb. Willow. Perennial,
native. Rare on established dunes. Only vegetative shoots seen.

Scrophulariaceae

Castilleja wrightii Elmer. Paintbrush. Perennial, native. Occasional
to common in seasonally wet (but disturbed) areas, also on steep, wet
hillsides; once seen on ocean-facing bluff; bracts and calyx variable in
color (yellow, dull red, bright red). Flowering sporadic, April-October.

Cordylanthus maritimus Nutt. ex Benth. Annual, native. Common in
the higher part of coastal salt marsh.

Mimulus aurantiacus Curt. Bush monkey-flower. Perennial, native.
Common in grassland and along roads away from coast. April-July.

M. guttatus Fisch. ex DC. ssp. létoralis Penn. Monkey-flower. Peren-
nial, native. Occasional in seasonally wet areas along roads. Flowering
sporadic, April-October.

Orthocar pus ertanthus Bench. var. roseus Gray. Johnny-tuck. Annual,
native. Abundant in rare patches in disturbed parts of grassland. April-
May.

Scrophularia californica C. & S. Figwort. Perennial, native. Rare
along roads. May.

Veronica americana Schwein. Speedwell. Perennial, native. Occcasional
near center of fresh-water marsh (with Sparganium eurycarpum). July.

Solanaceae
Solanum nodiflorum Jacq. Nightshade. Annual-perennial, native. Oc-
casional herb with flowering shoot over 2 m tall, along roads and in wet,
shaded gulley. June-July.
Umbelliferae
Angelica hendersonii Coult. and Rose. Perennial, native. Occasional
along ocean-facing bluffs. August-September.
Conium maculatum L. Poison hemlock. Occasional along roads and
in seasonally wet areas.
Daucus carota L. Queen Anne’s lace. Biennial, introduced. Occasional
along roads. July-September.
D. pusillus Michx. Ratlesnake weed. Annual native. Rare in dunes.
April.
Foeniculum vulgare Mill. Sweet fennel. Biennial-perennial, introduced.
Occasional along roads; odor of licorice. July-September.

310 MADRONO [Vol. 20

Heracleum lanatum Michx. Cow parsnip. Perennial, native. Common
on steep, wet hillsides. April.

Oenanthe sarmentosa Presl. Perennial, native. Abundant in narrow
strip at edge of saline flats near ocean, also in fresh-water marsh. May-
September.

Sanicula arctopoides H. & A. Yellow mats. Perennial, native. Occa-
sional in grassland very close to shore. March.

MONOCOTYLEDONEAE

Amaryllidaceae
Allium dichlamydeum Greene. Wild onion. Perennial, native. Rare on
grassland hilltops. June.
A. triquetrum L. Wild onion. Perennial, introduced. Rare in seasonally
wet area near road. March.
Brodiaea pulchella (Salisb.) Greene. Blue dicks. Perennial, native.
Occasional near crests of grassland hills. June-July.

Cyperaceae

Carex barbarae Dewey. Sedge. Perennial, native. Occasional along
road. June-July.

Cyperus eragrostis Lam. Umbrella sedge. Perennial, native. Occasional
in seasonally wet areas near roads, abundant in center of fresh-water
marshes. August-September.

Scirpus americanus Pers. Bulrush. Perennial, native. Common in saline
flats near ocean and in standing brackish water. April-June.

S. cernuus Vahl. var. californicus (Torr.) Beetle. Bulrush. Annual,
natve. Rare in dunes. July.

S. koilolepis (Steud.) Gleason. Bulrush. Annual, native. Common in
clumps at base of ocean-facing bluffs. August.

S. microcar pus Presl. Bulrush. Perennial, native. Largest of the Cype-
raceae; common in fresh-water marshes. July.

Gramineae

Aira carvophylla L. Hairgrass. Annual, introduced. Common to abun-
dant in grassland and in disturbed parts of grassland; contribution to
standing biomass often overlooked because of short size. April-June.

Agrostis exarata Trin. Bent grass. Perennial, native. Rare along
shaded streambank. July.

Ammophila arenaria (L.) Link. Beachgrass. Perennial, introduced.
Widely planted on dunes, probably escaped in other areas; abundant.
Flowering sporadic, along the outer dune in July, variable further inland.

Avena barbata Brot. Wild oat. Annual, introduced. Common along
roads. May.

Briza maxima L. Quaking grass. Annual, introduced. Rare along roads.
June.

B. minor L. Quaking grass. Annual, introduced. Rare along roads.
June.

— II, LI ————

1970] BARBOUR: BOGEDA HEAD Sil

Bromus carinatus H. & M. California brome. Perennial, native. Oc-
casional to common in grassland. May-June.

B. mollis L. Soft chess. Annual, introduced. Rare in disturbed areas.
June.

B. diandrus Roth. Ripgut. Annual, introduced. Abundant in grass-
land, occasional in disturbed areas. April.

Calamagrostis nutkaensis (Presl) Steud. Reedgrass. Perennial, native.
Occasional to common on steep, wet hillsides. November.

Cortaderia selloana (Schult.) Arch. & Graebn. Pampas grass. Peren-
nial, introduced. Possibly planted; near Marine Laboratory; rare. Sep-
tember.

Dactylis glomerata L. Orchard grass. Perennial, introduced. Rare
along roads. June.

Distichlis spicata (L.) Greene var. stolonifera Beetle. Perennial, na-
tive. Salt grass. Common in saline flats near ocean and at base of ocean-
facing cliffs. Only vegetative shoots seen.

Elymus glaucus Buckl. Rye grass. Perennial, native. Rare in wet,
shaded gulley. July.

E. vancouverensis Vasey. Rye grass. Perennial, native. Occasional in
stabilized dunes (with Lupinus arboreus), along roads, and near lip of
ocean-facing bluff. August.

Festuca dertonensis (All.) Arch. & Graebn. Fescue. Annual, intro-
duced. Occasional at edge of saline flat near ocean, with Holcus lanatus.
May.

Holcus lanatus L. Velvet grass. Perennial, introduced. Abundant in
low area of grassland (near Potentilla egedi var. grandis) and on saline
flat near ocean; occasional in disturbed areas of grassland. June-August.

Hordeum brachyantherum Nevski. Perennial, native. Occasional in
erassland. April.

H. depressum (Scribn. & Sm.) Rydb. Wild barley. Annual, native.
Common in grassland, occasional along roads. May.

H. leporinum Link. Farmer’s foxtail. Annual, introduced. Occasional
in disturbed areas. March-June.

Lolium multiflorum Lam. Italian ryegrass. Annual, introduced. To-
gether with Azra caryophylla, Bromus arvensis, Bromus rigidus, makes
up most of the grass cover in grassland. Abundant. May-June.

Poa douglast Nees. Sand bluegrass. Perennial, native. Occasional on
established dunes. March-April.

P. scabrella (Thurb.) Benth. Blue grass. Perennial, native. Occa-
sional in grassland. April.

Polvpogon mons peliensis (L.) Desf. Rabbit-foot grass. Annual, intro-
duced. Occasional in disturbed areas and along base of ocean-facing
bluff, rare along shaded streambank. June-September. Variable morphol-

ogy.

312 MADRONO [Vol. 20

Tridaceae
Tris douglasiana Herb. Wild iris. Perennial, native. Occasional in grass-
land close to and away from the shore. March-April.
Sisyrinchium bellum Wats. Blue-eyed grass. Perennial, native. Occa-
sional in low areas of grassland. April-July.

Juncaceae

Juncus balticus Willd. Rush. Perennial, native. Abundant in occa-
sional patches in wet, low parts of grassland. March.

J. bolanderi Engelm. Rush. Perennial, native. Rare in wet areas along
roads. August.

J. bufonius L. Toad rush. Annual, native. Smallest of the rushes at
the Head; rare in seasonally wet area near road (with Cyperus eragros-
tis. August.

J. effusus L. var. brunneus Engelm. Rush. Perennial, native. Occasional
along road. May-October.

J. leseurii Bol. Rush. Perennial, native. The most wide-spread and
common of the rushes at the Head; common in dunes near a pond, abun-
dant in fresh-water marsh, occasional along roads. May-October.

Luzula subsessilis (Wats.) Buch. Wood rush. Perennial, native. Com-
mon in grassland, especially near crests of hills. March.

Juncaginaceae

Triglochin maritima L. Arrow grass. Perennial, native. Common in

salt marsh. May-June.
Liliaceae

Chlorogalum pomeridianum (DC.) Kunth. Soap plant. Perennial,
native. Common in grassland; only vegetative shoots seen.

Fritillaria recurva Benth. Fritillary. Perennial, native. Rare, at lip of
steep hillside, in grassland. March.

Potamogetonaceae
Potamogeton crispus L. Perennial, introduced. Occasional in fresh
water ponds. Leaves broad. Only vegetative material seen.
P. pectinatus L. Perennial, native. Common in fresh water ponds.
Leaves linear. Only vegetative material seen.

Sparganiaceae
Sparganium eurycarpum Engelm. Bur-reed. Perennial, native. Occa-
sional in center of fresh-water marshes. July-September.

Typhaceae
Typha angustifolia L. Cat-tail. Perrenial, introduced. Occasional in
seasonally wet areas near road (often with Cyperus eragrostis). August-
September.
Zosteraceae
Phyllospadix torreyi Wats. Surf-grass. Occasionally thrown up on
beach from near the low-tide level. Perennial, native.

Department of Botany, University of California, Davis

1970] WEST: HETEROSIPHONIA O15

LITERATURE CITED

Hoover, M.B., H. E. Renscu, and E. G. Renscu. 1948. Historic spots in California.
Stanford Univ. Press.

Howe tt, J. T. 1937. A Russian collection of California plants. Leafl. W. Bot. 2:17—20.

. 1949. Marin flora. Univ. Calif. Press, Berkeley.

Hunt, R. D., and N. Sancuez. 1929. A short history of California. Thomas Y.
Crowell. New York.

Jackson, M. L. 1958. Soil chemical analysis. Prentice-Hall, Englewood Cliffs.

Kinnarb, L. 1966. History of the greater San Francisco Bay region. Lewis Historical
Publishing Co., New York. 3 vols.

Muwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.

THE CONSPECIFICITY OF HETEROSIPHONIA ASYMMETRIA
AND H. DENSIUSCULA AND THEIR
LIFE HISTORIES IN CULTURE

JouHN A. WEST

INTRODUCTION

The marine red algal genus Heterosiphonia (Ceramiales, Dasyaceae)
includes about 40 species which are widely distributed in temperate,
tropical and cold waters. The genus is characterized as having poly-
siphonous and corticated main axes which branch in a sympodial man-
ner. The lateral branches are alternate, distichous and either mono-
siphonous or polysiphonous. Spermatangia and tetrasporangia are borne
in specialized conical reproductive structures called stichidia (Kylin,
1956).

On the Pacific coast of North America five species are known. Heter-
osiphonia densiuscula and H. laxa were described from Friday Harbor,
Washington by Kylin (1925). Both species are known only from
northern Washington and southern British Columbia (Scagel, 1957).
Heterosiphonia asymmetria, described by Hollenberg (1945), has a
range extending from Santa Catalina Island to the Monterey Peninsula
in California (Hollenberg and Abbott, 1966). Gardner (1927) described
H. erecta which ranges from southern California to Baja, California
(Dawson, 1963). Heterosiphonia wurdemanniu Bérgesen is broadly dis-
tributed in tropical waters and is present in the Gulf of California (Daw-
son, 1963).

Heterosiphonia erecta and H. wurdemanni are described as having
four pericentral cells, H. asymmetria as having five, and H. laxa and
H. densiuscula as having six to nine. The first two species appear to be
morphologically distinct taxa, but H. asvmmetria, H. densiuscula and
possibly H. laxa appear very closely related, if not identical, for reasons
which will be brought out in the observations and discussion section of
this paper. Because of the apparent taxonomic problems involving these
three species, I considered it necessary to re-investigate various aspects
of their morphology and life histories.

314 MADRONO [Vol. 20

MATERIALS AND METHODS

For the culture of H. densiuscula, specimens were dredged from about
15 m depth at Partridge Bank, west of Whidbey I., Washington, on July
6, 1965. This clone has been maintained for over four year in unialgal
culture with Provasoli’s enriched seawater medium (Provasoli, 1966)
in 10 C, 16 hr daily photoperiod and 20-40 ft-c cool white fluorescent
lighting. The clone of H. asymmetria was isolated from material col-
lected in the drift at the south end of Carmel Beach, Monterey Penin-
sula, California, April 19, 1967. It has been cultured for more than two
years in 15 C, 15—150 ft-c cool white lighting and 16 hr daily photo-
period with Provasoli’s medium.

All seawater for these culture studies was obtained from the Bodega
Marine Laboratory, Bodega Bay, California. It was aged at least 30
days in the dark at 20-22 C before use. The salinity was adjusted to
30-31% by adding glass distilled water. The seawater was then steam
sterilized for 30 mintues and stored until use. The enrichment medium
was added to the sterile seawater just prior to use.

All cultures were maintained in either Pyrex (No. 3250) 100 x 80
deep storage dishes with 150-200 ml of the medium or in Pyrex 90 x 50
mm crystallizing dishes with 100 ml of the medium.

Tetraspores were allowed to attach to coverslips which were trans-
ferred to fresh medium every 14—30 days.

OBSERVATIONS AND DISCUSSION

Culture studies. Heterosiphonia densiuscula grows well in culture
without significant deviation from the morphology observed in field-
collected specimens. The initially tetrasporophytes produced tetraspores
which gave rise to tetrasporangia-bearing plants after 3 months at about
30 ft-c illumination. The plants in culture are 3.0 to 5.0 cm long when
reproductive, in contrast to plants from nature which often reach 10
to 15 cm in length when reproductive. Neither carpogonia nor sperma-
tangia were observed on any of more than 500 plants from four suc-
cessive generations. The tetrasporangia are normal in morphology and
release 4 spores. Each fertile segment of the stichidium bears 4 to 6
sporangia in a whorl (fig. 2). Spore germination is similar to that de-
scribed for most of the Ceramiales. The spore first divides into two
unequal cells. The smaller cell is the initial of the basal system. It forms
either an elongate multicellular rhizoid or a multicellular lobed at-

Fics. 1-5. 1-4, Heterosiphonia densiuscula; 5, H. asymmetria; 1, 2, same scale;
1, apex of cultured plant, sympcdial, alternate, distichous branching of main poly-
siphonous branch is evident; 2, cultured tetrasporophyte with pedicellate stichidi-
um borne on polysiphonous basal portion of lateral branch; 3, squashed prepara-
tion of type specimen showing branch bearing five pericentral cells enclosing axial
cell (ac) ; 4, young germling in monosiphonous stage, new apical meristem develop-
ing from intercalary cell and multicellular lobed attachment organs are evident;
5, habit photograph of typical specimen collected from same location at same date
as cultured specimen.

HETEROSIPHONIA

WEST

1970]

316 MADRONO [Vol. 20

Fic. 6. Heterosiphonia densiuscula, habit photograph of typical specimen dredged
from Hein Bank, south of San Juan I., Washington.

tachment disk (fig. 4). The larger of the two cells derived from spore
division is the precursor of the erect system. When the primary erect
monosiphonous filament becomes about 1 mm in length, the apical meri-
stem begins to branch sympodially. Additional meristems may be estab-
lished from intercalary cells by an oblique lateral division which pro-
duces a new initial (fig. 4). The meristems give rise to the polysipho-
nous sympodially branched axes of the developing plant (fig. 1).
Heterosiphonia asymmetria cultures also were started from a field-
collected tetrasporophyte and the tetraspores also gave rise to tetra-
sporangia-bearing plants for two successive generations. The cultured
plants were much smaller than plants from nature, rarely exceeding

1970] WEST: HETEROSIPHONIA SU,

1.0 cm in length. The third generation is morphologically aberrant and
does not grow well but it is being maintained for further study. It grows
primarily by proliferation of the uniseriately filamentous basal system
from which occasionally arise erect polysiphonous branches that bear
tetrasporangia.

Heterosiphonia densiuscula cannot tolerate light intensities above
50 ft.-c. Above 75 ft-c the germlings bleach and die within a week. On
the other hand, H. asvmmetria grows and reproduces in intensities from
15 to 150 ft-c.

Morphological studies on field-collected specimens. The types of H.
densiuscula and H. laxa were re-examined and found to have only 5 peri-
central cells per segment instead of 6—9 as recorded in the original de-
scription (Kylin, 1925). Kylin’s own figure (fig. 44, p. 68) shows only
three pericentral cells on one side which is indicative of five or, at the
most, six. Re-examination of H. asymmetria type specimen reveals 5
pericentral cells per segment as indicated in the original description
(Hollenberg, 1945). The pericentral cells of all three species are charac-
teristically arranged so that three are evident in face view on one side
of the branch and two are seen on the opposite side when the branch is
turned over.

Although all three species are similar, an insufficient number of H. laxa
specimens are available for a thorough re-examination of its morphology.
A comparison of the major morphological characteristics of H. densius-
cula and H. asymmetria (table 1) indicates that clear similarities exist.

The reproductive patterns of life histories of these two species also
appear similar. A survey of H. densiuscula from several herbaria shows
that of approximately 110 specimens, 95% bear tetrasporangia and
the remaining 5% are sterile. The original description likewise refers
only to tetrasporangiate plants. Recently, however, male and female
gametophytes were collected by Michael Wynne from Puget Sound in
Washington. Mature cystocarps were present on the female plants in-
dicating that both sexes possess functional sexual structures. The pres-
ence of gametophytes, even though they are extremely scarce, suggests
that this species exhibits two types of life histories in the same locality.
It is possible that two distinct genetic races have developed, one exhibit-
ing the typical sexual red algal life history and the other exhibiting a
non-sexual type. Gametophytes are not yet available for a culture study.

A survey of H. asvmmetria herbarium specimens from the Monterey
Peninsula reveals that 44 of 45 plants examined are tetrasporophytes.
The remaining plant is sterile. To the best of my knowledge gameto-
phytes have been collected only twice. The type specimen is a male col-
lected February 12, 1938 from Corona Del Mar, Orange Co., California.
Two cystocarpic female specimens were collected by E. Yale Dawson
from White’s Cove on Santa Catalina Island, California, October 31,
1948. It is apparent in this case that on the Monterey Peninsula a type

318 MADRONO LVol. 20

TABLE 1. MORPHOLOGICAL COMPARISON OF HETEROSIPHONIA ASYMMETRIA AND
H. DENSIUSCULA.

H. asymmetria H. densiuscula

Main branches

sympcdial, distichous and alternate — +
Main branches (1st & 2nd orders)

lacking pubescence of monosiphonous

branchlets — +
Number of segments between branch 2-3 zy,
Ratio of dimensions of segments Or “tol: x O5.to (5x

as long as broad as long as broad

Number of pericentral cells 5 5
Arrangement of pericentral cells transverse to transverse to

(in side view) slightly asymmetric _ slightly asymmetric
Arrangement of pericentral cells

(in cross section) 3+ 2 3+ 2
Cortication by longitudinal filaments

derived from pericentral cells + +

Stichida on monosiphonous &

poysiphonous branchlets + +
Stichidia pedicellate or sessile = =e
Pedicels monosiphonous or - +

polysiphonous 3-5 4-6
Number of sporangia/fertile segment

in stichidium up to 800 uw up to 800 u
Maximum diameter of main axis up to 15 cm up to 15 cm
Total height of plants intertidal & subtidal
Habitat subtidal

of life history similar to that observed for H. densiuscula occurs. Male
and female plants are reported only from the southern limits of its range
in California which suggests that only certain populations may exhibit
a life history characteristic for the higher Florideophyceae. However,
tetrasporophytes are not known to occur in the same locality.

The tetrasporangia from field-collected specimens of both species
appear normal morphologically and form tetrads of spores but evidently
are mitotic, in most instances, rather than meiotic, because in the cul-
tured clones tetrasporophytes develop repeatedly.

Although the non-sexual type of life history is not common among the
higher Florideophyceae, there is good evidence that some species are
represented only by the tetrasporophytes in nature. Svedelius (1937)
demonstrated that tetrasporangia of Lomentaria rosea are apomeiotic.
Sparling (1961) noted that Halosaccion glandiforme (Gmelin) Ruprecht
and Rhodymenia palmata f{. mollis Setchell and Gardner never or very
rarely have been found to bear spermatia, carpogonia or cystocarps.
She maintained tetraspore germlings of both species in culture for about
2 years, but was not able to observe any reproduction. This suggests,
at least, that heteromorphic gametophytes and tetrasporophytes do not
occur in these species and that mitotic tetraspores are the only means of
reproduction.

1970] WEST: HETEROSIPHONIA 319

On the basis of their morphological similarities and the evident simi-
larities in their life histories H. asymmetria Hollenberg should be con-
sidered a synonym of H. densiuscula Kylin. Heterosiphonia laxa and H.
densiuscula also appear closely related. In addition, as H.laxa is separated
from H. densiuscula only by the comparative sparsity of branching in the
former, it cannot be concluded at this time that it is conspecific. Insuffi-
cient material is available for morphological comparison and no living
plants are available for culture. Moreover, all three phases of H. /axa fre-
quently occur in nature, indicating that it may have a typical sexual
life history.

ACKNOWLEDGEMENTS

I am grateful to George Hollenberg for the loan of the type specimen
of Heterosiphonia asymmetria and to G. F. Papenfuss for obtaining the
type specimens of H. densiuscula and H. laxa from Almborn of the Bo-
tanical Museum of Lund University in Sweden. R. E. Norris, Michael
Wynne, R. F. Scagel, and J. S. Garth made specimens available. Michael
Wynne provided liquid preserved and living male and female specimens
of H. densiuscula. G. F. Papenfuss also offered many helpful comments
on this manuscript.

SUMMARY

Similarities in the morphology and life histories of H. asymmetria and
H. denstuscula indicate that they are conspecific. The two clones studied
in cultures formed successive generations of tetrasporophytes. This cor-
relates well with information on the field collected plants, 95% or more
of which are also tetrasporophytes.

Department of Botany, University of California, Berkeley
| LITERATURE CITED

Dawson, E. 1963. Marine red algae of Pacific Mexico. Part 8. Ceramiales: Dasya-
ceae, Rhodomelaceae. Nova Hedwigia 6:401-481.

GARDNER, N. 1927. New Rhodophyceae from the Pacific coast of North America.
VI. Univ. Calif. Publ. Bot. 14:99-138.

HOLieNBERG, G. 1945. New marine algae from southern California. III. Amer. J.
Bor. 32: 447-451.

.. and I. Aspotr. 1966. Supplement to Smith’s Marine Algae of the Mon-
terey Peninsula. Stanford Univ. Press.

Kyun, H. 1925. The marine red algae in the vicinity of the biological station at
Friday Harbor, Wash. Acta Univ. Lund 21(9) : 1-87.

—. 1956. Die Gattungen der Rhodophyceen. CWK Gleerup Forlag. Lund,
Sweden.

Provasoti, L. 1968. Media and prospects for cultivation of marine algae. Jn
Watanabe, A., and A. Hattori (eds). Culture and collections of algae. Proc.
U.S—Japan Conference Hakone, Sept. 1966, Jap. Soc. Pl. Physiol.

SCAGEL, R., 1957. An annotated list of the marine algae of British Columbia and
northern Washington. Bull. Natl. Mus. Canada 150.

SPARLING, S. 1961. A report on the culture of some species of Halosaccion, Rhody-
menia and Fauchia. Amer. J. Bot. 48:493-499.

SVEDELIuS, N. 1937. The apomeiotic tetrad division in Lomentaria rosea in com-

parison with the normal development in Lomentaria clavellosa. Symbol. Bot.
Upsal. 2(2) :1-54.

A NEW PROSTRATE VARIETY OF ERIOGONUM APRICUM
(POLYGONACEAE)

RopDNEY MYATT

The Ione Buckwheat, Eriogonum apricum Howell, is a narrow endemic
restricted to acidic, kaolinitic, clay soils of the Ione Formation (Allen,
1929) near Ione, Amador Co., California. The original population, dis-
covered by Howell in 1954, is located on an exposed red clay hill about
four miles south of Ione. A second population is located along California
Highway 88, about two miles south of Ione (Roderick, 1964). Recently
a third population was discovered by Gankin and Hildreth in February
1967 about five miles north of Ione, which differs from the previously
known populations in having prostrate stems.

Eriogonum apricum Howell var. prostratum Myatt, var. nov. A var.
apico differt caulibus prostratis, foliis minoribus. Similar to var. apricum
except that the basal rosette is 3—7 cm across, with glabrous prostrate
stems; leaves orbicular to ovate, truncate to slightly cordate, woolly-
tomentose beneath, glabrous above, (3—) 5-6 (—9) mm long (fig. 1).

Fic. 1. Eriogonum apricum: left, var. prostratum; right, var. apricum.

Type. Open areas among Arctostaphylos myrtifolia Parry on red clay
soil, near the Irish Hill Road about 3 miles from the junction with
Highway 104, about 5 miles north of Ione, Amador Co., California, ele-
vation ca. 300 ft., June 20, 1967, Myatt s.n. (DAV-—holotype).

All three known populations of £. apricum are located within exposed
areas of Arctostaphylos myrtifolia Parry vegetation, which is apparently
an azonal vegetation type (Gankin & Major, 1964).

The new variety differs from E. apricum var. apricum in several subtle
but consistent ways. The var. prostratum flowers from mid-June to early

320

1970] PARNELL: CLARKIA 321

July, with the seeds being shed by mid July. The two populations of
var. apricum, however, flower from mid July through September and
seeds are shed from August through late October. The leaves of var.
prostratum are generally smaller than those of var. apricum, the former
averaging 5-6 mm and are often 10-11 mm long. The only anatomical
difference evident is that the cortex cells in the flowering stems of var.
prostratum are of a longer, palisade type, being 2—3 times the length of
those in var. apricum. Greenhouse studies have shown that plants grown
from seeds of var. prostratum retain these distinctive characteristics
under uniform conditions.

I would like to thank Grady Webster for the Latin diagnosis and his
advice in preparing this article, and also John M. Tucker for his help
with the description.

Department of Botany, University of California, Davis

LITERATURE CITED

ALLEN, V. T. 1929. The Ione formation of California. Univ. Calif. Publ. Geol.
18:347-448.

GANKIN, R., and J. Major. 1964. Arctostaphylos myrtifolia, its biology and rela-
tionship to the problem of endemism. Ecology 45:792-808.

Roperick, W. 1964. A new station for Eriogonum apricum. Leafl. W. Bot. 10:136.

CLARKIA JOLONENSIS (ONAGRACEAE), A NEW SPECIES
FROM THE INNER COAST RANGES OF CALIFORNIA

DENNIS R. PARNELL

In the most recent taxonomic discussion of Clarkia deflexa, Lewis and
Lewis (1955) pointed out that this endemic California species “‘shows
considerable variation, particularly from population to population and
to some extent regionally.” A further study of this species (Parnell,
1968) has clarified that, in fact, what has been included under the name
C. deflexa are two morphologically distinct population groups effectively
separated from each other by both geographical and internal barriers
to gene exchange.

The first group of populations is found in the outer Coast Ranges of
California from Orange Co. north to San Luis Obispo Co. Although
there is a high degree of morphological variation between these popu-
lations, hybrids between them are highly fertile (Lewis, 1953; Parnell,
1968).

The second group of populations is known only from the inner Coast
Ranges of Monterey Co. As in the case of the first group, interpopula-
tional hybrids are fully fertile.

Except for two individuals from populations in the outer Coast Ranges
who were heterozygous for a single translocation, all individuals includ-

322 MADRONO [Vol. 20

ing the hybrid progeny regularly formed nine pairs of chromosomes at
meiotic metaphase I.

Intergroup hybrids have been obtained only seven times (four of the
individuals came from the seeds of a single capsule) and only after
numerous pollinations had been attempted. The difficulty in obtaining
these hybrids stems from embryo inviability caused by genetic and pos-
sibly cytoplasmic differences between the two groups. The seeds of the
plants belonging to the two groups are markedly different and provide
the only consistent way of telling them apart. In the Monterey Co.
plants the seeds are heavily covered with scales and appear dark gray.
Seeds from other populations are covered primarily by small! papi-
liform projections and appear black or brown in color.

In view of these morphological differences and the well developed bar-
riers to hybridization, it seems appropriate to recognize the Monterey
County populations as a distinct species.

Clarkia jolonensis Parnell, sp. nov. Herba erecta, altitudine ad 6 dm;
caulibus simplicibus vel ramosis; foliis 2-6 cm longis, 2-5 mm lato; caly-
cis limbo 9-15 mm longo, 2—3 lato; petalis 11-19 mm longis, 9-14 mm
latop stylo 9-14 mm longo, quam staminibus longiore vel longitudine
aequa; semina cinera propter squamas.

Type. California, Monterey Co., 9 mi. N.W. of Bradley along Jolon
Road, June 3, 1963, R. F. Thorne & P. Everett 32156. (DS, LA-holo-
type).

Specimens examined. Monterey Co: Road to Pleyto, 0.4 mi. south of
Bradley-Jolon Road, Lewis & Epling 192 (LA); 3.1 mi. north of San
Antonio Road, Lockwood-San Lucas Road, Hardham 4318 (LA); Shale
Hills, w. side Hames Valley, Jolon-Bradley Road, Hardham 1299 (LA);
Mill Creek, road to Adler Creek, Hardham 6049 (LA); foot of grade
to King City, Dudley (DS).

Most populations of C. jolonensis are uniform in appearance except
for one population located 9.1 mile east of the San Lucas Road turnoff
in Lockwood. When compared with other populations, the seedlings and
adult plants of this population tend to be smaller. The petals are 3-8
mm shorter in length and 3—5 mm narrower in width as well as being
a much more pale pink. The style is shorter than the stamens and very
seldom has it been observed to fall clear of them. Since pollen is being
shed at the time the stigma becomes receptive some degree of self-
pollination undoubtedly occurs. This is indicated by the full seed set
that is obtained when plants from this population are left unattended
in the greenhouse. Although this does not exclude a certain amount of
outcrossing, it does indicate that self-pollination is probably the norm
for the population in contrast to all other populations which ordinarily
set no seed under similar conditions and show no sign of fertilization
(e.g., swelling of the ovary accomplished by abscission of the flower.)

Department of Biological Science, California State College, Hayward

1970] KOWALSKI: LAMPRODERMA 625

LITERATURE CITED

Lewis, H. 1953. The mechanisms of evolution in the genus Clarkia. Evolution
7:1-20.
. and M. E. Lewis. 1955. The genus Clarkia. Univ. Calif. Publ. Bot.
20:241-342.
PaRNELL, D. 1968. Reproductive barriers in Clarkia deflexa. Brittonia 20:387-394,

CONCERNING THE VALIDITY OF LAMPRODERMA
ECHINOSPORUM

DONALD T. KOWALSKI

In 1924 Meylan described Lamproderma echinosporum on the basis
of several collections from the Jura Mountains of Switzeralnd. This
species was a typical snowline Myxomycete, i.e., found only at high ele-
vations near the melting snow. Lister (1925) did not mention this taxon
in her monograph, probably because it was described too late to be in-
cluded in her work. In 1924 Macbride and Martin recognized the species
as valid, but apparently did not have any material for observation.
Dennison (1945), however, placed L. echinosporum under the heading
of doubtful species. She had no material for study and on the basis of
the description, she thought it was very close to L. echinulatum (Berk.)
Rost. Hagelstein (1944) and Martin (1949) did not include it in their
monographs because it was not reported from North America. During
my work in the western United States, I have made five collections of a
species which does not fit any of the generally recognized taxa, but
which match perfectly with five of Meylan’s collections of L. echinos-
porum. These five collections were obtained on loan from the Musée de
Botanique, Lausanne, Switzeralnd. I believe that L. echinosporum is a
good species and my collections (3601, 3668, 6240, 8284, 82856) ap-
parently represent the first time that this taxon has been reported from
the Western Hemisphere. All of my collections are from northern Cali-
fornia and 8284 has been deposited in the University of Iowa Herbarium.

Meylan did not specifically designate a type collection and his species
diagnosis was, in my opinion, incomplete. Thus, I am designating his
April, 1923 collection from Prise as the lectotype and including a de-
tailed English description with accompanying paragraphs dealing with
the majoror characteristics and relationships of this taxon.

LAMPRODERMA ECHINOSPORUM Meylan. Sporangia (fig. 1) scattered
to loosely clustered in small groups of 3—6, sessile or briefly stipitate,
broadly ovoid to occasionally globose, 1.0-1.5 mm in diameter, color
variable, dark brown to blue, dull, occasionally slightly iridescent;
stipes, when present, short, averaging about 1.0 mm in length, shiny
brownish-black; peridium membranous, thin, usually long persistent,

324 MADRONO [Vol. 20

Fics. 1-4. Lamproderma echinosporum. 1, sporangia, X 30; 2, spores, X 1130;
3, columella and capillitium x 50; 4, spores, & 1740.

splitting irregularly, covered with numerous depressed, dark brownish-
black areas, giving it a mottled appearance, depressed areas circular to
slightly elongated, few near the base of the sporangium, becoming plen-
tiful in the upper half; hypothallus poorly developed, discoid, thin, trans-
parent, often merging at the margins into adjacent hypothalli, reddish-
brown; columella (fig. 3) black, tapering only slightly towards the
truncate or bluntly pointed apex, usually attaining two-thirds the height
of the sporangium, often branching at the apex where it forms the

1970] KOWALSKI: LAMPRODERMA 625

primary branch of the capillitium; capillitium (fig. 3) forming a dis-
tinct, dense net, with abundant free ends, dark brown throughout,
becoming hyaline only at the extreme tips, arising predominantly from
the apex of the columella, threads of predominantly uniform thickness,
2—4 w in diameter, usually not expanded in the axils, often covered with
numerous nodules, averaging 5—10 » in diameter; spores (figs. 2, 4)
globose, dark brown in mass, dusky brown by transmitted light, cov-
ered with large irregularly distributed spines, often reaching 1.0 p» in
length, including ornamentation, 13-16 » in diameter; plasmodium
unknown.

This species is known only from Switzerland and California. All of
the Switzerland collections examined appear to be on herbaceous plant
debris while the California collections are all on fallen coniferous twigs.

There is no characteristic that can be used to identify this species in
the field. Typically, it forms small fruitings, consisting of only 20-30
sporangia. Small fruitings, however, can be found in several other
species of Lamproderma. There is, however, one distinctive feature
which makes L. echinosporum very easy to identify under the stereo-
scopic microscope, and that is the presence of the dark depressed areas
on the peridium. It is interesting that Meylan did not mention the pres-
ence of this characteristic in his diagnosis and as far as I can determine,
it has not been mentioned by anyone in the literature. In 1919, five years
before describing L. echinosporum, Meylan described L. gulielmae from
the mountains of Switzerland. This is the only other species in the genus
that also has the dark depressed areas on the peridium. I do not believe,
however, that these taxa are closely related. There are many differnces
which distinguish them. In L. gulielmae the sporangia are small, 0.3—0.5
mm in diameter, the peridium is silvery-blue, the stipe, in relation to
the size of the sporangium, is long, about four times the diameter of the
sporangium, and the capillitium forms a lax net which is distinctly
hyaline or pallid. In L. echinosporum the sporangia are large, 1.0-1.5 mm
in diameter, the peridium is dark brown or blue, the stalk, in relation
to the size of the sporangium, is short, about equal to, or less than, the
diameter of the sporangium, and the capillitium forms a dense net which
is dark brown.

Although Meylan (1924) thought ZL. echinosporum was related to
as L. atrosporum Meylan, and Dennison (1945) believed it was near L.
echinulatum, I believe L. echinosporum is probably most closely related
to L. sauteri Rost. These taxa share many characteristics in common.
Both have large sporangia, 1-2 mm in diameter; short stalks, 1.0 mm
or less in length; peridia that are dull, rarely iridescent; capillitia which
form dense dark brown nets; and spore sizes which overlap, 13-16 p» in
L. echinosporum and 12-15 p» in L. sauteri. There are, however, signifi-
cant differences between these two species. In L. echinosporum the
peridium is covered with dark depressed areas while L. sauteri lacks

326 MADRONO [Vol. 20

this feature entirely. The spores of L. echinosporum are uniformly
dusky in color and covered with irregularly distributed spines which
may reach 1.0 » in length, while in L. sauteri the spores are dark violet
brown and distinctly lighter on one side, and regularly and densely
spinulose, the spines attaining only 0.5 » in length.

This study as supported by the National Science Foundation (Grant
GB-5799).

Department of Biology, Chico State College, Chico, California

LITERATURE CITED

Dennison, M. L. 1945. The genus Lamproderma and its relationships. I. Mycologia
37:80-108.

HacELsTEIN, R. 1944. The Mycetozoa of North America. Mineola.

Lister, A. 1925. A monograph of the Mycetozoa. 3rd ed. Revised by G. Lister,
Brit. Mus. Nat. Hist. London.

MacpripkE, T. H., and G. W. Martin. 1934. The Myxomycetes. MacMillan.

Martin, G. W. 1949. North American Flora 1:1—-190.

Meytan, C. 1919. Notes sur quelques especes de Myxomycetes. Bull. Soc. Vaud.
Sci. Nat. 52:447-450.

. 1924. Recherches sur les Myxomycetes du Jura en 1921-22-23. Bull. Soc.

Vaud. Sci. Nat. 55:237-244.

PERENNATION IN ASTRAGALUS LENTIGINOSUS AND
TRIDENS PULCHELLUS IN RELATION TO RAINFALL

JANICE C. BEATLEY

In late winter-early spring (March 15—April 12) of 1965, at a season
when precipitation is not predictable, rains of extraordinary frequency
and magnitude fell over most parts of south-central Nevada. Rainfall
during this period on the Nevada Test Site, Nye Co., was from 2.5 to
more than 5 inches at elevations below 5,000 feet, where the communi-
ties and flora belong to the Mojave Desert (Beatley, 1969). Late autumn
rains the same year were even more extraordinary, and for the 1965
calendar year precipitation totalled 8 to more than 15 inches over the
Test Site, with many areas receiving in excess of 10 inches.

A number of biological phenomena following the spring rains, and in
the 1966 spring season following the autumn rains, were as exceptional
as the rainfall which preceded them. Among these was the appearance
in certain areas of conspicuous numbers of seedlings of Astragalus len-
tiginosus Dougl. var. fremonti (Gray) Wats. (Leguminosae) and T7i-
dens pulchellus (HBK.) Hitchc. (Gramineae) in the spring of 1965,
and spectacular flowering populations of the Astragalus in the spring of
1966. On permanent study sites located in these areas, year-round envi-
ronmental measurements and plant data collections in the spring of each

1970] BEATLEY: PERENNATION 327

year enabled a history of the populations to be quantitatively document-
ed in relation to certain environmental variables.

It is the objective of this report to record the fate of the Astragalus
seedlings on five sites, and 7ridens on one site, as a contribution to un-
derstanding of the annual-biennial-perennial habit in these, and perhaps
other desert species, in relation to precipitation.

METHODS

Four of the sites (Plots 16, 17, 66 and 67) were located on the upper
bajada of eastern Jackass Flats, a major drainage basin in the south-
western part of the Test Site and at the northern edge of the Mojave
Desert. Vegetation was Larrea-Lycium andersoniu-Grayia, which charac-
terizes the high Mojave Desert vegetation of the region (Beatley, 1969).
Soils were predominantly sand and essentially without desert pavement.
The sites had an altitudinal range of around 500 feet. Plot 16 was about
three miles downslope from Plots 66 and 67, with Plot 17 about half-way
between. Plots 66 and 67 were at the divide between Jackass Flats and
Frenchman Flat on the undisturbed remnant of a townsite (Wahmonie)
laid out in the 1920’s; the former was in undisturbed shrub vegetation,
and on the latter, in an adjacent clearing for a street of the townsite,
there were occasional scattered shrubs of the same species.

On a fifth site (Plot 42), in Mid Valley to the north of Jackass Flats,
the Coleogyne shrub cover had been destroyed by fire in 1959, and the
site was essentially without shrub vegetation during the period of this
study. Elevation was over 400 feet higher than that of the Wahmonie
sites, and nearly 1000 feet higher than the lowest of the Jackass Flats
plots. Desert pavement of the soil surface was in a disturbed condition.

The sixth site (Plot 36) was on the bajada below the Ranger Moun-
tains in southeastern Frenchman Flat, where the vegetation was Larrea-
Atriplex-Lycium shockleyi, and the well developed desert pavement was
typical of that below limestone mountain ranges of the region.

In May 1965, seedlings of Astragalus were counted in 50 0.1 m?
quadrats, and the number/m‘? calculated. Data for Astragalus or Tridens
plants intercepting 11 permanent parallel transect lines, 100 feet long
x inch wide and 10 feet apart, were recorded in late May or early June
of consecutive years. For each intercepting plant were recorded: 1, the
locations of the beginning and ending intercept (to tenths of feet) along
each of the 100-foot lines; 2, height, measured to the nearest inch;
3, whether living or dead; and 4, if living, whether flowering, fruiting,
or vegetative only. From interception values for the total 1,100 feet of
transect percentage cover of the soil surface was calculated for the
species.

Rainfall was recorded year-round on each site, using an 8-inch funnel
feeding into a two-liter bottle buried below the soil surface. Bottles
were emptied weekly, and the measured milliliters converted to inches
of precipitation.

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328

1970] BEATLEY: PERENNATION 329

ASTRAGALUS LENTIGINOSUS

Astragalus lentiginosus var. fremonti is the common Astragalus of
the region, where it is usually associated with sandy soils derived from
volcanic rocks. In extreme western Jackass Flats, at 3,000-3,300 feet
elevation, it is entirely replaced by var. variabilis Barneby on deep
sands of the bajada below the volcanic Calico Hills. Where soils are
derived from limestones and dolomites, the common perennial Astragalus
is A. tidestromiu (Rydb.) Clokey.

Seedlings of var. fremontii were conspicuous, though scattered, and
rather uniformly spaced in the spring of 1965; density was 3-7/m? on
the five sites under consideration. On all of the plots there was an occa-
sional plant that was not a seedling, but none intercepted the lines at
time of the 1965 spring data collections. Apparently all, or nearly all, of
the seedlings survived the relatively dry summer and autumn, and fol-
lowing the heavy late autumn rains grew uniformly and rapidly, espe-
cially during late winter. By mid-April 1966 nearly all plants were flow-
ering profusely, and by early May were in abundant fruit; plants were
commonly a foot or more in diameter, and averaged 7-8 inches high.
Over several square miles of eastern Jackass Flats and the larger “burn”
of northern Mid Valley, flowering and fruiting populations were the
highly colorful landscape feature noted by Barneby (1964) to be char-
acteristic of this variety of A. lentiginosus.

In Table 1 are the quantitative data for the populations on each of
the five sites, for the 1966 and following spring season. It is apparent
from the data that numbers (and size) of plants, and total cover of the
soil surface in 1966 were remarkably similar on the Jackass Flats plots.
The Mid Valley population was about half the size of the populations in
Jackass Flats.

Percentage survival of individuals from 1966 to 1967 in Jackass Flats
was in a gradient upslope, and was directly correlated with a precipita-
tion gradient from the lower to the higher elevations. Essentially none
survived at the lowest elevation, where there was the least rainfall during
all periods. Nearly all survived on the Mid Valley site, at the highest
elevation and where rainfall was consistently the greatest from time of
germination (and plants in 1967 were commonly two feet or more in
diameter). There were somewhat fewer and smaller plants (average 6.6
inches high) in 1966 on the disturbed Plot 67, as compared with the
undisturbed Plot 66 (average height 7.8 inches), but the percentage sur-
viving until the next year was somewhat higher on the disturbed site.

The large majority of plants, which germinated in the spring of one
season and flowered and fruited the next spring season, and also died
the second season, were distinctly biennials; the annual, biennial, and
short-lived perennial habit are all ascribed to this variety of A. lenti-
ginosus by Barneby (1964). Whether populations or individual plants
are biennial or perennial appears from the data to be under the control

330 MADRONO [Vol. 20

of soil moisture. The large flowering populations, which appear in the
spring on the basin floors of southern Nevada and then vanish for a
period of years, apparently are associated with the occurrence the year
before of an exceptionally heavy rain, at a time when temperatures were
not limiting to germination, and most of the plants were destined from
time of germination to be biennials; the number surviving on a given
site until the next season is related to the local (and probably micro-
variations in) soil moisture regimes. The predominantly perennial pop-
ulations are those of the higher elevations (4,500-7,500 feet on the Test
Site), where there is more frequent and greater rainfall, and greater
precipitation effectiveness at the lower prevailing temperatures; even
these are usually short-lived, though occasional plants may persist for
several years.

It is possible that the rain “triggering” mass germination may also be
autumn precipitation, when at least one heavy rain is predictable in this
region. Mass germination of this species in the autumn, however, has
not been observed at the Test Site, although occasional seedlings com-
monly occur with the winter annual seedlings after autumn germination.
In those flowering the next spring but not surviving to the following year,
the life cycle is identical to that of the winter annuals.

Variety variabilis, the representative of the species in western
Jackass Flats, also apparently germinated following the early spring
rains of 1965, but flowered and fruited into mid-summer of the same
season, and died during the summer. Perennial plants are scattered in
this area, but the largest flowering populations, at least in this case,
exhibited an annual growth habit. The variety is described by Barneby
(1964) as a perennial of short duration or monocarpic.

TRIDENS PULCHELLUS

This low, tufted perennial grass, of usually calcareous soils, germi-
nated in large numbers in Frenchman Flat following the 1965 spring
rains. In Table 2 are the data for this species on one site for four years
along the same 1,100 feet of transect. The seedlings matured, though
most of the plants at maturity were less than an inch in diameter, and
flowered and fruited abundantly through the late spring of 1965. In
the spring of 1966, three-fourths of the plants yet present were dead;
apparently 15-20 percent of the 1965 plants survived as perennials
through 1966 and 1967. In the 1965 population, around 80 percent
behaved as annuals, with a life cycle of several weeks only.

On another site (Plot 33, on the east slope of Frenchman Flat), there
were 427 plants in 1967 along the 1,100 feet of transect (where there
had been only 13 plants in 1966). The plants germinated following a
mid-September rain, 1966, which was locally 1.5 inches on the east slope
of Frenchman Flat, and only 0.5 inch or less on Plot 36 and elsewhere
on the Test Site. Seeds of this species, therefore, may germinate either
in early spring or early autumn. Data were not collected again in 1968,

1970] BEATLEY: PERENNATION 33

TABLE 2. NUMBER OF PLANTS OF TRIDENS PULCHELLUS INTERCEPTING 1,100 FEET
OF LINE, PERCENT COVER, AND PRECIPITATION, PLOT 36, SOUTHEASTERN FRENCHMAN
Frat, Nevapa Test Site, Nye Co., NEVADA (3,085 feet elevation). 1964-1967.

1964 1965 1966 1967

Number of plants 28 218 125 42

% apparently dead ss 0.0 74.4 0.0

% flowering/ fruiting — 99.1 25.6 81.0

Percentage cover 0.5 223 1.4 0.7
Precipitation (inches)

Mar—June 1965 (germination through flowering/ fruiting) 2.64

July 1965—June 1966 6.64

July 1966-June 1967 5.60

but observation indicated that living plants were rare in the spring of
1968 here and elsewhere on this slope of Frenchman Flat, and for the
most part the plants had exhibited a winter annual growth regime.

DISCUSSION

In Astragalus lentiginosus and Tidens pulchellus it appears the large
and importantly reproducing populations in the northern Mojave Desert
are biennials or annuals, which germinate following unusually heavy
rainfall in the spring or autumn. Only a limited number successfully per-
ennate where precipitation is irregular and variable from season to
season. In these environments, those that do become perennial plants
flower and fruit during the years between large germinations and give
continuity to the presence of the species in the community (of poten-
tial significance especially to any dependent consumers in the commu-
nity). The large perennial populations of Astragalus are confined to the
higher valley floors and mountains, where there is relative constancy
of precipitation above a minimum necessary for the perennial habit.
Tridens does not occur at the higher elevations in this region, and the
perennial populations are those which survive the vagaries of precipita-
tion at the lower elevations.

A potential in higher-elevation perennials for the biennial or annual
life cycle at lower elevations could be expected to enable such species to
occur over a greater altitudinal range, and hence belong to a greater
diversity of plant communities in desert regions, if variables other than
rainfall are not limiting at the lower elevations. Astragalus lentiginosus
var. fremonti, in fact, is nearly unique in the herbaceous perennial flora
of the Test Site region for its altitudinal range from the lowest to the
highest elevations (3,000—7,500 feet), and its occurrence to a greater
or lesser degree in nearly all kinds of plant communities, except where
soils are highly calcareous. The diversity of communities in which it
occurs—from Larrea—Franseria to Artemisia — Pinyon — Juniper — is
matched only by that of Orvzopsis hvmenoides (R. & S.) Ricker.

332 MADRONO [Vol. 20

The facultative life cycle may characterize a number of other desert
herbaceous species usually considered perennial, but which perhaps may
in fact flourish intermittently as biennials or annuals during periods
following extremes in amount or timing of rainfall. Other perennials in
the Test Site flora suspect of having a facultative life cycle are especi-
ally the species of Sphaeralcea, Mirabilis pudica Barneby, and Eriogonum
inflatum T. & F., in which large and conscpicuous populations one year
are often absent the following season.

Work performed under Contract No. AT(04—1) Gen—12 between the
U. S. Atomic Energy Commission, Division of Biology and Medicine,
and the University of California.

Laboratory of Nuclear Medicine and Radiation Biology,
University of California, Los Angeles

LITERATURE CITED

BarneBy, R. C. 1964. Atlas of North America Astragalus. Mem. New York Bot.
Gard. 13(2).

BEATLEY, J. C. 1969. Vascular plants of the Nevada Test Site, Nellis Air Force
Range, and Ash Meadows (northern Mojave and southern Great Basin deserts,
south-central Nevada). UCLA 12-705. Lab. Nucl. Med. & Rad. Biol., Univ.
Calif. Los Angeles.

REVIEWS

Principles and Methods of Plant Biosystematics. By Otto T. Sorpric. xii + 226
pp. MacMillan Company, New York. 1970. $9.95.

Solbrig states that his objective has been “to present the theoretical and tech-
nical aspects of systematics that are not adequately covered in most of the presently
available text-books.” Within his self-imposed limits and the limitations of space
(possibly imposed by the publisher) he has succeeded remarkably well. As reflected
in the title and the quoted excerpt from the preface, the book provides a synopis
of current principles and methods used by practicing biosystematists. The book is
in two parts. The first summarizes the current rationale behind biosystematic re-
search; the second briefly reviews various “modern” data-gathering techniques
employed by biosystematists. The latter section is well suited to development of a
series of laboratory exercises to complement the discussions in the former section.

Organization of material, format, illustrations, etc. are very good and the
breadth of treatment of particular topics is generally uniform and adequate for
elementary students. For advanced classes the discussions form a sound base from
which more thorough analyses of particular principles or methods may be developed.
The following list of chapter headings indicates the scope of the text: Part I—
Introduction and Historical Background, Synthetic Theory of Evolution, Patterns
of Phenetic Variability, Breeding Systems, Speciation, Hybridization, and The
Species Problem and Classification; Part II—Genetics, Cytology, Chemistry, Mathe-
matics and Statistics, and Conclusion.

Most of the discussions of theoretical points are clear, concise, and well supported
through reference to published work. There are, however, several distressing syn-
tactical monstrosities which should never have reached the printed page.

1970] REVIEWS O00

“Under such conditions the plants in each population that are most dissimilar in
their requirements can grow where a minimum number of plants of the other popu-
lation can grow.” p. 85.

“Biological phenomena are never undimensional.” p. 109.

“The important thing is not to make biosystematic or chemical conclusions in-
consistent with the data at hand.” p. 162.

“|. what constitutes a ‘character’ is therefore somewhat irrelevant to the
problem at hand. What is important is to be sure when establishing relationships
that comparable characters are considered.” p. 183.

Further, there is an alarming number of spelling errors and minor errors of fact,
e.g., “annus” for annuus (pp. 7 & 8), “6(107) — 640” (P) (p. 21), “subtrite for
subtribe (p. 38), “Macmillan” for McMillan (pp. 41-43—the work of the pub-
lisher?), “chrysosthoma” for chrysostoma (p. 112), herbaceous Baptisias are said
to be shrubby (p. 164), “betalins” for betalains (p. 164 and glossary). It is hoped
that such errors will be corrected in future printings.

It is unfortunate that in such a small book several pages are wasted in dupli-
cating material. An adequate page of Contents is followed by four pages of Detailed
Contents. “Genetic system” is defined in the text on p. 49 and again in a footnote
on p. 157. The glossary of some twelve pages could easily have been left out and
needed definitions made parenthetically. “Chromatin” is defined both in the glossary
and parenthetically in the text (p. 143). Additionally, many of the glossary entries
are more misleading than informative: An “achaene” (sic) is described as dehiscent
while no mention of dehiscence is made in the entry for “capsule”. “‘Chiasma’’ is
said to be “an exchange of partners .. .”’. “Relationship. A statement about two or
more objects that is either true or false.”

While the general coverage of topics germane to biosystematics is quite good,
there are a few conspicuous omissions. There is no discussion of apomixis even
though this phenomenon is of major importance in the biosystematics of many
genera of flowering plants. Ecotypic variation is very well treated but there is no
mention of clinal variation. Finally, perhaps a minor point, there is no mention
of the importance of voucher specimens for documenting biosystematic research.

Apart from the points raised above, I feel that the author has realized his
objective. No other textbook approximates to such a neat synopsis of current
thought and method in today’s biosystematics. Solbrig is to be congratulated for
recognizing an empty niche and capably filling it. It will be interesting to see
whether his text will succeed or be displaced by competitors, which are sure to
come.—JOHN L. StroTHER, Herbarium, University of California, Berkeley.

The Native Cacti of California. By LYMAN BENSON. xii + 243 pp., illus. Stanford
University Press. 1969. $7.95.

Lyman Benson, leading specialist in the taxonomy of the cacti of the United
States and Canada, has produced a well-organized and thoroughly illustrated book
on the more than 50 different taxa of this family that occur naturally within Cali-
fornia. This compact publication will appeal to a broad range of readers.

A two and one-half page key introduces the nine genera that are covered. Each
genus is treated in detail with a description and, where needed, a key to the species
and varieties. The individual taxa are described and their distribution within the
state is mapped; nearly all are also illustrated, often more than once.

Four color plates by L. C. C. Krieger are superb; the colors, shading, detail, and
accuracy may well be the best that have been employed to illustrate cacti. Also

334 MADRONO [Vol. 20

very good are the black and white photographs by David Griffiths and the line
drawings by Lucretia B. Hamilton. Some of the color photographs taken in the field
are less inspiring but are nearly always helpful for identification. Most of the
legends are informative; but a few would have been more useful if the taxonomic
significance of the flower parts had been commented on, rather than the names of
these parts merely pointed out. In one unfortunate instance there are eight pages of
color plates inserted between a photographic figure and its legend.

Some of the distributional information is, of necessity, incomplete. The author
has constructed distribution maps primarily on the basis of herbarium specimens
examined; however, as cacti are seldom collected by most botanists and only rarely
collected in large duplicate sets, the herbarium record often leaves a part of the
distribution undocumented. For example, the distribution of Opuntia prolifera
actually includes four other Southern California off-shore islands in addition to the
three indicated. Further, O. oricola and O. littoralis var. littoralis occur on all the
islands of this group; and the distribution of O. phaecantha var. major, as it is
designated in this publication, extends into the southernmost portion of Santa Bar-
bara County rather than merely along its northern fringe as shown. It should also
be noted that this latter taxon is the only native flat-jointed Opuntia known from
San Luis Obispo County. The mapping of O. 1. var. littoralis in this county is in
error.

With the anticipated publication of his more technical and comprehensive Cacti
of the United States and Canada, the present book is, in many ways, intentionally
popular. In deference to the layman, the descriptions have been written in non-
metric terms; but many readers will find fractions of an inch, such as 1/128, more
awkward than decimal equivalents in millimeters.

In addition to the features already mentioned, this book contains the following
special sections: a taxonomic summary inside the front and back covers, a long
general introduction, a historical section, and a pertinent biographical list. The
taxonomic summary serves as a convenient index and outline of general classification
and distribution; it is surprising to note that within California nearly as many
kinds of cacti occur in the chapparal as in the Mojave Desert. Also of interest is
the large number of plant communities in which one can expect to find the beaver-
tail cactus, Opuntia basilaris var. basilaris.

The first 60 or so pages form a relatively elementary introduction that is con-
spicuously different from the rest of the text. This introductory portion contains
general botanical information on structure, identification, classification, nomencla-
ture, distribution, climate, paleobotany, and vegetation types. Such material will
be an aid primarily to the beginning student or amateur cactus enthusiast, but
everyone will appreciate the characteristically handsome photographs with which
these concepts are illustrated.

The historical section was contributed by David L. Walkington and is extremely
interesting and pertinent in a plant group where man has played such a significant
role in altering recent evolution. This chronological account of cacti in California
should, however, have begun with Portola’s observation of Opuntia along the Cali-
fornia coast in 1769, a date which is 24 years earlier than the first event considered
here.

The taxonomic portion of the book brings together a number of relatively recent
name combinations at the generic, specific, and subspecific level. A few readers,
especially those who are familiar with the author’s conservative point of view, may
be surprised at the recognition of three genera, Sclerocactus and Neolloydia (segre-
gates of Echinocactus) and Coryphantha (a segregate of Mammillaria).

On the whole, this book is an admirable combination of convenience, attractive-
ness, and completeness—a book that will be used by cactus enthusiasts, students,
and amateur and professional botanists——RatpH N. Purvprick, Santa Barbara
Botanic Garden.

1970] REVIEWS 335

Modern Methods in Plant Taxonomy. Edited by V. H. Heywoop. xvi + 312
pp. Academic Press, London and New York. 1968. 84s.

This collection of papers is one of the best surveys of current trends in plant
taxonomy published to date. The papers were presented at the conterence on
Modern Methods in Plant Taxonomy which was held at the University of Liverpool
in September of 1967. Included in this volume, in addition to the introduction by
V. H. Heywood, are 16 papers grouped somewhat arbitrarily under the following
four headings: The Continuing Role of the Modern Herbarium in Taxonomic
Research; The Role of Experimental Data; Biochemistry, Computers, and Taxon-
omy; and Geography and Ecology.

In general, the various authors reviewed recent developments, discussed particu-
lar problems, and presented original work in areas of interest to the plant syste-
matist. However, not only do these papers form a comprehensive discussion of
Modern Methcds in Plant Taxonomy, but when viewed collectively they also
emphasize the various philosophical differences which exist within the field. One
has only to read, for example, the papers by Cronquist, Solbrig, and Johnson and
Holm, to detect that the meaning or value assigned to such concepts as “phylo-
genetic classification” and “biological ‘species,’ are quite different, if not daia-
metrically opposed. Nevertheless, this dimension of the book should not obscure
its basic importance, an excellent review and analysis of modern plant taxonomy
as well as an indication of its future directions—DeENNIS R. PARNELL, Department
of Biological Science, California State College, Hayward.

Pacific Northwest Ferns and Their Allies. By THomas M. C. Taytor. x + 248
pp., illustrated. Univ. of Toronto Press, Toronto and Buffalo. 1970. $15.00.

Renewed interest in pteridoyphytes during the past two decades has resulted
in an eminently more natural classification system. A consequence of this research
is the outdating of older floristic treatments. Taylor’s book on the Pacific North-
west ferns and their allies helps fill this gap by bringing together our present knowl-
edge of these plants in a well-documented but aesthetically pleasing format.

By the author’s estimate, about one-quarter of the known species of pteri-
dophytes occurring on the North American continent north of Mexico are to be
found in the area treated (Oregon, Washington, British Columbia, and Yukon Ter-
ritory, and Alaska excluding the Aleutian Islands). Not surprisingly, 45% of the
97 species Taylor records for the area are circumboreal.

Keys and concise descriptions are provided for families, genera, and species.
Synonymy, although not complete, is adequate for the purposes of a flora. Refer-
ences are given to earlier floristic treatments and illustrations. Habitat and range
are given for all species. Brief comments include mention of diagostic features,
cytological data, and taxonomic problems requiring additional study. Full page
original line drawings, showing both habit and details, are provided for each
species; although somewhat stylized, these are entirely adequate in most cases for
identification. Chromosome numbers (with references) and lists of species grouped
by distributional patterns are listed in appendices. That the pteridophtes of the area
are exceptionally well studied cytologically is evident from the fact that only 18
taxa remain uncounted.

The author wisely relies on the judgments of monographers in treating many
of the critical genera, including Pellaea, Woodsia, Cystopteris, Botrychium, and
Equisetum, while incorporating the most recent biosystematic data available in
the treatment of such genera as Polystichum, Dryopteris, and Polypodium. He
recognizes such “splinter” genera as Mecodium (Hymenophyllum) and Aspidotis
(Cheilantes), but maintains Thelypteris and Lycopodium in a broad sense.

336 MADRONO [Vol. 20

The book is written both for the professional botanist and for the amateur
fern enthusiast. The latter will find the brief introduction and glossary helpful.
However, common names are not given, nor is reference made to the culture of
these ferns, many of which would make attractive additions to a garden of native
plants.

The book is nearly free of both typographical and factual errors, and those that
were found do not detract from the usefulness of the flora. One possible source of
confusion is the placement of unlabeled distribution maps of some species on pages
where another taxon is treated.

Pacific Northwest Ferns and Their Allies will doubtless be a useful reference
for identification and a source of information for many years to come. One suspects
that only its price might preclude its reaching a wider audience——ALANn R. SMITH,
Department of Botany, University of California, Berkeley.

NOTES AND NEWS
NEW PUBLICATIONS

Vascular Plants of Wells Gray Provincial Park and Its Vicinity, in Eastern
British Columbia. By Leena HAMeEtT-AnTI. Annales Botanici Fennici 2:138-164.
1965. This paper will be of interest to anyone concerned with the flora and plant
geography of British Columbia.

Flora of Montana. Part II. By W. E. Booru and J. C. WricutT. 305 pp. Depart-
ment of Botany, Montana State University, Bozeman. 1966. This part of the
Flora of Montana contains the Dicotyledons and is a corrected and amended ver-
sion of the first edition of 1959.

The Redwood Trail. By P. H. Brypon. 24 pp., illustrated. Strybing Arboretum
Society, Golden Gate Park, San Francisco. 1963. This is a guide to the redwood
trail of the Arboretum.

A Monograph of Lemnaceae. By Epw1n Horace Dauss. Illinois Biological
Monographs 34: vii + 1-118. University of Illinois Press, Urbana. 1965.

Ecology of Soil-Borne Plant Pathogens. Edited by KENNETH F. BAKER and
WittiAmM C. Snyper. xlii + 571 pp. University of California Press, Berkeley and
Los Angeles. 1965.

Plants in Perspective, A Laboratory Manual of Modern Biology. By Etpon H.
NEWCOMB, GERALD C. GeERLOFF, and WILLIAM F. WHITTINGHAM. 218 pp. W. H.
Freeman and Company, San Francisco. 1964.

Dictionary of Word Roots and Combining Forms. By Donatp J. Borror. vi +
134 pp. National Press, Palo Alto, California. 1960.

A WEST AMERICAN JOURNAL OF BOTANY

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Institutional abbreviations in specimen citations should follow Lanjouw
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Abbreviations of botanical journals should follow those in Botanico-
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Footnotes should be avoided whenever possible.

Membership in the California Botanical Society is normally considered
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VOLUME 20, NUMBER 7 JULY, 1970

Contents |

EMBRYOLOGY OF CHRYSOTHAMNUS (ASTEREAE,
ComposiTaE), Loran C. Anderson 327

Two NEw SPECIES OF LAMOUROUXIA (SCROPHULARIACEAE)
In Mexico, Wallace R. Ernst and Michael F. Baad 342

A New TETRAGASTRIS (BURSERACEAE) FROM
PanaMA, Duncan M. Porter 346

Notes on SOME MEXICAN SPECIES OF GOSSYPIUM
(MatvaceaE), Paul L. Fryxell 347

OENOTHERA BRANDEGEEI FROM BAJA CALIFORNIA, MEXICO, AND A
REVIEW OF SUBGENUS PacHYLoPHuS, Peter H. Raven 350

PoLLeEN APERTURE VARIATION AND PHYLOGENY IN DICENTRA
(FuMARIACEAE), Kingsley R. Stern 354

Fossit LEAVES oF LYONOTHAMNUS, Satish C. Banwar 359

CHROMOSOME STUDIES IN MELAMPODIUM (COMPOSITAE,
HELIANTHEAE), Tod F. Stuessy 365

Harotp Ernest Parks, Lee Bonar 3/3

New ReEcorps oF MyxXOMYCETES FROM CALIFORNIA IV.
Donald T. Kowalski and Dwayne H. Curtis Sid,

Notes aND NEws: RECORDS AND OBSERVATIONS ON
A Rare PLANT, OXALIS LAXA IN CALIFORNIA, John Weiler 349
EDITORSHIP OF MaApRONO 364
FASCIATION OF CoasTaL REpwoops, Rudolf W. Becking;
Back Issues oF MaproNo 382

REVIEWS: JOHN THomMas Howe Lt, Marin Flora, Manual of
Flowering Plants and Ferns of Marin County
(Peter H. Raven) ; Cartes B. HEIsEr, Jr., Nightshades,
The Paradoxical Plants (Dennis R. Parnell) ; Roxana S.
Ferris, Flowers of the Point Reyes National Seashore
(John H. Thomas) 383

A WEST AMERICAN JOURNAL OF BOTANY > SPO

FBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
| ERM HSON;>
Se Py

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~,

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SPR

0 1971
NO SRARI pee

MADRONO th:
A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established —
1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional
subscription price $12.00 per year. Published quarterly in January, April, July, and
October by the California Botanical Society, Inc., and issued from the office of
Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley,
California. Orders for subscriptions, changes in address, and undelivered copies
should be sent to the Corresponding Secretary, California Botanical Society, Depart-
ment of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS
CLASS OF:

1970—Lyman BEnson, Pomona College, Claremont, California
Mitprep E. Marutas, University of California, Los Angeles

1971—Marion OwnBEy, Washington State University, Pullman
Joun F. Davipson, University of Nebraska, Lincoln

1972—Ira L. Wiccins, Stanford University, Stanford, California
REED C. Rois, Harvard University, Cambridge, Massachusetts

1973—WALLACE R. Ernst, Smithsonian Institution, Washington, D.C.
Roy L. Taytor, University of British Columbia, Vancouver

1974—-KEenTON L. CHAMBERS, Oregon State University, Corvallis
EMLEN T., LitTEL, Simon Frazer University, Burnaby, British Columbia

1975—ArtTuRO GoMEz Pompa, Universidad Nacional Autonoma de México
Duncan M. Porter, Missouri Botanical Garden, St. Louis

Editor — Jonw H. THomas
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCasxKILy
P.O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

The Council of the California Botanical Society consists of the officers listed
above plus the immediate past President, Harry Thiers, Department of Ecology and
Systematic Biology, San Francisco State College; the Editor of Madrofio; and
three elected Council Members: Robert Ornduff, Department of Botany, University
of California, Berkeley; Malcolm Nobs, Carnegie Institution of Washington, Stan-
ford; and Elizabeth McClintock, Department of Botany, California Academy of
Sciences, San Francisco.

EMBRYOLOGY OF CHRYSOTHAMNUS
(ASTEREAE, COMPOSITAE)

LORAN C. ANDERSON

Embryological information is scanty on the wide-spread Chryso-
thamnus of western America. “Embryology” is used here in its broad
sense, following Davis (1966) and others. Snow (1945) reported details
of gametogenesis for a population referrable to C. nauseosus (Pallas)
Britt. ssp. albicaulis (Nutt.) H. & C. Although gametogenesis is similar
in some other subspecies of C. nauseosus, it certainly is not homogeneous
throughout the genus. Embryological data on all species of Chryso-
thamnus are recorded here as part of an intensive study (Anderson,
1970) on the floral anatomy of the genus. Those features of embryology
that characterize the genus or the entire family are treated only briefly.

METHODS AND MATERIALS

Over 90 populations were studied. Most materials were preserved in
FPA in conjunction with general anatomic studies. That fixative was
acceptable but caused considerable shrinkage in the embryo sacs. Kill-
ing heads in Craf V (Sass, 1958) resulted in much better preservation.
Fixed tissues were dehydrated in a graded TBA series, embedded in
paraplast, and cut at 9-12), depending on age of the flowers. Sectioned
tissues used in general studies were stained in safranin-fast green with
tannic acid-ferric chloride mordanting. Tissues prepared specifically
for this study were stained in Harris hematoxylin followed by fast green
and mounted in permount. Many of the slides were prepared by Kent
Fish.

EMBRYOLOGY

Growth of flowers in one head is closely synchronized. Although in-
dividual flowers develop acropetally on the receptacle (Snow, 1945),
they usually undergo either microsporogenesis or megasporogenesis
simultaneously regardless of differences in individual flower size. The
least synchronous species is C. linifolius Greene. In it, older flowers
of the head have mature embryo sacs when the youngest flower under-
goes meiosis in the ovule.

Anthers in Chrysothamnus are tetrasporangiate. Method of wall for-
mation is the dicotyledonous type (Davis, 1966) with a glandular
tapetum. Many aspects of microsporogenesis have been considered
earlier (Anderson, 1966).

The ovules are anatropus, unitegmic, and tenuinucellar. Four cellular
megaspores result from meiosis. The chalazal megaspore usually pro-

Manprono, Vol. 20, No. 7, pp. 337-384. March 17, 1971.

337

[Vol. 20

MADRONO

338

1970] ANDERSON: CHRYSOTHAMNUS 339

duces the embryo sac; however, a functional micropylar megaspore
was observed in one sample of C. linifolius (Anderson 2738, KSC).
The nucellus is intact at the tetrad stage but breaks down concurrently
with three megaspores before the enlarged functional megaspore divides.
On disintegration of the nucellus, an endothelium (integumentary tape-
tum) develops; its cells are often binucleate.

Taxa of Chrysothamnus develop a Polygonum type embryo sac.
Mature embryo sacs (figs. 1-8) are typically long and narrow. The
longest observed (fig. 1) was 495u (44% of total ovule length at that
age). In most samples embryo sacs range from 150-300 (15-20% of
total ovule length). The wider embryo sac shown in fig. 7 probably is
a post-fertilization condition.

The synergids are somewhat vacuolated and usually extend into the
micropyle, especially in C. nauseosus (fig. 7). The egg is character-
istically overarched by the central cell (figs. 2, 7) in which the polar
nuclei fuse prior to fertilization. Average antipodal size is 20-30un.
The largest is usually the chalazal antipodal, over 200y in C. viscidi-
florus (Hook.) Nutt. ssp. lanceolatus (Nutt.) Piper (fig. 1) and C. ves-
cidiflorus ssp. puberulus (D.C.Eat.) H. & C. The largest basal anti-
podal observed was 80u (fig. 2). Occasionally, one of the interstitial
antipodals is the largest, but never over 50y long.

The antipodal apparatus in the megagametophyte varies widely.
Three trends were observed: 1, antipodal cells reduced to two, one
with two nuclei (fig. 8); 2, nuclei multiplied so that one or more of
the three antipodals are multinucleate (fig. 2), and some individual
nuclei may be multinucleolate; 3, divisions continued to more than three
antipodals (fig. 3). The last two trends are interrelated; several anti-
podals may be present with one or more multinucleate (fig. 4). Anti-
podal cell number and frequency for all species are listed in Table 1.

In the Compositae, antipodal multiplaction often results in persistent,
haustorial cells (Davis, 1966). In Chrysothamnus, truly haustorial anti-
podals are infrequent. A strong lateral haustorium was observed only
in C. viscidiflorus ssp. humilis (Greene) H. & C. (fig. 5). An indication
of a weak lateral haustorium was seen in C. linifolius (fig. 6). The pro-
duction of an haustorium by elongation of the chalazal antipodal (figs.
1-5) is more frequent than lateral haustoria. If the chalazal antipodal
surpasses the endothelium, it is considered haustorial. Nonetheless, haus-
torial activity is not prolonged in Chrysothamnus because the antipodals
usually become inactive and disintegrate shortly after fertilization or

Fics. 1-8. Mature embryo sacs and endothelium in Chrysothamnus: 1, C. viscidi-
florus ssp. lanceolatus (Anderson 2717, KSC); 2, C. nauseosus ssp. leiospermus
(Anderson 1995, KSC) ; 3, C. pulchellus (Anderson 3213, KSC); 4, C. parryi affin.
ssp. nevadensis (Anderson 2966, KSC); 5, C. viscidiflorus ssp. humilis (Anderson
2950, KSC) ; 6, C. linifolius (Anderson 2500, KSC); 7, C. nauseosus ssp. nauseosus
(Anderson 2707, KSC) ; 8, C. viscidiflorus ssp. viscidiflorus (Anderson 2900, KSC).

340 MADRONO [ Vol. 20

TABLE 1. NUMBER OF ANTIPODALS AND THEIR OBSERVED FREQUENCY IN EMBRYO
Sacs IN CHRYSOTHAMNUS.

NAUSEOS|
C. parryi

C. nauseosus

CHRYSOTHAMNUS

. linifolius

. Spathulatus

. albidus Ox
C. greenei

C. viscidiflorus Ox
PULCHELLI

C. vaseyi

(Sp eS

C. molestus

P 2 99S OF
2
|

C. depressus

C. pulchellus
PUNCTATI

C. paniculatus

29 9909 g
> OO

2
96

C. teretifolius

occasionally prior to fertilization as in Anderson 2927 (KSC, C. nause-
osus ssp. albicaulis). Relatively persistent chalazal haustoria were
found in C. albidus (Jones) Greene and C. greenez (Gray) Greene.

Shortly after fertilization the embryo sac becomes laterally distended
in all taxa (as in fig. 7). Embryogeny is the asterad type. Endosperm
formation is nuclear at first. Further growth precipiates wall formation
that progresses to form a cellular endosperm.

DISCUSSION

Size of embryo sacs in Chrysothamnus varys widely. Extremes (figs.
1, 8) are found in C. viscidiflorus, the only species known to contain
polyploids (Anderson, 1966). Polyploidy evidently has no significant
bearing on embryo sac size; the largest as well as the smallest come
from diploids.

Howe (1959) found lateral haustoria in species of Grindelia, Gutier-
rezia, and Haplopappus. He used that phenomenon (less frequent than
chalazal haustoria in Astereae) to suggest interrelationship of the taxa.
Nevertheless, the presence of a lateral haustorium in C. viscidiflorus ssp.
humilis does not suggest that it is related to those taxa also. It does,
however, point to the distinctness of that taxon. In fact, unique features
in its floral anatomy reinforced my decision to recognize it as a separate
species (Anderson, 1970).

1970] ANDERSON: CHRYSOTHAMNUS 341

Species are listed in table 1 in a phylogenetic sequence (assumed from
gross morphology). Patterns in antipodal numbers are fairly well in
accord within sections, except for the unusual bimodal pattern in C.
paniculatus (Gray) Hall. In section Chrysothamnus, and particularly in
the Pulchelli, the average number reflects the assumed phylogeny.

Antipodal apparatus in Vauseosi is especially interesting as both spe-
cies of that section are considered primitive and have numerous sub-
species. In a statistical study of floral features (Anderson and Fisher,
in press) C. parryi (Gray) Greene ssp. parryi was determined the most
primitive taxon in the genus. If the postulate that an antipodal number
of three is primitive, it is noteworthy that embryo sacs in C. parryi ssp.
parryi usually have five to seven antipodals. The more primitive taxa of
C. nauseosus have megagametophytes with three to five antipodals. In
both species the higher numbers of antipodals are found in taxa adapted
to xeric environments. If that is a trend, it is not universal; in section
Chrysothamnus, C. albidus and C. viscidiflorus with two to three anti-
podals are fairly well adapted to xerism.

Hall (Hall and Clements, 1932) stated C. paniculatus was the most
stable and primitive species in the genus. Its uniformity in external
morphology is not seen in its embryology. Contrary to Hall’s view, that
species is very specialized (table 1). I had considered it less so than
C. teretifolius (Dur. & Hilg.) Hall; however, embryological information
reinforces a more recent determination (Anderson and Fisher, in press)
that it is more specialized.

The close similarity of the Punctati to Haplopappus, section Erica-
meria (Anderson, 1966), now includes their embryology. Haplopap pus
propinquus Blake (Raven 16802, RSA) has four antipodals in the meg-
agametophyte, and H. trianthus Blake (Anderson 3032, KSC) has five.
Similarly, the Nauseosi are close to Haplopappus, section Macronema
(Anderson and Reveal, 1966); H. macronema Gray from California
(Anderson 2922, KSC) has five to seven antipodals, and another popu-
lation from Colorado (Anderson 2540, KSC) has 10 antipodals.

In the Astereae, a multinucleate antipodal apparatus is known for
several genera related to Chrysothamnus; i.e., Haplopappus (Harling,
1951), Petradoria (Anderson, 1963), and Solidago (Beaudry, 1958).
In none, with the possible exception of Haplopappus, is the latitude of
variability in the megagametophyte so great as it is in Chrysothamnus.

This study was supported by National Science Foundation grant GB
3058 and this paper is contribution number 987 of the Division of
Biology, Kansas Agricultural Experiment Station, Kansas State Uni-
versity.

Division of Biology, Kansas State University, Manhattan

342 MADRONO [Vel. 20

LITERATURE CITED

ANDERSON, L. C. 1963. Studies on Petradoria (Compositae): anatomy, cytology, tax-
onomy. Trans. Kans. Acad. Sci. 66:632-684.

. 1966. Cytotaxonomic studies in Chrysothamnus (Astereae, Compositae).
Amer. J. Bot. 53:204-212.

. 1970. Flcral anatomy of Chrysothamnus (Astereae, Compositae). Sida
3:466-503.

. and J. L. REvEAL. 1966. Chrysothamnus bolanderi, an intergeneric hybrid.
Madrono 18:225-233.

.. and P. S. Fisuer. Phylogenetic indicators frem floral anatomy in Chryse-
thamnus (Astereae, Compositae). Phytomorphology (in press).

BrAupry, J. R. 1958. Studies on Solidago. III. Megasporogenesis, development of
megagametophyte and mode of repreduction in Solidago altissima. Proc. Genet.
Soc. Canada 3:7-14.

Davis, G. L. 1966. Systematic Embryology of the Angiosperms. Wiley, New York.

Harr. H. M. and F. E. CLements. 1923. The phylogenetic method in taxcnomy
The North American species of Artemisia, Chrysothamnus, and Atriplex. Publ.
Carnegie Inst. Wash. 326.

Hariinc, G. 1951. Embryological studies in the Compositae. III. Astereae. Acta
Hort. Berg. 16:73-120.

Howe, T. D. 1959. Recent studies of the female gametophyte in the Compositae,
especially in the tribe Astereae. Proc. IX Int. Bot. Cong. 2:171-172.

Sass, J. E. 1958. Botanical Microtechnique, 3rd ed. Iowa State Univ. Press.

Snow, E. 1945. Floral morphology of Chrysothamnus nausecsus speciosus. Bot.
Gaz. (Crawferdsville) 106:451-462.

TWO NEW SPECIES OF LAMOUROUXIA
(SCROPHULARIACEAE) IN MEXICO

WALLACE R. ERNsT and MICHAEL F. BAap

The genus Lamourouxia H. B. K., nom. cons. (Taxon 18: 479-480.
1969), allied to the Rhinanthoideae of Scrophulariaceae, is distributed
from northern Mexico through Central America to South America about
as far south as Lima, Peru. Of the approximately 26 species, the following
two have been studied jointly. They are placed in section Hemispadon
Bentham, having long tubular, red corollas with a pair of large anthers
and a pair of sterile staminal filaments. Search through the literature
and at least 25 herbaria, including the major ones of Europe, has failed
to disclose other names or collections for these two species. They are
being described here to make the names available before a taxonomic
revision and a discussion of their morphological relationships to avoid
a longer author citation.

Lamourouxia colimae Ernst & Baad, sp. nov. Herba suffruticosa,
erecta. Folia glabriuscula, elliptica vel ovata, pauce dentata, attenuata
basin versus, nervatura supra impressa. Calyx glaber, limbis patulis,
late triangulatis. Corolla coccinea, cylindrica, labio superiore bifido, lobis
rotundatis parum cuspidatis, non vel minime reflexis, labio inferiore

1970] ERNST & BAAD: LAMOUROUXIA 343

EARS PSR

PRET EO

RSENS

DEER WINES

SEE EERE IO BEBE EC RRER

Fic. i. Holotype, Lamourouxia colimae.

344 MADRONO [Vol. 20

brevissime trifido. Stamina inferiora fertilia, superiora breviora, parva,
sterilia. Affinis sectionis Hemispadon, verosimiliter L. gracilis vel L.
lanceolata.

Type: Colima: mountain summits near pass ca. 11 miles south-south-
west of Colima on Manzanillo road, elevation 500 m, Rogers McVaugh
18077 (& H. F. Loveland, R. W. Pippen) (MICH-holotype), Sept. 21,
1958 (fig. 1).

Stems erect to 1 m tall. Leaves glabrous or with very few hairs, el-
liptical or ovate, 18-26 mm long, 6-10 mm broad, attenuate basally,
mostly with a very short or indistinct petiole, margins revolute with 3—6
teeth on a side, veining standing out below, the midrib and 3-6 laterals
recessed above. Inflorescence erect; pedicels 3—4 mm long. Calyx gla-
brous or microscopically papillate and possibly glandular, 6—7 mm long,
4 mm broad, the lobes 4-6 mm long, broadly triangular, spreading.
Corolla scarlet, 30-35 mm long, 6-7 mm broad, the upper lip 12-13 mm
long, the lobes shallow, rounded, slightly cuspidate, little or not re-
reflexed; the lobes of the lower lip about 1 mm long. Upper pair of
stamens /% or less the length of the corolla, their anthers vestigial and
glabrous; lower pair of stamens as long as the corolla, the filaments
slightly expanded distally, the anthers shaggy pubescent, their lobes
short acuminate at base. Style with a few short hairs.

This species is known only from the type collection, thus the discus-
sion cannot reflect a sense of variation. The new species seems some-
what similar to L. gracilis Robinson & Greenman, a rare species in Guer-
rero and Morelos, having proportionately broader corollas and narrower,
basally more attenuate leaves. The flowers recall those of L. lanceolata
Bentham in DC. in Oaxaca and Central America but in that species the
inflorescences are dependent, the flowers resupinate, and the serrate
leaves are longer and narrower. The flowers also resemble those of
L. gutierrezii Oersted in Bentham & Oersted in Central America but
that species has scabrous leaves and calyx.

Lamourouxia jaliscana Ernst & Baad, sp. nov. Herba suffruticosa,
erecta. Folia pubescentia, anguste lanceolata, grosse dentata, attenuata
basin versus. Calyx glandulo-pubescens, limbis subulatis. Corolla rubra,
cylindrica, labio superiore bifido, lobis late attenuatis, reflexis, labio in-
feriore anguste trifido. Stamina inferiora fertilia, superiora breviora,
parva, sterilia. Affinia sectionis Hemispadon, specie proxima dubia.

Type. Jalisco: Sierra de Caule, southwest of Talpa de Allende, south-
west of Piedra Rajada, elevation 1800-2250 m, Rogers McVaugh 14250
(& J. Sooby, Jr.) (MICH-holotype, and duplicate), Nov. 19-21, 1952,
(fig. 2).

Other material examined. Jalisco: 11-12 miles south of Talpa de Al-
lende, headwaters of west branch of Rio de Talpa, elevation 1200-1700
m, McVaugh 21325 (& C. Feddema, R. Pippen) (MICH) Nov. 23-25,
1960.

1970] ERNST & BAAD: LAMOUROUXIA 345

Fic. 2. Holotype, Lamourouxia jaliscana.

346 MADRONO [Vol. 20

Stems erect to 1.5 m tall. Leaves very pubescent, possibly scabrous
above, soft below, narrowly lanceolate, 50-80 mm long, 6-10 mm broad,
attenuate apically and basally, revolute, dentate with 12-22 coarse teeth
on a side. Inflorescence erect; the pedicels 5-8 mm long. Calyx glandular
pubescent, 14-16 mm long, 5 mm broad, the lobes narrowly subulate,
10-12 mm long. Corolla red, cylindrical, 56-63 mm long, 9-10 mm broad,
the upper lip 16-25 mm long, the lobes broadly attenuate, 5-7 mm long,
reflexed; the lobes of the prominent lower lip narrow, 5-8 mm long.
Upper pair of stamens half as long as corolla, the anthers vestigial and
glabrous; lower pair of stamens nearly as long as corolla, the filaments
slightly dilated distally, the anthers shaggy pubescent, the lobes acu-
minate at base. Style with spreading hairs. Fruits ovoid, 12-13 mm
long, 7-8 mm broad.

This species, known from only two collections, is distinguished by its
large, long, narrow, coarsely-toothed and very pubescent leaves, the long
narrow, glandular calyx lobes, and the large size of the corolla. The size
of the leaves and flowers somewhat recalls L. longiflora Bentham of
section Lamourouxia, which has entire leaves. The texture of the leaves
somewhat recalls L. viscosa H. B. K., but the new species does not
seem to be closely allied to any other species.

Department of Botany, Smithsonian Institution, Washington, D. C.

Department of Biological Sciences, Sacramento State College,
Sacramento, California

A NEW TETRAGASTRIS (BURSERACEAE) FROM PANAMA
DUNCAN M. PORTER

The following new species of Tetrvagastris was discovered during a
survey of the Burseraceae for the Flora of Panama. It is apparently en-
demic to the Republic of Panama.

Tetragastris tomentosa D. M. Porter, sp. nov. A 7. panamensis
(Engl.) O. Ktze. quadrimerus floribus et foliolis costa infernis conspicuis
luteis-tomentosis statim diagnoscenda.

A tree ca. 8 m high; branchlets densely yellowish-tomentose. Leaves
once-pinnate, 37 cm long or longer and to 24.5 cm wide; petioles striate,
densely yellowish-tomentose, canaliculate above, 84 mm long; leaflets
7(?), membranaceous, elliptic to ovate, abruptly acuminate apically,
slightly oblique basally, the main vein densely yellowish-tomentose be-
low, minutely puberulent above, the secondary veins and blade with
scattered trichomes on both surfaces, to 18.5 cm long and 9 cm wide,
the laterals largest, the lowermost smallest and reflexed; petiolules
densely yellowish-tomentose, canaliculate above, swollen apically, the

1970] PORTER: TETRAGASTRIS 347

laterals 15-24 mm long, the terminals 48-59 mm long. Staminate inflor-
escences axillary panicles, branched from the base, spreading, densely
yellowish-tomentose, to 19 cm long. Staminate flowers cream-yellow,
4-merous; pedicels sparsely yellowish-tomentose, 1.5-3 mm long; calyx
broadly cupular, sparsely yellowish-tomentose, ca. 1 mm high and 2.5
mm wide, the lobes 4, acute, spreading; corolla yellowish, tubular,
densely yellowish-tomentose without, pubescent within, ca. 3 mm long,
the lobes 4, thick, acute, 1-1-5 mm long and ca. 1 mm wide, with an
incurved apical process adaxially; stamens 8, ca. as high as the corolla
tube, the filaments subulate, inserted at the base of the disc between
the lobes, the 4 opposite the sepals adnate basally to the corolla below
the clefts, the anthers sagittate, basifixed; disc sulcate, 8-lobed, glab-
rous, half as high as the ovary; ovary tomentose, ovoid, ca. 1 mm in
diameter. half immersed in the disc, the style columnar. Fruits unknown.

Type. Panama: Bocas del Toro Province, Fish Creek Hills, H. von
Wedel 2398 (GH, MO-holotype, US) May 7, 1941. Known only from
the type collection.

Tetragastris tomentosa is easily separated from T. panamensis
(Engl.) O. Ktze., the only other species of the genus known from Pan-
ama, by a number of characters, the most obvious being its 4-merous
flowers and the conspicuous yellow tomentum on the midribs of the
leaflets. Tetragastris panamensis has 5-merous flowers and sparingly
pubescent to glabrate lower leaflet midribs. A flower will be illustrated
in the forthcoming treatment of the Burseraceae for the Flora of

Panama. : : ; : ;
Missouri Botanical Garden, St. Louis, Missouri

NOTES ON SOME MEXICAN SPECIES OF
GOSSYPIUM (MALVACEAE)

PAUL A. FRYXELL

Gossypium aridum (Rose & Standley) Skovsted was originally de-
scribed as Erioxvwlum aridum Rose & Standley and based on material
from the state of Sinaloa (Culiacan, Rose, Standley & Russell 14199,
US). Rose and Standley regarded this species as distinct from Erioxvlum
palermt (Rose) Rose & Standley based on a collection from Colima
(Palmer 1316, GH, MEXU, US). Prokhanov accepted this view and
transferred the latter species, then known only from the type, to Gossy-
pium as G. rosei Prokh. Consequently, G. aridum has long been regarded
as endemic to Sinaloa.

The distinctions between these two taxa are slight. Indeed, more re-
cently collected material shows that they are conspecific. Moreover, this
species ranges southeastward far beyond the states of Sinaloa and
Colima, covering at least 1000 miles of the west coast of Mexico, and
extending to beyond Tehuantepec. The following is a list of specimens

348 MADRONO [Vol. 20

of G. aridum excluding numerous collections from Sinaloa.

COLIMA: Tecoman, Miranda 9108 (MEXU); Tecolapa, McVaugh
15543 (MEXU, US). JALISCO: Navidad, McVaugh 11882 (MEXU).
MICHOACAN: Infiernillo, Bratz s.n., 28.xi1.1964 (MEXU). GUER-
RERO: 95 miles NW of Acapulco, Fryxell 625 (TAES); 57 miles NW
of Acapulco, Anderson & Laskowski 4489 (MICH-n.v.), Fryxell 624
(GH, MICH, TAES, UT); Acapulco, Miranda 4350 (MEXU). OAX-
ACA: 5 miles W of Tehuantepec, Fryxell 753 (F, MEXU, MO, NA,
TAES, U, UC, US), Smith 3221 (n.v.: MEXU, PH, US); 7 miles W of
Nilotepec, Fryxell & Bates 908 (BH, TAKES).

Two recent articles (Fryxell, 1965; Fryxell and Parks, 1967) have
dealt with the distribution of Gossypium trilobum (Moc. & Sess. ex DC).
Skov. An additional specimen has come to my attention from Polotitlan
(Miranda 27172, MEXU) that extends our knowledge of the distribution
of this species. Polotitlan is in the northern extremity of the state of
México at an elevation of 2400 m. Previous collections have all been
made between 800 and 1800 m elevation.

This record notably extends not only the geographical and altitudinal
ranges of G. trilobum but also those of the entire subgenus, composed
of the American diploid species of Gossypium, of which G. trilobum is
the type species. These plants are distributed solely on the Pacific
(western )slopes of the New World, except in the Isthmus of Tehuan-
tepec, where G. gossypioides (Ulbr.) Standley also crosses a short dis-
tance to the east of the continental divide north of Oaxaca. The present
specimen was collected very near the continental divide and, in fact,
from an area that drains to the east. This additional trivial exception
emphasizes the distinctive western distribution of the group. No col-
lections have previously been reported, to my knowledge, from elevations
this high.

The basionym of Gossypium gossypioides (Ulbr.) Standley is Selera
gossvpioides Ulbr., and was based on a specimen from Oaxaca (San
Bartolo Yautepec, C. & E. Seler 1700, Jan. 6, 1896). The holotype was
at the Berlin herbarium but is now lost. Since no isotypes are known to
exist, and Ulbrich cited no other material, it becomes necessary to desig-
nate a neotype.

Such a designation raises no significant problems, since the species
shows relatively little variability and is quite distinctive, both in mor-
phology and in distribution. Ulbrich indeed chose to place it in a mono-
typic genus. The specimen chosen as neotype was collected within ap-
proximately 10 km of the type locality, at what appears to be the lower
elevational limit of the species, these limits being approximately 800—
1400 m.

Neotype of Selera gossypioides Ulbr.: OAXACA: 39 km W of Tequi-
sistlan, on Hwy. 190 at K 706", in rocky hills. Elev. 2900 feet. Tree to
15 feet tall. Fryxell 757 (F, MEXU, MO, NA, US-neotype, TAKES),
Sepe. 9, 1968:

1970] FRYXELL: GOSSYPIUM 349

Note added in proof: The distribution of G. aridum may also be ex-
tended inland to the state of Puebla on the basis of the following speci-
mens. PUEBLA: Tecomatlan, C. Pollatzin, Miranda 2609 (MEXU);
Tecuatitlan San Martin, near Tecomatlan, 3000 ft. alt., Fryxell 759
(ARIX, MEXU, MICH, NA, NY, US), Fryxell & Bates 918 (BH,
MEXU, NA, US).

Plant Sciences Research Division, U.S. Department of Agriculture at
Texas A & M University, College Station

LITERATURE CITED

FryYxeELL, P. A. 1965. A further description of Gossypium trilobum. Madrono
18:113-118.

. and C. R. Parks. 1967. Gossypium trilobum: an addendum. Madrono
19:117-123.

NOTES AND NEWS

RECORDS AND OBSERVATIONS ON A RARE PLANT, OXALIS LAXA IN CALIFORNIA. —
Oxalis laxa H. & A. has previously been reported as sparingly naturalized at Stinson
Beach, Marin Co. and near San Andreas, Calaveras Co., California where it has
been introduced from Chile (Munz, A California flora, 1959). Recent field collec-
tions and subsequent investigation of previously undetermined specimens in the
Fresno State College Herbarium have revealed several additional populations, some
very extensive, outside the range reported in Munz. The new populations are docu-
mented by herbarium vouchers filed at Fresno State Coliege Herbarium. My col-
lections have been widely distributed to other institutions.

The sites reported below are centered in an area less than ten miles wide on
either side of the San Joaquin River extending both upstream and downstream
from the former site of Fort Millerton. All collections were made between 300 and
700 feet elevation. Those sites some distance removed from the river are found in
drainage basins of creeks where the plants usually grow on high, dry soil away
from streamside. Soil is thin, of decomposed granite, and usually supports sparse
vegetation. Oxalis at these sites may grow fully exposed to sun or occasionally in
dense shade provided by boulders or scattered shrubs of Ceanothus cuneatus or
Lupinus albifrons and trees of Quercus douglasii.

Plants grown from seeds taken at the Madera Co. site yielded chromosome
counts of n = 10 from pollen mother cells squashed in aceto-carmine and proved
to be self-compatable, setting abundant seeds when cultivated individually in pots.
In this respect the plants are illustrative of the idea proposed by Baker (Evolution
9:347-348, 1955) that establishment after long distance dispersal is greatly enhanced
if the organism is self compatable. It remains to be seen whether this group of popu-
lations is to be regarded as resulting from a separate introduction or whether it is
part of a much broader undetected distribution in the sierran foothills ranging
southward from the San Andreas site.

Fresno Co.: along the San Joaquin River near Fort Millerton, Quibell 1158, April
4, 1929; Temperance Flat east of Friant, B. Brock 415, March 19, 1959; along the
San Joaquin River 2 miles downstream from Friant Dam, Field & Munger, May 1,
1960; along Little Dry Creek near its crossing with Millerton Road 2 miles east of
Auberry Road junction, Weiler 65024, April 13, 1965.

Madera Co.: near Cottonwood Creek 5 miles north of the San Joaquin River
between Friant and North Fork, Weiler 65007, March 5, 1965.—JoHN WEILER,
Depariment of Biology, Fresno State College, Fresno.

OENOTHERA BRANDEGEEI FROM BAJA CALIFORNIA,
MEXICO, AND A REVIEW OF SUBGENUS PACHYLOPHUS
(ONAGRACEAE)

PETER H. RAVEN

In December 1887, while collecting for the Smithsonian Institution,
Edward Palmer made two collections of an interesting annual Oenothera
on stony ridges near Bahia de los Angeles on the east coast of Baja Cali-
fornia (542, GH, and 582, US). These collections, each of a single
plant, were determined by Sereno Watson as O. caespitosa Nutt. They
remained under this name until 1930, when they were studied by P. A.
Munz (1931) for his revision of Oenothera subg. Pachylophus. Munz
considered them a distinctive unnamed variety of O. caespitosa which
he named var. brandegeei Munz (1931), selecting no. 542 as the holo-
tvpe. Munz assumed this plant to be the same as the one mentioned by
T.S. Brandegee (1889) as “Oenothera caespitosa Nutt. var. Leaves finely
divided and villous——E] Campo Aleman”; but Brandegee’s specimen
(El Pozo Aleman, 23 April 1889, UC) had long since been determined
by Katherine Brandegee as O. primiveris Gray, and Munz himself con-
curred when he examined the specimen in 1932. In 1965, treating the
Onagraceae for the North American Flora Munz raised this rare and
local endemic to the rank of subspecies as O. caespitosa Nutt. ssp. bran-
degeei (Munz) Munz. Until 1966, Palmer’s two plants remained the only
known representatives of O. caespitosa var. brandegeei.

Recently, Reid Moran very kindly sent me a collection he had ob-
tained 22 April 1966 on Isla Angel de la Guarda in the Gulf of Cali-
fornia. The plants grew among volcanic rocks on the north slope of the
peak southwest of Pond Island, ca. 350 m elevation, near 29°01’N,
113°10’ W, 12983 (DS, RSA, SD). Moran found occasional woody dead
plants from earlier years’ growth with the capsules adhering, and also
a few dozen living ones with leaves mostly 3—4 cm long and one capsule
per plant (Palmer’s had leaves respectively ca. 8 and ca. 15 cm long).
In these depauperate plants, the terminal lobes of the leaves are less
prominent than in Palmer’s collections, and the flowers are smaller:
hypanthium 5 mm. long, sepals 5 & 0.7 mm, petals about 8 & 4.5 mm,
filaments 5 mm long, anthers about 3 mm long in Moran’s material, and
respectively 38 mm, 12 & 3 mm, 16 & 15 mm, 7 mm. and 5 mm in
Palmer’s 582. In every other way, however, Moran’s plants are identical
with Palmer’s, and there is no doubt that all three represent the same
entity. The dead plants of earlier years that Moran collected were much
more robust, with about 30 capsules per plant.

When I examined Moran’s material, it became clear to me that these
slender annuals of Baja California should not be considered conspecific

350

1970| RAVEN: OENOTHERA Sol

with O. caespitosa Nutt., itself an exceedingly polymorphic species, but
a robust, tufted perennial with much larger flowers. O. caespitosa is
basically a species of the Great Basin of the western United States,
extending south to the San Bernardino Mts. of southern California and
the Huachuca Mts. of southeastern Arizona, but not known from Mexico.
Oenothera caespitosa is, as far as is known, always self-incompatible
(Gregory, 1963; Klein, pers. comm.) whereas “‘var. brandegeet,” with its
small flowers and stigma surrounded by the shedding anthers at anthesis,
is highly autogamous as shown by three plants grown at Stanford from
Moran’s collection. The change from self-incompatibility to autogamy
is known often to accompany a change from the perennial to the annual
habit in angiosperms.

This discussion to this point establishes the desirability of separating
“var. brandegeei” from O. caespitosa at the specific level. There is,
however, another basically annual species with small white flowers which
is closely related to O. caespitosa, namely O. cavernae Munz (1941).
Oenothera cavernae is so similar to “var. brandegeet” that Munz anno-
tated a specimen of the former (‘‘Utah, Capt. Bishop, 1872,” US) as
follows: ‘““Oenothera caespitosa var. brandegeet ... This is a plant from
Lower California. The data on the label certainly incorrect. PAM—
1930.” This was, of course, before Munz was aware of the existence of
O. cavernae as a distinct entity. Despite their overall similarity, there
are a few differences which clearly distinguish O. cavernae from “var.
brandegeet.”” As pointed out by Munz, the leaves of “var. brandegeei”’
are distinctive in the O. casespitosa alliance in being deeply divided into
narrow, acuminate, lobes which are directed forward, toward the apex
of the leaf. These lobes are much reduced, and the treminal lobe is very
prominent in well-developed individuals. In O. cavernae, on the other
hand, as is usually the case in O. caespitosa, the lateral lobes are acute
or obtuse and stand out at right angles to the rachis. The terminal lobe
of the leaf is much less prominent than in “var. brandegeet.”’ The cap-
sules of the iwo entities likewise differ modally, those of var. brandegeei”’
being short and stout, 14-18 mm. long, with very prominent, well sepa-
rated tubercles along the lines of dehiscence; whereas those of O. cav-
ernae are often longer, 15-38 mm long, with an acuminate apex and less
prominent or distinct tubercles.

In summary, their morphological distinctiveness and wide geographi-
cal separation suggests that these two white-flowered, autogamous annual
species were derived independently from O. caespitosa as the deserts of
western North America expanded and the available habitats became pro-
gressively less favorable for their perennial ancestor. The distinctive
leaves of “var. brandegeei” suggest that it may have been the earlier
derivative, an hypothesis consistent with its present geographical sepa-
ration from O. caespitosa. Oenothera cavernae occurs on the desert slopes
of southern Nevada (Clark Co.) and southeastward to Toroweap and

$52 MADRONO [Vol. 20

Havasu Canyon on the Colorado River in northwestern Arizona. O. caes-
pitosa occurs at higher elevations and presumably in more mesic sites,
often associated with juniper woodland and sometimes with pinyons,
in the same region. In view of these considerations, a new combination
seems appropriate.

OENOTHERA brandegeei (Munz) Raven, comb. nov. O. caespitosa
Nutt. var. brandegeei Munz, Amer. J. Bot. 18:732. 1931; O. caespitosa
ssp. brandegeet (Munz) Munz, N. Amer. FI. II. 5:101. 1965.

As I have earlier pointed out the importance of a modern and com-
prehensive reevaluation of sectional and subgeneric alignments in Oeno-
thera (Raven, 1964), it may be appropriate at this point to offer a few
comments concerning the relationships of the six species currently re-
ferred to subg. (sec.) Pachylophus and the overall constitution of the
group. First, it is clear that O. caespitosa, O. cavernae, and O. brande-
geei form a close-knit alliance. In the protologue of O. cavernae, Munz
compared it with the yellow-flowered desert annual O. primiveris Gray,
but these two species do not appear to be closely related. On the other
hand, the annual O. primiveris does appear to be related to the yellow-
flowered perennial O. xylocarpa Cov., a narrow endemic found along the
east flank of the southern Sierra Nevada in California and Nevada. Un-
like O. caespitosa, O. xylocarpa has swollen, fleshy underground parts.
In this, as in the morphology of the capsule, it closely resembles the
sixth species of the group, the white-flowered (not yellow, contrary to
the prediction of Munz (1931; 1965), O. tubifera Sessé & Mocino ex
Ser. of central Mexico.

Oenothera tubifera in turn is obviously closely related to another
white-flowered perennial Mexican species currently referred to subg.
Raimannia: O. muelleri Munz. Although the flowers of O. muzlleri are
much larger, these two species can be crossed easily in cultivation, and
the seeds germinate readily to produce healthy F, individuals. These
two species are identical in capsule morphology and in habit, the plants
producing a series of decumbent flowering branches from a central
rosette.

Ancther Mexican species currently referred to subg. Raimannia, O.
macrosceles Gray, is similar in habit, but has yellow flowers and much
more slender capsules. It is clearly not as closely related to O. muelleri
and O. tubifera as they are to one another. Oenothera macrosceles can
easily be hybridized with O. muelleri and O. tubifera in cultivation, how-
ever, but we have not yet succeeded in germinating the seeds. On the
other hand, Cleland (1968) has recently shown that O. macrosceles does
not hybridize readily with any species of Raimannia. On the balance,
it would seem that O. macrosceles should be placed in subg. Pachylophus.

Although the yellow-flowered O. maysillesiit Munz of Durango, Mexico,
is similar in habit and has been compared with O. muelleri and O. macro-
sceles, its status is currently being investigated, and it is best retained

1970] RAVEN: OENOTHERA 353

at least for the time being, in subg. Raimannia, as originally placed.
Oenothera subg. Raimannia (revised by Munz, 1935) is a relatively
homogeneous group in South America, but has been made to include a
much more diverse assemblage of North American species. Oenothera
macrosceles and O. muelleri, as suggested above, seem best referred to
subg. Pachylophus, and O. albicaulis Pursh and O. coronopifoliaT. & G.
are best romoved to a ditypic group of their own—sect. Kleinia Munz
(1965), perhaps best thought of as intermediate between subg. Raiman-
nia and subg. Anogra. Oenothera organensis is now regarded as belong-
ing to a monotypic sect. Emersonia (Munz, 1965) perhaps intermediate
to subg. Oenothera ( Euoenothera). With these subtractions, subg. (sect. )
Raimannia appears to be a reasonably natural group, although rich in
species. Interestingly, all of the remaining species would have yellow
flowers. .
These rearrangements would leave Oenothera subg. Pachylophus with
a total of eight species, with O. macrosceles and O. xylocarpa yellow-
flowered perennials, O. primiveris a yellow-flowered annual, O. tubifera,
O. muelleri, and O. caespitosa white-flowered perennials, and O. brande-
geet and O. cavernae white-flowered annuals. Relationships within this
group need further clarification by biosystematic studies, but it appears
at present that O. caespitosa, O. cavernae, and O. brandegeei; O. xylo-
carpa and O. primiveris (which have been hybridized experimentally,
although the seeds could not be germinated; Klein, pers. comm.) ;
O. macrosceles; and O. muelleri and O. tubifera constitute four distinct
subgroups. Three species, O. caespitosa, O. xylocarpa, and O. primiverts,
are self-incompatible (Klein, pers. comm.) ; two, O. muelleri and O. mac-
rosceles, are self-compatible but modally outcrossing; and two, O.
brandegeei and O. cavernae, are autogamous. In Oenothera tubifera,
self-pollination is frequent but since a relatively small load of pollen is
deposited on the stigma, full seed set does not normally result.
Oenothera subg. Pachylophus as constituted here appears to include
an assemblage of relatively closely related species, and to embody a use-
ful taxonomic concept. The four groups mentioned above might reason-
ably be regarded as distinct sectioins, but further studies of the entire
genus will be necessary to determine the best systematic treatment for
the group as a whole. It might be noted in closing that O. caespitosa
and O. primiveris include several distinct races best recognized at the
subspecific level, but the other species appear relatively homogeneous.
T would like to thank Ralph E. Cleland, William M. Klein, Reid V.
Moran, and Philip A. Munz for their useful comments on this paper.
This work was supported by National Science Foundation Grant GB
7949X.

Department of Biological Sciences, Stanford University

354 MADRONO [Vol. 20

LITERATURE CITED

BRANDEGEE, T.S. 1889. A collection of plants from Baja California. Proc. Calif. Acad.

Sei, I. 2:11 /=2 16:
CLELAND, R. 1968. Cytogenetic studies on Oenothera, subgenus Raimannia. Jap.

J. Genet. 43 :329-334.
Grecory, D. P. 1963. Hawkmoth pollination in the genus Oenothera. Aliso 5:357-419.
Muwz, P.A. 1931. Studies on Onagraceae VII. The subgenus Pachylophus of the
genus Oenothera. Amer. J. Bot. 18:728—738.
————. 1935. Studies in Onagraceae IX. The subgenus Raimannia. Amer. J. Bot.

22:645-663.
. 1941. Interesting western plants. Leafl. W. Bot. 3:49-53.
————. 1965. Onagraceae. North American Flora IT. 5:1-278.
Raven. P. H. 1964. The generic subdivision of Onagraceae, tribe Onagreae. Brit-
tonia 16:276-288.

POLLEN APERTURE VARIATION AND PHYLOGENY IN
DICENTRA (FUMARIACEAE)

KINGSLEY R. STERN

Dicentra Bernh., comprising some 20 species of perennial and biennial
herbs and climbers of North American and East Asian distribution, was
monographed by Hutchinson (1921) as part of a larger treatment.
Fedde (1936) largely followed Hutchinson’s treatment in his discussion
of the Papaveraceae, although both earlier works were incomplete. In
my revision of the genus (1961; 1967), phylogenetic trends, based pri-
marily on morphological and anatomical features, were discussed. Berg
(1964), studying seed dispersal ecology in Dicentra independently,
reached essentially similar conclusions about the intrageneric phylogeny,
as did Fahselt and Ownbey (1968) while investigating the flavonoid
components. Cytological evidence obtained by Ryberg (1960), Ernst
(1965), Stern (1968) and others suggests the development of a poly-
ploid series accompanying morphological and chemical advancement,
but further extensive study is needed before the role of polyploidy in the
evolution of the genus, and cytotaxonomic interrelationships in general
can be clearly portrayed.

After brief mention of pollen morphology in my 1961 monograph, I
studied Dicentra pollen grains in more detail (Stern, 1962), and found
the interspecific variation not only extensive, but specifically constant
enough to permit distinguishing between all except two of the species on
the basis of pollen morphology alone. Such interspecific variation is ex-
ceptional, although not wholly unique, as the representative studies of
Dahl (1952), Fasbender (1959), Helmich (1963) and Lewis (1965)
suggest. My 1962 study included descriptions and dimensions of the
pollen grains and mention of phylogenetic trends. This study amplifies

1970] STERN: DICENTRA 255)

and details the extensive aperture variation found, indicates correlations
between pollen morphology and other features, and, on the basis of new
evidence, revises some phylogenetic concepts.

Pollen grains for this study were mounted in lactic acid (ca. 85%),
after removal from herbarium specimens, and slides were made semi-
permanent by the addition of ringing cement and cover glasses. Addi-
tional mounts in Dahl’s medium (Stern, 1961), Calberla’s solution, sili-
cone oil (Anderson, 1960) and glycerine jelly, following acetolysis,
(Erdtman, 1960) were made for comparison, although it was found
that the latter preparations were of more value in exine studies than
in aperture studies. A duplicate set of acetolyzed pollen slides has been
deposited in the collections of the Palynologiska Laboratoriet, Stockholm-
Solna, Sweden.

D. burmanica Stern: Kaulback 267 (BM, E).

D. canadensis (Goldie) Walp.: Hone 179 (MIN); Shafer 130 (UC);
Stern 190 (UC); Stern 192 (UC); Umbach 1570 (S).

D. chrysantha (H. & A.) Walp.: Bacigalupi & Holmgren 3179
(UC); Meyer 745 (UC); Sharsmith 4277 (S); Sowder 431 (UC);
Stern 157 (MIN); Van Dyke s.n. (CAS, F, POM).

D. cucullaria (L.) Bernh.: Anderson 661 (UC); Bush 13228 (S);
Nielsen 2399 (MIN); Stern 191 (UC); Stern 193 (UC); Umbach s.n.
(F, MICH, UC, US, WIS).

D. eximia (Ker) Torr.: McVaugh 5714 (UC); Stern 2021 (UC);
Stern 197 (UC); Stern 202 (UC).

D. formosa (Haw.) Walp.: Brown s.n. (MIN); Everett & Balls
9458 (S); Henry sn. (DS); Kruckeberg 4990 (UC); Leach & Leach
1360 (ORE) ); Stern 775 (UC).

D. grandifoliolata (Merrill) Stern: Ward 143 (Vernay-Cutting
Expdn.) (GH, NY).

D. lichiagensis Fedde: Handel-Mazzetti 4329 (GH, US); Maire
3265B (UC); Schneider 2004 (B, US); Tsai 56060 (AAH).

D. macrantha Oliv.: Forrest 26601 (E, NY, W, US); Smith 2098
(CUES)

D. macrocapnos Prain: El.C. Kew Distr. No. 119 (GH, K, L, LE,
M, S); Stainton, Svkes & Williams 4254 (BM); Stainton, Svkes &
Williams 5009 (BM).

D. nevadensis Eastw.: Cronquist 2148 (MO); Darland s.n. (UC);
Stern 166 (MIN).

D. ochroleuca Engelm.: Clokey & Templeton 4615 (UC); French
332 (UC); Gifford 195 (UC); Howell 4079 (CAS); Pollard s.n. (S);
Stern 158 (MIN).

D. pauciflora Wats.: Brown 418 (UC); Haddock 14 (DS, UC); Pur-
pus 3140 (UC); Rowntree s.n. (CAS).

D. paucinervia Stern: Ludlow & Sheriff 15838 (BM).

356 MADRONO [Vol. 20

D. peregrina (Rudolph) Makino: Hiroe 7054 (UC); Jochelson 228
ENDS

D. rovlei Hook. f. & Th.: Lace 1516 (E); Ten 1367 (B).

D. scandens (D. Don) Walp.: Ownbey, s.n. (WS); Schneider 3244
(B); Tsai 52955 (GH).

D. spectabilis (L.) Lem.: Bazilevski s.n. (LE); Maire 2714 (UC);
Umbach s.n. (WIS, UC, F).

D. torulosa Hook f. & Th.: Cooper 3129 (E); Ducloux 948 (E);
Maire 725 (BM, E); Tsiang 8866 (UC).

D. uniflora Kell.: G. N. Jones 9935 (GH); M. E. Jones s.n. (POM,
UC); Steward & Gilkey (OSC).

Figure 1 illustrates, via diagrams, the various aperture arrangements
occurring in pollen grains of Dicentra species. The pollen diagrams
themselves are superimposed on a diagram revised and adapted from
Stern (1961), which indicates presumed relationships between species,
based on advancement indices derived primarily from morphology and
anatomy. As observed by Alston and Turner (1963), such base diagrams
do not indicate the factor of time for the assumed branching, since the
angles of divergence, etc., are strictly diagrammatic, and are not in-
tended to signify constant rates of evolution. Nevertheless, in the ab-
sence of extensive genetic and experimental evidence. they do serve a
useful purpose as a framework for future investigation. As indicated
earlier, since the base diagram appeared in its original form, support for
many of the phylogenetic positions indicated has been derived from seed
dispersal ecology studies and chemotaxonomy. An exception to this is
the position of D. spectabilis, which was originally included in the sub-
genus Chrysocapnos Engelm. A reconsideration of the pollen exine mor-
phology suggests its affinities lie closer to members of the subgenus Di-
centra, and the chemical evidence presented by Fahselt and Ownbey
(1968) tends to substantiate this. The morphology and anatomy of
the species, however, is sufficiently distinctive to warrant its relegation
to a monotypic subgeneric ranking of its own. Dicentra macrantha also
is here accorded subgeneric ranking, since its floral morphology differs
so markedly from that of other members of the subgenus Chrysocap-
nos; further, although it is not scandent, it does appear, vegetatively, to
be more closely related to members of the subgenus Dactvlicapnos
(Wall.) Stern.

Subgenus Hedycapnos (Planch.) Stern, stat. nov. Capnorchis subg.
Hedycapnos Planch. Fl. Serres 8:193. 1853. Eucapnos Sieb. & Zucc.
Abh. Math.-Phys. Cl. Konigl. Bayer. Akad. Wiss. 3:721. 1840, non
Bernh. Linnaea 8:468. 1833. Dicentra subg. Chrysocapnos sect. Hedy-
capnos (Planch.) Stern, Brittonia 13:21. 1961. Type species: Dicentra
spectabilis (L.) Lem. FI. Serres I. 3:pl. 258. 1847.

Fic. 1. Diagram indicating presumed phylogenetic relationships and correlations
of pollen aperture types in Dicentra.

STERN: DICENTRA Sov

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358 MADRONO [Vol. 20

Subgenus Macranthos (Stern) Stern, stat. nov. Dicentra subg. Chryso-
carpnos sect. Macranthos Stern, Brittonia 13:24. 1961. Type species:
Dicentra macrantha Oliv. For further discussion and species synonymy
see Stern (1961).

When the pollen aperture diagrams are added to the presumed phy-
logeny base diagram, (fig. 1), certain correlations between gross mor-
phology and pollen morphology become apparent. The more primitive
species possess numerically constant (3 or 6) apertures, which are also
distinct. Dicentra ochroleuca, and occasionally D. macrantha, do, in
addition to the basic 3 or 6 apertures, exhibit anomotreme pollen grains,
often with bizarre aperture configurations, which are, however, always
distinct. The proportion of aperture to non-aperture surface area in these
and other Dicentra pollen grains appears to be more or less constant,
regardless of the particular configurations or numbers of apertures. To
demonstrate this mathematically in prolate spheroids with so many
grain-to-grain variables would, however, be a most challenging task.

The more advanced species, in the subgenera Dicentra and Dactyli-
capnos, in general, have numerically inconstant apertures, and, in the
latter subgenus in particular, the apertures become less distinct. Also,
in the subgenus Dicentra, there is a trend toward more numerous aper-
tures, and eventual fusion of the apertures. Although an occasional
anomotreme pollen grain will appear in the four most primitive species
(D. formosa, D. nevadensis, D. eximia and D. peregrina), fusion of
apertures has not been observed. In D. pauciflora, however, some pollen
grains are 8-aperturate, the apertures consisting of 6 separate rugae or
colpi, plus 6 more rugae coalesced into 2 triradiate apertures. In D. can-
adensis, some basically 12-aperturate pollen grains become synapertur-
ate, and in D. cucullaria, various configurations, but all synperturate,
are typical. If increase in numbers of apertures, and fusion of apertures,
as well as decreases in the distinctness of aperture margins in the pollen
grains of Dicentra may be considered advancement, such advancement
appears to have accompanied morphological and anatomical advance-
ment in the genus.

This was part of a larger study, for which partial support from the
National Science Foundation, under grant GB-4498, is gratefully ac-
knowledged. The author is also grateful to G. Erdtman for the use of
the facilities of the Palynologiska Laboratoriet, in Stockholm-Solna,
Sweden.

Department of Biological Sciences, Chico State Ccllege, Chico, California

LITERATURE CITED

ANDERSON, S. T. 1960. Silicone oil as a mounting medium for pollen grains. Danske
Geol. Unders@ggelse. IV. 4:1-24.

Aston, R. E., and B. L. Turner. 1963. Biochemical systematics. Prentice Hall,
Englewood Cliffs, N.J.

Berc, R. Y. 1964. Adaptation and evolution in Dicentra, with special reference to
seed, fruit, and dispersal mechanism. Amer. J. Bot. 51:688.

1970] BANWAR: LYONOTHAMNUS 359

Dau, A. O. 1952. The comparative morphology of the Icacinaceae, VI. The Pollen.
J. Arnold. Arbor. 33:252-295.

ErpTMAN, G. 1960. The acetolysis method. A revised description. Svensk Bot.
Tidskr. 54:561-564.

Ernst, W. R. 1965. Documented chromosome numbers of plants. Madrono 18:24.

FanseEtt, D., and M. Ownsey. 1968. Chromatographic comparison of Dicentra
species and hybrids. Amer. J. Bot. 55:334—-345.

FasBENDER, M. V. 1959. Pollen grain morphology and its taxonomic significance in
the Amherstieae, Cynometreae, and Sclerobieae (Caesalpiniaceae) with special
reference to American genera. Lloydia 22:107-162.

FeppE, F. 1936. Papaveraceae-Fumarioideae. Jn Engler, Natiirlichen Pflanzenfamilien
17c:121-145.

Hertmicu, D. E. 1963. Pollen morphology in the maples. Pap. Michigan Acad. Sci.
48:151-164.

HutTcHInson, J. 1921. The genera of the Fumariaceae and their distribution. Kew
Bull. 1921:97-115.

Lewis, W. H. 1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia
(Rubiaceae). Amer. J. Bot. 52:257-264.

Ryserc, M. 1960. A morphological study of the Fumariaceae and the taxonomic
significance of the characters examined. Acta Hort. Berg. 19:121-248.

STERN, K. R. 1961. Revision of Dicentra (Fumariaceae). Brittonia 13:1-—57.

. 1962. The use of pollen morphology in the taxonomy of Dicentra. Amer.
J. Bot. 362-368.

. 1967. A new species of Dicentra from Burma. Brittonia 19:280—282.

. 1968. Cytogeographic studies in Dicentra I. Dicentra formosa and D.
nevadensis. Amer. J. Bot. 55:626-628.

FOSSIL LEAVES OF LYONOTHAMNUS
SATISH C. BANWAR

Leaves of the extant genera Lyonothamnus, belonging to the family
Rosaceae, and Comptonia, belonging to the family Myricaceae, are very
similar in external appearance. Many paleobotanists who have examined
fossil leaves of Lyvonothamnus were at times led to identify them as
Comptonia. This study was conducted to examine and compare leaves
of Lvonothamnus, both extinct and extant, and those of Comptonia, so
that differences and similarities in shape, nature, and venation could be
established, which would then help to distinguish them.

Fossil leaves, identified as those of Lyonothamnus, have been col-
lected in various localities in the western United States from Washing-
ton to Oregon, California, and Nevada. The ages of all these fossil leaves
range from Miocene to Pliocene. All the specimens collected so far are
comparable to leaves of the extant L. floribundus Gray ssp. asplenifolius
(Greene) Raven. The leaves of subspecies asplenifolius are so distinctive
that similarities with the fossil forms are easily recognized (figs. 2, 4).
So far, to my knowledge, no one has reported the presence of fossil leaves
which may be comparable to foliage of subspecies floribundus (fig. 1).

The first fossil specimens to be identified as Lyonothamnus were col-
lected by Axelrod in 1939 from the Tehachapi area of California; he

360 MADRONO [Vol. 20

AGRO ERS
‘ st GEN Ne
WSS Se we
(A Sorcerer
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pentane:
; 2

1970] BANWAR: LYONOTHAMNUS 361

described and named these specimens as L. mohavensis (Axelrod, 1939).
The age of these specimens was determined as Middle Miocene; they
were in small fragments and only portions of leaflets were present, but
the general appearance, shape, and venation were similar to those of
leaves of living L. floribundus ssp. apslenifolius. Since then Axelrod has
reported the occurence of L. mohavensis from many other localities in
California and Nevada. The sites and their ages are as follows: Furnace
Creek Flora, California (Mio-pliocene), Mint Canyon Flora, California
(Upper Miocene), Upper Modella Flora, California (Upper Miocene),
Coal Valley Flora, Nevada (Upper Miocene), Mulholland Flora, Cali-
fornia (Middle Pliocene) (Axelrod, 1940). In 1956, while examining the
Mio-pliocene floras from west-central Nevada, the same author described
many fossil specimens as C. parvifolia, because to him they resembled
the leaves of members of the extant genus Comptonia (Axelrod, 1958).
These leaf specimens reported as Comptonia were much larger and bet-
ter preserved than those reported earlier as Lyonothamnus.

Wolfe, while investigating the Miocene floras from Fingerrock Wash,
Nevada, reassigned C. parvifolia to L. parvifolia (Axelrod) Wolfe. He
justified his reassignment on the basis of venation pattern (Wolfe, 1964).
Wolfe also believed he could distinguish two forms of fossil Lyonotham-
nus: one form having seven to nine leaflets or pinnae and the other
fewer than seven leaflets. The former represents L. parvifolia, the latter
L. mohavensis. He was of the opinion that these two forms should be
regarded as two different species, and that L. mohavensis is closer to the
extant forms.

Page (1964) has recently described as Lyonothamnoxylon nevaden-
sis, fossil wood from the lower Pliocene of Nevada. The wood of L.
nevadensis most closely resembles that of Lyonothamnus.

The genus Lyonothamnus comprises a single living species with two
subspecies (Raven, 1963), namely LZ. floribundus Gray ssp. floribundus
and L. floribundus ssp. asplenifolius (Greene) Raven. The genus is en-
demic to the islands off the shores of southern California; no natural
population exists on the mainland. The distribution of the subspecies on
the islands is also very interesting. Subspecies floribundus is confined to
Santa Catalina I., while ssp. asplenifolius is restricted to Santa Rosa,
Santa Cruz, and San Clemente islands. The two subspecies are distin-
guished from each other by the nature and form of the leaves they bear.
The leaves of ssp. floribundus are simple, lanceolate, and have an entire
margin (fig. 1), while those of ssp. asplenifolius are pinnately compound

Fics. 1-7. Lyonothamnus and Comptonia: 1, leaf of L. floribundus ssp. flori-
bundus; 2, leaf of L. floribundus ssp. asplenifolius; 3, leaf of C. peregrina showing
the lobes (Transylvania Co., North Carolina, Fox 704, UC); 4 fossil specimen of
L. parvifolia, collected by the author at Stewart Springs area, Mineral Co., Nevada;
5, a leaf lobe of L. floribundus ssp. asplenifolius, showing venation pattern; 6, por-
tion of leaf C. peregrina, showing venation of lobes; and 7, a leaf lobe of L.
parvifolia, showing venation pattern.

362 MADRONO [Vel. 20

and have a dissected lamina (fig. 2).

The members of the genus Comptonia are confined in their distribu-
tion to the eastern United States. The leaves of C. peregrina (L.) Coult.
(fig. 3) are simple, and the laminae are dissected somewhat like those of
L. floribundus ssp. asplenifolius.

MATERIALS AND METHODS

Fossil specimens were examined in the excellent collections of the
Museum of Paleontology, University of California, Berkeley. Some
specimens were collected personally from Stewart Springs, Mineral Co.,
Nevada. They [fossil leaves] were examined under a dissecting micro-
scope, and the venation patterns were observed by wetting the specimens
with clove oil.

Leaves of living plants of Lyonothamnus were obtained from the
Rancho Santa Ana Botanical Gardens at Claremont and from the Uni-
versity of California Campus at Berkeley. At both of these places the
trees are cultivated. Leaves of C. peregrina were obtained from herbar-
ium specimens housed in the University of California Herbarium, Berke-
ley.

The leaves of Lyonothamnus and Comptonia were found to be very
rich in tannin and thus the clearing technique had to be modified slightly.
Both dried and fresh leaves were boiled in water and then in alcohol to
remove the chloroplasts. The leaves were then washed in running tap
water for some time and soaked in My-pro Bleach (manufactured by
Crown Products, San Francisco), which removed all the tannin and
made the leaves white. The timing in this bleach had to be adjusted for
each leaf because a prolonged treatment in bleach macerated all the
tissues. After this the leaves were washed again in tap water for several
hours and then soaked in a saturated solution of chloral hydrate. They
were then washed in water and stained in the usual way with | percent
alcoholic safranin. It was found that dried plant materials yielded better
results than did similarly treated living materials. The leaves were
mounted in Picolyte on large glass plates.

OBSERVATIONS

Most of the fossil specimens examined are only fragments of large
leaves. Some of these fragments are fairly large while others are small,
but even from the latter it was evident that the original laminae were
pinnately compound and that each pinna had several dissected lobes.
The definite number of pinnae present on each leaf could not be deter-
mined because of the small size of the fragments or their arrangement.
Some of them have as few as three pinnae, while others have as many as
nine. The lobes are not exactly opposite each other on a pinna but are
“staggered” or arranged alternately. Each lobe is somewhat triangular in
shape with an acute pointed apex. The lobes are sessile and attached by
one complete side to the axis of the pinna (fig. 4).

1970] BANWAR: LYONOTHAMNUS 363

In each lobe, several secondary veins are given off from the axis of the
pinna. These extend nearly to the margin of the lobe. Each secondary
vein produces several tertiary veins, which also divide further and form
a fine network of veins. The finer details and the details of the vein end-
ings could not be examined as they were not well preserved in the fossils.
Among th secondary veins, it is the uppermost (or first) secondary vein
which terminates in the apex of the lobe, while the other secondary veins
form a loop at the distal margin and join with the secondary vein above
(fig. 7).

In the genus Lyonothamnus, only leaves of ssp. asplentfolius were ex-
amined in detail because it is in this subspecies that similarities have
been noted with the various fossil leaves mentioned and with leaves of
Com ptonia.

Leaves of L. floribundus ssp. asplenifolius are opposite (although
members of the Rosaceae are usually considered to have alternate
leaves) and pinnately compound with several pinnae (fig. 2). The
number of pinnae varies between three and seven on different trees and
even on different branches of the same tree. No leaf was observed to
have as many as nine pinnae. Each pinna is divided into several lobes,
the sinuses extending clear into the axis of the pinna. These are slightly
staggered or alternate to each other on the axis of the pinna, and are
sessile and triangular in outline. Each lobe has an acute apex and one
complete side is attached to the axis of the pinna (fig. 2). In each lobe,
several secondary veins are given off from the axis of the pinna and ex-
tend nearly to the margin of the lobe. Each secondary vein produces sev-
eral branches which divide further and produce a fine network of veins
with free vein endings. It is the uppermost (first) secondary vein which
terminates in the apex of each lobe (fig. 5), while the other secondary
veins form loops near the distal margin and join with the adjacent sec-
ondary vein towards the apex.

Leaves of C. peregrina are alternate and simple. The lamina is dissect-
ed into nearly opposite lobes, but the sinuses do not extend all the way
to the midrib (figs. 3, 6). The size of the leaves varies and so also does
the number of the lobes. But it is evident that the lobes are very dif-
ferent from those of Lyonothamnus. Each lobe has an obtuse apex. Gen-
erally three secondary veins develop in each lobe, the first extending
only to the sinus, and (in contrast to the situation in Lyonothamnus )
the second or central secondary vein terminating in the apex of the
lobe (fig. 6). Each secondary vein produces several tertiary veins which
divide further and produce a fine net-work of veins with free vein end-
ings.

DISCUSSION

From the external morphology of the leaves studied, it is very clear
that the fossil leaves of Lyonothamnus are very similar to those of the
extant ssp. asplenifolius. The similarity with leaves of Comptonia is

364 MADRONO [Vol. 20

very superficial. The leaves of Comptonia are simple, the incisions of the
lamina are shallow, and the lobes are opposite or nearly so, while those
of Lyonothamnus are compound with a number of pinnae and the in-
cisions of the lamina are deep, extending right up to the axis of the
pinna. The shape, position, and nature of the lobes and their apices are
also very different.

The venation patterns are also distinct. The venation of both extant
and fossil leaves of Lyonothamnus examined were very similar. In both
it is the first, or uppermost, secondary vein which terminates in the apex
of the lobe. In Comptonia it is the second, or the central, secondary vein
which does so. Thus on the basis of venation pattern also the fossil
leaves show a greater similarity with those of the extant ssp. aspleni-
folius. The similarity with those of Comptonia is very superficial.

This study agrees with the views of Wolfe in reassigning all the fossil
leaves described as C. parvifolia Axelrod to Lyonothamnus parvifolia
(Axelrod) Wolfe, although the creation of two species among the fossil
leaves is debatable.

The author wishes to thank Lincoln Constance, A. S. Foster, and
W. L. Fry, of the University of California, Berkeley, for guiding and
helping me in this study. This study was a part of the doctoral thesis
submitted at the University of California, Berkeley. The author is also
very thankful to the directors of the Paleontological Museum and
Herbarium of the University of California, Berkeley, for allowing the
use of their collections.

Hearngunje, Hazaribagh, Bihar, India

LITERATURE CITED

AXELROD, D. I. 1939. A Miocene flora from the western borders of the Mohave

Desert. Publ. Carnegie Inst. Wash. 516.
. 1940. A record of Lyonothamnus in Death Valley, California. J. Geol.

48:526-531.

———. 1958. Evolution of the Madro-Tertiary geoflora. Bot. Rev. (Landcaster)
24:433-509.

Pace, V. M. 1964. Lyonothamyxylon from the lower Pliocene of Nevada. Madrono
17:257-266.

Raven, P. H. 1963. A flora of San Clemente Island, California. Aliso 5:289-347.

Wo Fe, J. 1964. Miocene floras from Fingerrock Wash, southwestern Nevada. U. S.
Geol. Survey Prof. Pap. 454N.

NOTES AND NEWS

EpirorsHie OF MaproNo.—Some time during the summer of 1971, Dr. Robert
Ornduff, Department of Botany, University of California, Berkeley, will become
the Editor of Madrofo. At the same time, Dr. John Strother, also at Berkeley, will
become Managing Editor. In preparation for the change of Editors, it is requested
that all new manuscripts intended for publication in Madrono be sent to Berkeley
rather than Stanford.

CHROMOSOME STUDIES IN MELAMPODIUM
(COMPOSITAE, HELIANTHEAE)

Top F. STUESSY

A number of chromosome counts for the genus Melam podium already
have been reported (Negodi, 1938; Jackson, 1957; Turner, Beaman,
and Rock, 1961; Turner and Johnston, 1961; Turner and King, 1962;
1964; Turner and Flyr, 1966). These studies surveyed 21 of the 35
presently recognized in the genus (Stuessy, 1968) and indicated the pres-
ence of intraspecific polyploidy in several taxa (Turner and King, 1962).
In addition, base numbers for the genus have been tentatively proposed
as x = 9 (Negodi, 1938) and x = 10 (Turner and King, 1962). The
present studies significantly add to previous work by 1, most importantly,
putting the old counts into the framework of the recent taxonomic con-
cepts and nomenclature (Stuessy, 1968), 2, adding new counts for two
species, 3, surveying intra- and inter-populational chromosome variabil-
ity throughout the ranges of many species which more clearly indicates
the significance of polyploidy in each species and in the whole genus,
and 4, clarifying certain problems mentioned in earlier chromsome treat-
ments.

METHODS AND MATERIALS. Buds for meiotic counts were collected in
the field in modified Carnoy’s fixative, 4 parts chloroform, 3 parts abso-
lute alcohol, and 1 part glacial acetic acid, and refrigerated in the labo-
ratory (Walters, 1952), until subsequently counted by acetocarmine
squash techniques. Material stored in this manner lasted as long as one
year and still produced clear meiotic figures, although some hardening of
the cells was noticed.

Seeds for the few mitotic counts were treated as in Speese and Bald-
win (1952), first being germinated on filter paper in petri dishes, pre-
treated for an hour in a saturated aqueous solution of PDB, then fixed
in 3:1:1 (3 absolute alcohol: 1 chloroform: 1 glacial acetic acid) and
finally squashed under acetocarmine stain.

RESULTS AND Discussion. Table 1 lists the various chromosome
counts known for the genus Melam podium. Due to problematical nomen-
clature, many previous counts were reported under names now regarded
as synonyms (Stuessy, 1968). In addition, the recognition of new vari-
eties in M. cinereum, M. leucanthum and M. montanum (Stuessy, 1968)
has required putting old counts referred to these species into the pro-
posed respective varietal categories.

Two species, M. glabrum and M. hispidum, are here reported for the
first time asn = II (fig. 1) and n = 20 (fig. 2), respectively. Melam-
podium glabrum thus coincides cytologically with the morphologically
smilar species, M. perfoliatum (n = 11, 12), which strengthens the

365

366 MADRONO [Vol. 20

Fics. 1-3. Camera lucida drawings of chromosomes of species of Melampodium:
1, M. glabrum, diakinesis, n= 11, Stuessy 708; 2, M. hispidum, metaphase II
(% of cell drawn, n= 20, Stuessy 1038; and 3, M. sericeum, diakinesis, n = 30,
Stuessy 364.

inferred close phyletic relationship of these two taxa. Melampodium his-
pidum is a mountainous species very similar morphologically to M. seri-
ceum (n = 30; 20 large and 10 small bivalents, fig. 3). It may be that
an ancestor of M. hispidum was involved in the evolution of this hexa-
ploid species, perhaps contributing the 20 larger bivalents. The close
morphological similarity of M. sericeum to another species, M. sp. nov.
(P. Goldsmith 133; chromosomally unknown but placed in the x = 10
section of the genus), suggests that an ancestor of the latter may have
donated the smaller set of 10 bivalents. This speculative origin of M.
sericeum contrasts markedly with that indicated by Turner and King
(1962). While they noted the close relationship to M. hispidum (first
perceived by Robinson, 1901), they suggested the possible involvement
of an ancestor of M. camphoratum (n = 16) by the incorporation of
some of its small chromosomes into the genome of the incipient VM. seri-
ceum through amphiploidy. But since M. camphoratum, in my opinion,
belongs to the rather distantly related genus Unxia (closely related to
Polymia), the likelihood of such an origin seems remote.

Extensive surveying of many populations of species previously counted
(118 new population reports) has shown that all but three species are ap-
parently uniformly diploid throughout most of their ranges. The diploid-
tetraploid races in M. cinereum and M. leucanthum noted by Turner and
King (1962) have been verified in the present study (table 1) and will
be discussed at length elsewhere. Melampodium dicoelocarpum is the
only other species that has been found to possess polyploid races, being
diploid at n = 12 (new report, fig. 4) and aneuploid (at the tetraploid
level) with n — 23 (fig. 5). Although quantitative morphological differ-
ences are present between the latter two races, too few counts have been
made and too few herbarium records are available to comment on sig-
nificant geographical or ecological differences.

Melampodium longipilum has been counted previously only once

1970] STUESSY: MELAMPODIUM 367

: o « °

Fics. 4-9. Photographs of chromosomes of species of Melampodium: 4, 5, M.
dicoelocarpum, metaphase II; 4, n= 12, Stuessy 693; 5, n= 23, Stuessy 716;
6-8, M. longipilum; 6, 7, anaphase I, n= 10, Stuessy 373; 8, metaphase, 2n = 20,
Stuessy 634; and 9, M. perfoliatum, diakinesis, n = 11, Stuessy 379. All approxi-
mately 1800.

(Turner and King, 1962), the authors commenting that, ‘This collec-
tion is interesting in that its habit and floral features are similar to M.
divaricatum, but the achenes bear well-developed hoods such as are char-
acteristic of the section Melam podium. Its chromosome number, n = II,
however, would be exceptional for the latter section.” Although on close
examination this species is quite distinct within the genus, it seems mor-
phologically closest to M. diffusum (n = 10) and on this basis would be
expected to fall into the section Melampodium. Since meiotic (and one
mitotic) counts have been made from six populations of this species in the
present study (table 1) and all have yielded unequivocal counts of
n — 10 (figs. 6-8), it is probable that the normal chromosome number
of this species isn = 10.

The reported counts of M. perfoliatum (Turner and King, 1962;
1964) have been both n = II and n = 12. Although the present study
recorded only counts of n = 11 for this species (fig. 9), more survey
work is needed to discover the factors involved in the establishment and
maintenance of these two chromosomal levels, if two indeed exist. The

368 MADRONO [Vol. 20

chromosomal voucher specimens of the two races cannot be distin-
guished morphologically.

The early counts of n = 10 for both M. divaricatum and M. perfoli-
atum reported by Negodi (1938) accompanied by descriptions and pho-
tographs of the plants, contrast with the consistent subsequent counts
of other workers of n = 12 and n = I] and 12, respectively. It is likely
that these unusual n = 10 counts represent either very anomalous con-
ditions or perhaps erroneous observations.

Negodi (1938) was the first to discuss the taxonomy of Melam podium
in a phyletic sense. Based on counts of n = 9 and n — 10 for three spe-
cies, he felt that n = 9 was the ancestral base, followed by an aneuploid
gain to n = 10. Turner and King (1962) however, comment that, “It is
obviously impossible to know what the ancestral basic chromosome num-
ber for the genus might have been, but it does seem significant that the
number, n = 10, is found in a wider range of morphological types than
is any other number.” The fact that a large number of morphologically
diverse species within a genus has a certain characteristic (e.g., x — 10)
by no means designates this unequivocally as a primitive trait. But it
may be suggestive, especially in recently evolved groups such as the
Compositae. Nevertheless, phylogenetic speculations including base
number hypotheses must be based on all available evidence, not numer-
ology or one or two characters alone. Further insight into aneuploid
trends in Melam podium must wait for a compilation of evidence that is
accumulating on other aspects of the genus.

Field work for this investigation was supported in part by NSF
Traineeship 4128, NSF grant GB-1428, and a Sigma Xi Grant-in-Aid.
Appreciation is expressed to P. H. Raven and B. L. Turner for per-
mission to include several unpublished chromosome counts, and to R. S.
Irving, J. L. Strother, and B. L. Turner for several bud collections. This
study represents a portion of a dissertation (supervised by B. L. Turner)
submitted to the Graduate School of the University of Texas at Austin
in partial fulfillment of the requirements for the degree of Doctor of
Philosophy.

Note added in proof: M. hispidum has been counted recently as
n — 20 by Powell and Sikes (1970).

TABLE 1. CHROMOSOME COUNTS IN MELAMPODIUM

All voucher specimens cited in this study are deposited in the University of
Texas Herbarium, Austin.
All citations are meiotic counts showing clear bivalents unless otherwise indi-
cated at the end of each voucher citation.
Superscripts after voucher specimens refer to counts not made by the author
but found in the following references:
* Negodi (1938)
b Jackson (1957)
© Turner, Beaman and Rock (1961)

1970] STUESSY: MELAMPODIUM 369

d Turner and Johnston (1961)

e Turner and King (1962)

! Turner and King (1964)

h Powell, A. M., and B. L. Turner (unpublished)

kK Raven, P. H. (unpublished)

m Turner, B. L. (unpublished)

1 Turner, B. L., W. L. Ellison, and R. M. King (unpublished)

Different voucher numbers from the same locality refer to counts from buds
from an individual plant (listed first) and from a populational sample. Irving,
Stuessy and Turner collections of M. cinereum and M. leucanthum, however, are
all individual plants.

M. americanum L. n= 10, GUATEMALA. Baja Verapaz: near Salama, King
3260 (reported as M. americanum var.)°; Progreso: 35 mi NE of Guatemala,
Stuessy 602. MEXICO. Chiapas: Santa Isabel, Stuessy 632; Colima: Alzada,
Stuessy 727; Guerrero: 9 mi NW of Taxco, King 4168 (reported as M. kunthi-
anum)°; 25 mi NE of Acapulco, King 4178 (reported as M. kunthianum)®° ; Michoa-
can: 7 mi S of Ario de Rosales, Stuessy 688,689; Nayarit: 25 mi N of Tepic, King
3699; Vera Cruz: 20 mi. E of Cuitlahuac, King 2679°; 26 mi E of Cuitlahuac, King
2682°; 9 mi NW of Alvarado, King 2709°; 9 mi SE of Alvarado, King 2718; 24 mi
E of Cuitlahuac, Stwessy 314, 315; 27 mi S of jct rtes 110 & 105; Stuwessy 469, 470
(ca 10); 19 mi S cf Diamante, Stuessy 481; 34 mi NW of José Cardel, Stuessy
484, 485; 14 mi E of La Tinaja, Stwessy 516; Salinas, Stuessy 518, 519; near Cat-
emaco, Stuessy 522.

M. aureum Brandg. n = 33. MEXICO. Michoacan: 20 mi W of Ciudad Hidalgo,
King 3617°; 21 mi NW of Ciudad Hidalgo, Stuessy 683, 684; Oaxaca: 7 mi NE of
Nochistlan, Stuessy 663 (& frag.).

M. cinereum DC. var. cinereum. n = 10. MEXICO. Tamaulipas: 59 N of
Sabinas Hidalgo, Stwessy 857a. TEXAS. Hidalgo Co.: 6 mi E of Sullivan City,
Turner 4490 (reported as M. cinereum)*; Webb Co.: ca 22 mi NW of ject rtes 83
& 81 (35), Stuessy 869; 37 mi N of Zapata, Thompson 174 (reported as M. cin-
ereum®; Zapata Co.: 14.6 mi N of San Ygnacio, Strother 556; 27 mi N of Zapata,
Thompson 175 (reported as M. cinereum)*®; Zavala Co.: 11 mi SE of Batesville,
Sullivan & Turner 22 (reported as M. cinereum)*; 6 mi S of Batesville, Turner
5006 (reported as M. cinereum)®.

M. cinereum DC. var. cinereum. n = 20, TEXAS. Duval Co.: 25 mi N of Heb-
bronville, Stwessy 429; Jim Hogg Co.: 10 mi E of Hebbronville, Stuessy 423
(ca 20) ; near Hebbrcnville, Stuessy 425; Hebbronville, Stuessy 426; near Hebbron-
ville, Stwessy 428; Hebbronville, Thompson 177 (repcrted as M. cinereum)*; Jim
Wells Co.: near Orange Grove, Strother 565; Live Oak Ca.: ca 32 mi S of Whitsett,
Stuessy 772, 773; 14 mi S of George West, Thompson 180 (reported as M. ciner-
eum)®*; Zapata Co.: 17 mi NE of Zapata, Thompson 176 (reported as M. ciner-
eum) ©

M. cinereum DC. var. nov. n= 10. MEXICO. Coahuila: near Nueva Rosita,
Stuessy 902a; 21 mi S of Monclova, Stuessy 912; Nuevo Leon: 15 mi N of Sabinas
Hidalgo, Stuessy 854; 26 mi N of Sabinas Hidalgo, Stuessy 855a; 38 mi N of
Sabinas Hidalgo, Stwessy 856a.

M. cinereum DC. var. ramosissimum (DC.) A. Gray. n = 10. MEXICO. Tamaul-
ipas: San Fernando, Stuessy 450, 541; Reynosa, Stuessy 778, 779; 27 mi S of
Reynosa, Stuessy 787.

M. cupulatum A. Gray. MEXICO. Sinaloa: near Culiacan, Flyr 112 (reported
as M. rosez)*.

M. dicoelocarpum Rob. n= 12. MEXICO. Michoacan: 25 mi S of Ario de

370 MADRONO [Vol. 20

Resales, Stuessy 693 (fig. 4).

M. dicoelocarpum Rob. n = 23, MEXICO. Michcacan: near Cotija, King &
Soderstrom 4646 (reported as M. microcephalum)®; 15 mi S of jct & rtes 15 & rd to
Cotija, Stuessy 715, 716 (fig. 5).

M. diffusum Cass. n= 10. MEXICO. Guerrero: 26 mi S of Acapulco, Powell
758°; Acapulco, Stuessy 366.

M. divaricatum (Rich. in Pers.) DC. n=10. Plants obtained from bot. gard.
Goteborg".

M. divaricatum (Rich. in Pers.) DC. n= 12. EL SALVADOR. Santa Ana: near
Santa Ana, Stuessy 609. GUATEMALA. Alta Verapaz: 28 mi E of San Miguel
Uspantan, Stuessy 588; Jutiapa: 25 mi E of Cuilapa, Stwessy 605. MEXICO. Cam-
peche: Champoton, Stuessy 532; Chiapas: Tapachula, Stuessy 626; Morelos: 6 mi
NW of Cuautla, Stuessy 351; 10 mi S of Cuernavaca, Stuessy 358, 359; Oaxaca:
Huajuapan de Leon, Stuessy 341; Zimatlan, Stuessy 655; Tabasco: near Villa
Hermosa, Stuessy 547; Vera Cruz: 23.2 mi SE of Alvarado, Stuessy 319; 12 mi
S of Tantoyuca, Stuessy 473 (& 2-3 frag.), 474; 23 mi S of Tecolutla, Stuessy 480;
Jalapa, Stuessy 486; 34 mi NW of Tehuacan, Stwessy 506; Fortin, Stuessy 507;
20 mi S of Alvarado, Stuessy 520 (& 3-5 frag.) ; 49 mi SE of Catemaco, Stuessy 526
(& 2 frag.); many Mexican states: 29 different population counts*®. NICARAGUA.
Granada: Granada, Stwessy 620; Managua: Managua, Stwessy 616; Matagalpa:
Sebaco, Stuessy 614, 615.

M. sp. nov.n = 25+ 1, COSTA RICA. Cartago: Turrialba, King 5348 (reported
as M. cf. flaccidum)".

M. glabrum Wats. n = Il. MEXICO. Jalisco: near La Barca, Stwessy 707, 708
(fig. 1) ; Michoacan: 9 mi S of jct rte 15 & rd to Cotija, Stuessy 714.

M. gracile Less. n= 9. MEXICO. Campeche: Champoton, Stwessy 530, 531; 16
mi N of Champoton, Stuessy 533; Chiapas: 28 mi SE of Comitan, King 3042
(reperted as M. cf. brachyglossum)*; 17 mi S of Tuxtla Gutierrez, King 3096 (re-
ported as M. cf. brachyglossum)®; 32 mi SE of Comitan, Stuessy 573; Michoacan:
near Jiquilpan, King 3636 (reported as M. cf. brachyglossum)*; 3 mi NW of
Zamora, Stuessy 393; 25 mi S of Ario de Rosales, Stuessy 694; Morelos: 7 mi NW
of Cuautla, Stwessy 354, 356; San Luis Potosi: El] Salto, King 3887 (reported as
M. microcarpum)*; Tamaulipas: 6 mi N of Antiguo Morelos, Stuessy 454, 455;
8 mi E of Antiguo Morelos, Stuessy 458, 459; 33 mi N of Ciudad Valles, Stuessy
464; 18 mi § of jet rtes 110 & 105, Stuessy 466; Yucatan: 13 km N of Mérida,
Stuessy 536, 537; Vera Cruz: 13 mi W of Orizaba, Graham & Johnston 4777°, 13
mi W of Orizaba, Johnston 4777 (reported as M. microcarpum)*; 7 mi SW of
Morelos, Powell 646 (reported as M. cf. brachyglossum)®.

M. hispidum H. B. K. n= 20. MEXICO. Chihuahua: Cuauhtémoc,Stuessy 1038
(fig. 2).

M. leucanthum Torr. & A. Gray var. leucanthum. n= 10. ARIZONA. Coconino
Co.: near Sedona, Turner 5738; Gila Co.: 16 mi NW of Globe, Turner 5736;
Pima Co.: near Greaterville, Turner 5735™. COLORADO. Fremont Co.: near Port-
land, Irving 823-1, 823-2, 823-3; near Canon City, Turner 5638; Prowers Co.:
31 mi S of Lamar, Jrving 825. MEXICO Chihuahua: 59 mi N of Villa Ahumada,
Stuessy 1122. NEW MEXICO. Bernalillo Co.: Jackson 2082 (reported as M. leu-
canthum)”; DeBaca Co.: 6 mi E of Yeso, Turner 5673"; Dona Ana Co.: Organ
Mts, San Augustin Pass, Turner 5748; Eddy Co.: near Whites City, Turner 5653;
Hidalgo Co.: 7 mi S of Road Forks, Turner 5719; Santa Fe Co.: 22 mi SW of
Santa Fe, Turner 5676™; Torrance Co.: 3 mi NE of Duran, Raven 19130". OKLA-
HOMA. Cimarron Co.: 6.8 mi N of Cimarron River on rte 287, Irving 824-A,
824-B. TEXAS. Blanco Co.: near Johnson City, Thompson & Graham 17 (re-
ported as M. leucanthum)*; Brewster Co.: Marathon, Stuessy 230, 231; near
Brewster-Pecos Co. line on rte 90, Stwessy 235 (ca 10); Culberson Co.: 6 mi S of
Van Horn, Turner 4738"; El Paso Co.: 28 mi SE of El Paso, Stuessy 1126; Loving

1970| STUESSY: MELAMPODIUM oil

Co.: Mentone, Stuessy 182; Oldham Co.: 16 mi N of Vega, Turner 5632, 56326;
Presidio Co.: near Marfa, Stuessy 201, 202, 203, 204 (& frag.), 206, 207 (ca 10),
213; Redford, Stwessy 227, 228; Travis Co.: near Austin, Thompson & Graham 87
(reported as M. leucanthum)*; Mt. Bonnell, Austin, Stuessy 138; Winkler Co.:
1 mi N into Winkler Co. on rte 18, Stwessy 152, 153, 154; Kermit, Stuessy 167,
168, 169.

M. leucanthum Torr. & Gray var. leucanthum. n = 20, TEXAS. Blanco Co.: 10
mi N of Johnson City, Thompson & Graham 18 (reported as M. leucanthum)® ;
Hays Co.: Dripping Springs, Thompson & Graham 16 (reported as M. leucan-
thum)®; Travis Co.: near Austin, Thompson & Graham 15 (reported as M. leu-
canthum)*; near jct Balcones Rd & 2222, Stuessv 418; Mansfield Dam, Stuessy 420
(ca 20); 7 mi SW of Zilker Pk, Austin, Stuessy 752 (2n = 40). 755-3, 755-4.

M. linearilobum DC. n= 10. EL SALVADOR. San Salvador: 24 mi E of
turnoff to San Vicente, Stuessy 612, 613. GUATEMALA. Jutiapa: 8 mi NE of
Jutiapa, Stwessy 606, 607. MEXICO. Michoacan: Apatzingan, Stuessy 697; Oaxaca:
40 mi W of Tehuantepec, King 2891°; 11 mi E of Zanatepec, King 3449°; 37 mi W
of Tehuantepec, King 3454*°. NICARAGUA. Granada: Granada, Stuessy 618, 619.

M. longifolium Cerv. ex Cav. n = 9. MEXICO. Oaxaca: Las Sedas, Stuessy 659;
San Luis Potosi: 22 mi E of San Luis Potosi, Powell 551%. Plants obtained from
bot. gard. Copenhagen®.

M. longipes (A. Gray) Rob. n = 10. MEXICO. Jalisco: Tequila, King 3662°;
Tequila, Stwessy 396, 737, 738.

M. longipilum Rob. n = 10. MEXICO. Guerrero: 19 mi N of Chilpancingo,
Stuessy 373 (figs. 6, 7), 374; Oaxaca: 13 mi NW of Tehuantepec, Stwessy 328, 329;
3.8 mi NW of Huajuapan de Leon, Stuessy 343; 10 mi NW of Tehuantepec, Stuessy
633, 634 (n = 10 & 2n = 20, fig. 8) ; near Huajuapan de Leon, Stwessy 666; Puebla:
Tehuitzingo, Stuessy 667.

M. longipilum Rob. n = 11. MEXICO. Oaxaca: 64 mi SE of Oaxaca, King
3461 (reported as M. sp. nov.)°.

M. microcephalum Less. n= 9. GUATEMALA. Huehuetenango: 6 mi S of
Huehuetenango, King 3425 (reported as M. oblongtfolium)*. MEXICO. Chiapas:
10 mi SE of Tonala, Stuwessy 627, 628; Michoacan: near Ciudad Hidalgo, King
3607° (reported as M. oblongifolium)*; 6 mi NW of Tuxpan, Stuessy 383, 384;
near Ciudad Hidalgo, Stuessy 680, 681; Oaxaca: Monte Alban, Stuessy 638.

M. montanum Benth. var. montanum. n= 11. MEXICO. Oaxaca: 10 mi N of
ject rtes 190 & 175, King 3492 (ca 11; reported as M. cf. montanum)°.

M. montanum Benth. var. nov. n= 11. GUATEMALA. Huehuetenango: be-
tween Chemal & San Juan Ixcoy, Beaman 3043 (reported as M. montanum)°.
MEXICO. Chiapas: 17 mi W of San Cristobal de Las Casas, King 2796 (reported
as M. montanum)°; 5 mi E of San Cristobal de Las Casas, King 2801 (reported as
M. montanum)*; Tecpisca, King 2843 (reported as M. montanum)®*; 34 mi S of
Ishuatan, Stuessy 559, 560; 20 mi W of San Cristobal de Las Casas, Stuessy 566.

M. paniculatum Gardn. n = 18. GUATEMALA. Alta Verapaz: near San Pedro
Carcha, King 3329 (reported as M. mimulifolium)*; San Pedro Carcha, Stuessy
594; Huehuetenango: near Huehuetenango, King 3417 (reported as M. dicoelo-
carpum)*; Huehuetenango, Stwessy 578; 12 mi E of Huehuetenango, Stuessy 582;
Solola: near Panajachel, King 3242 (reported as M. cf. mimulifolium)®.

M. perfoliatum (Cav.) H. B. K. n = 10. Plants obtained frem bot. gard.
Goteborg®.

M. perfoliatum (Cav.) H. B. K.n = 11. COSTA RICA. Cartago: Turrialba,
King 5350'; near Cartago, King 5407'. GUATEMALA. Guatemala: Guatemala, King
3248 (reported as M. cf. perfoliatum)*®; Huehuetenango: Huehuetenango, King 3410
(reported as M. ct. perfoliatum)*; Huehuetenango, Stuessy 576. MEXICO. Mi-
choacan: 45 mi W of Morelia, King 3635 (reported as M. cf. perfoliatum)®; 8 km
S of Uruapan, King & Soderstrom 4707' ; Ciudad Hidalgo, Powell & Edmondson 816

32 MADRONO [ Vol. 20

(reported as M. cf. perfoliatum)*; Zitacuaro, Stuessy 379 (fig. 9), 380; Oaxaca:
Zimatlan, Stuessy 654.

M. perfoliatum (Cav.) H. B. K.n = 12. MEXICO. Michoacan: 11 mi W of
Michoacan-Mexico state border, rte 15, King 3600°; Puebla: near Puebla, King
35608,

M. rosei Rob. n = 10. MEXICO. Simaloa: Mazatlan, Flyr 138%; 13 mi N of
Rosario, King 3710°; 21 mi N of Rosario, King 3712°; near Mazatlan, King
3715°; 10 mi NE of jct rtes 40 & 15, King 3716"; Isla Piedra, Stuwessy 747, 748; near
Mazatlan, Stuessy 749, 750.

M. sericeum Lag. n = 30. MEXICO. Guerrero: Petaquillas, Stwessy 364 (fig.
3); Michoacan: Zitacuaro, Stuessy 377 (ca 30); 7 mi S of Ario de Resales,
Stuessy 690 (& frag.) 691; Oaxaca: 53 mi S of Tehuacan, Powell 660 (reported
as M. sericeum var. sericeum)*; Las Sedas, Stuessy 660 (ca 30); Querétaro: 6 mi
W of Querétaro, Powell & Edmondson 579 (ca 30, reported as M. sericeum var.
exappendiculatum)®; near Querétaro, Rock M-442 (reported as M. sericeum var.
exappendiculatum)®.

M. tenellum Hook. & Arn. n= 10, MEXICO. Nayarit: 38 mi S of Sinaloa-
Nayarit border, King 3703 (reported as M. cupulatum)*; 28 mi S of Sinaloa-
Nayarit border, King 3704 (reported as M. cupulatum)®; ca 21 mi S of Sinaloa-
Nayarit border, King 3705"; Sinaloa-Nayarit border, King 3706 (reported as M.
cupulatum)®; 27.9 mi SE of Nayarit-Sinaloa border, Stuessy 401; 10 mi NW of
jet rte 15 & rd to Tuxpan, Stuessy 744 (ca 10), 745.

Faculty of Organismic and Developmental Biclogy and The Herbarium,
The Ohio State University, Columbus.

LITERATURE CITED

Jackson, R. C. 1957. Documented chromosome numbers of plants. Madrono
14:111.

Necopi, G. 1938. Contributo alla cariclogia ed alla morfologia di alcuni Melam-
pedium (Ccmpositae, Tubuliflcrae-Heliantheae-Melampodinae). Ann. Bot.
(Rome) 21:495-502.

Powe Ll, A. M., and S. Sikes. 1970. Chromosome numbers of some Chihuahuan
Desert Compositae. Southw. Naturalist 15:175-186.

Rosinson, B. L. 1901. Synopsis of the genus Melampodium. Proc. Amer. Acad.
Arts 36:455—466.

SPEESE, B. M., and J. T. Batpwin, Jr. 1952. Chromosomes of Hymenoxys. Amer.
J. Bot. 39:685-688.

StuEssy, T. F. 1968. A systematic study of the genus Melampodium (Compositae-
Heliantheae). Ph.D. dissertation. Univ. of Texas at Austin.

TurNER, B. L., J. H. BEAMAN, and H. F. Rocx. 1961. Chromosome numbers in the
Compositae. V. Mexican and Guatemalan species. Rhodora 63:121-129.

————., and D. F ryr. 1966. Chromosome numbers in the Compositae. X. North
American species. Amer. J. Bot. 53:24-33.

. and M. C. Jonnston. 1961. Chromosome numbers in the Composite-III.
Certain Mexican species. Brittonia 13:64—-69.

. and R. M. Kinc. 1962. A cytotaxonomic survey of Melampcdium (Com-
positae-Heliantheae). Amer. J. Bot. 49:263-269.

oa . and — . 1964. Chromosome numbers in the Compositae. VIII.
Mexican and Central American species. Southw. Naturalist 9:27-39.

Watters, J. L. 1952. Heteromorphic chromosome pairs in Paeonia californica.
Amer. J. Bot. 39:145-151.

HAROLD ERNEST PARKS
LEE BONAR

Harold Ernest Parks was born at Albany, Oregon, August 5, 1880.
Some of his early years were spent in Tacoma, Washington. Following
the death of his father the family moved to California in 1890, living
in various localities in the San Francisco Bay area while young Harold
attended public school.

He became a member of the California National Guard in San Rafael,
and his unit was mustered into federal service as part of “K’’? Company,
First California U.S. Volunteer Division, April 27, 1898, and sailed on
the transport, City of Pekin, May 28, 1898. ‘““K”’ Company participated
in the capture of Guam, June 1898, and arrived in the Philippines,
June 30, 1898, taking part in the battles around Manila and the capture
of the city, August 13, 1898. The company continued service in that
area during the Philippine Insurrection and Parks was wounded in ac-
tion, February 28, 1899. He served later that year on Negros Island and
was discharged from the Army, September 21, 1899, at San Francisco.

He moved to Tacoma, and worked as a salesman for H. J. Henry and
the National Biscuit Company from 1900 through 1910.

While in Tacoma he married Bessie A. Reynolds. Three sons were
born to this marriage: Robert Wayne, Wendell King, and Laurance
Dale. The family moved to Santa Cruz, California, where Parks worked
as mill worker and salesman until 1914, when he went to San Jose, Cali-
fornia, as a special clerk in the Post Office. Mr. and Mrs. Parks were
divorced in 1924.

During these years in California Parks took a number of courses in
language and business training from the International Correspondence
School of Scranton, Pennsylvania. Also during this period he developed
an interest in the study of plants, especially certain groups of fleshy
fungi. He wrote letters of inquiry seeking advice from professional men
and finally settled on a program of research in fungi with W. A. Setchell
as advisor and authority to whom he submitted collections for identifica-
tion. Parks soon developed an especial interest in the collection and
study of the hypogeous fungi of his area.

During the period 1916 to 1921 he collected, traveling by bicycle, in
the hill areas west of San Jose. All his free time was devoted to collect-
ing, compiling notes, and sending out collections. A file shows 30 letters
written him by Setchell during the calendar year 1918 giving answers to
questions and identifications for specimens. Some letters gave up to
25 identifications.

With the encouragement of W. A. Murrill and Setchell, Parks wrote
two papers on fungi during this time: Notes on California Fungi (My-
cologia 9:10-21. 1919), and California Hypogeous Fungi—Tuberaceae
(Mycologia 13:301-314. 1921).

373

374 MADRONO [Vol. 20

He also wrote, with suggestions and advice from certain zoologists, a
paper relating to the habits of the wood rats and their use of hypogeous
fungi as food: The genus Neotoma in the Santa Cruz Mountains (J.
Mammology 3:241-253. 1922).

Setchell once said that this man Parks bombarded him with so much
work that he decided that he had better try to get him to come to Berke-
ley and work for the Botany Department. On August 23, 1921, Setchell
offered Parks a position as helper and general handy man at a salary of
$150 per month; Parks served as technical assistant from September 1,
1921 through June 30, 1922.

He was appointed Collector for the Department of Botany, Univer-
sity of California, July 1, 1922, and continued in this position until he
resigned June 30, 1928. He was Associate Curator of the University
of California Herbarium, without salary, July 1, 1928, to June 30, 1950,
and held the title of Honorary Curator until his death.

His extensive general knowledge of field botany made Parks a very
valuable employee of the department. His main duties were collecting
class and research material for staff and graduate students. He also had
charge of the departmental storeroom and was general handy man for
the department. During this time he combined his interest in fungi and
obtaining specimens of them with collecting class material.

Parks became known as an outstanding student and collector of hy-
pogeous Gasteromycetes and contributed very extensive amounts of
material to S. M. Zeller and C. W. Dodge for some twenty years, start-
ing in 1918. His collections are repeatedly cited in their monographic
studies published in the Annals of the Missouri Botanical Garden.

Parks had extensive experience in collecting during several summer
trips to certain Pacific Islands. He went as assistant to W. A. Setchell
on the University of California-Carnegie Institution Expedition to Ta-
hiti, May 16-July 19, 1922. With W. A. Setchell, C. B. Setchell, J. E.
Hofmeister, and J. M. Ostergard on an Expedition to the Tonga Islands
under the auspices of the Bernice P. Bishop Museum, May 31-August
23, 1926. Parks spent most of his time in a botanical survey of Eua
Island. During the summer of 1927 he made a collecting trip to Fiji as
Research Associate of the Bishop Museum.

He was married to Susan Priscilla Thew in October 1927. They made
a trip, on their own, May to July 1930, to Raratonga and the Cook
Islands.

Parks made general collections on these expeditions but gave special
care to the collection of fungi and distribution of specimens following
the return of the expedition. He sent material to various specialists over
the world and compiled publications on some of it. James R. Weir, C. L.
Shear, and other members of the staff in charge of the Pathological Col-
lections of the United States Department of Agriculture were most
helpful with this work. Duplicates deposited in the University of Cali-

H. i. PARKS

BONAR

1970]

376 MADRONO [Vol. 20

fornia Herbarium have greatly enriched our collections. Published rec-
ords on some of these collections are as follows: Lichenes a W. A. Set-
chell et H. E. Parks in insula Tahiti a 1922 collecti, by E. A. Vainio
(Univ. Calif. Publ. Bot. 12:1-16. 1922); Report on a collection of ferns
from Tahiti, by W. R. Maxon (Univ. Calif. Publ. Bot. 12:17-44. 1924);
Tahitian mosses collected by W. A. Setchell and H. E. Parks, determined
by V. F. Brotherus (Univ. Calif. Publ. Bot. 12:45-48. 1924); Tahitian
fungi collected by W. A. Setchell and H. E. Parks, by H. E. Parks
(Univ. Calif. Publ. Bot. 12:49-59. 1926); Tahitian algae collected by
W. A. Setchell, C. B. Setchell, and H. E. Parks, by W. A. Setchell (Univ.
Calif. Publ. Bot. 12:61-142. 1926); Tahitian spermatophytes collected
by W. A. Setchell, C. B. Setchell, and H. E. Parks, by W. A. Setchell
(Univ. Calif. Publ. Bot. 12:143-230, 1926); The Tonga expedition of
1926, by W. A. Setchell (Science 64:440-442. 1926); Ferns of Fiji, by
E. B. Copeland (Bernice P. Bishop Mus. Bull. 59:1-106. 1929); Rara-
tonga ferns, collected by Harold E. and Susan Thew Parks, and miscel-
laneous oriental pteridophytes, by E. B. Copeland (Univ. Calif. Publ.
Bot. 12:395-381. 1931); and New Plants from Fiji—I, by J. W. Gilles-
pie (Bernice P. Bishop Mus. Bull. 74:1-99. 1930).

During 1931 the Parkses became established in a home overlooking
the Pacific at Trinidad, Humboldt Co., California. This was an area little
known mycologically, and Parks continued his botanical work in this
area for twenty-two years. He formed a close working relationship with
another enthusiastic botanical student, Joseph P. Tracy (UC ’03) of
Eureka, California, and the two very often collected together to advan-
tage, since Tracy was well-known for his studies on the higher plants of
that region. Parks collected fungi generally, but his emphasis was on
parasitic fungi and all sorts of micro-fungi. His series of collection num-
bers of California fungi reached approximately 9000; these were almost
all sent to the University of California Herbarium. This material came
with a goodly portion of it already identified, and was collected, when
possible, in sufficient quantity to make distribution sets of 25-30 dupli-
cates. The fact that Parks collected 696 of the first 1225 sets of Fungi
of California distributed by the University of California Herbarium
exemplifies the extent of his contributions.

Parks frequently received requests from specialists for collections of
particular fungi in which they were interested and had correspondence
with many mycologists. He collected Discomycetes for Edith K. Cash
and Thelephoraceae for H. S. Jackson.

The following thirty of Parks’ collections, including one new genus,
were named in his honor; these represent widely different types of
plants from different parts of the Pacific Basin: Parksia libocedri Cash,
California; Asplenium parksu Copel., Raratonga; Belonidium parksii
Cash, California; Cyathea parksiae Copel., Raratonga; Cyrtandra park-
sit Setch., Tahiti; Dennstedtia parksti Copel., Tongatabu; Ectocarpus

1970] BONAR: H. E. PARKS ST

parksii Setch. & Gardn., California; Eryrotrichia parksu Setch. & Gardn.,
California; Freycinetia parksiu Mart., Fiji; Fucus parksiu Setch. &
Gardn., California; Gautieria parksiana Zell. & Dodge, California; Hyd-
nangium parksu Zell. & Dodge, California; Hymenogaster parksu Zell.
& Dodge, California; Hypoxylon parksu Lloyd, Tahiti; /ridophycus
parksii Setch. & Gardn., California; Languas parksi Gill., Fiji; Loxo-
gramme parksu Copel., Fiji; Lycopodium parksu Copel., Fiji; Maesa
parksii Gill., Fiji; Melanogaster parksu Zell. & Dodge, California;
Oleandra parksii Copel., Fiji; Pandanus odoratissimus L. var. parksit
Mart., Tonga; Peridermium parksianum Faull, California; Polypodium
parksiu Copel., Fiji; Portia parksit Murr., California; Puccinia farksiana
Cumm., Fiji; Salix parksii Ball, California; Spireanthemum parksu Gill.,
Fiji; Strigula (Melanothele) parksii Ras. ex Sbarb., Cook Islands; and
Aglaia parksu A. C. Smith, Fiji.

The Parkses became well-known and active members of the com-
munity in the Trinidad and Eureka area and many visitors enjoyed the
hospitality of their pleasant home. With advancing age and increasing
health problems they disposed of their home in 1953 and spent the re-
mainder of their time in traveling, living in guest hotels and health re-
sorts. Parks died after protracted illness at Calistoga, California, March
5, 1967. His wife, Susan Thew Parks, followed him in death on January
29, 1968. They are buried at Visalia, California. The University of Cali-
fornia Herbarium owes much to the numerous contributions of Harold
Ernest Parks.

Department of Botany, University of California, Berkeley

NEW RECORDS OF MYXOMYCETES FROM CALIFORNIA IV.
DONALD T. KOWALSKI AND DWAYNE H. CurTIS

The new records of slime molds listed in our last paper (Kowalski
and Curtis, 1968) brought the total number of Myxomycetes recorded
in print for California to 190. Since then, seven new species and two new
records have been reported from the state (Kowalski, 1968a; 1968b;
1969a; 1969b; 1970). Ten new records are reported in this paper. This
brings the total number of slime molds found in California to 209 spe-
cies. All collections cited here have been deposited in the Herbarium of
the University of California at Berkeley. Unless otherwise stated, the
numbers are those of the authors, labeled K and C respectively. With
the exception of Diderma umbilicatum Pers., the names of the organ-
isms are those accepted by Martin (1949). This investigation was sup-
ported by National Science Foundation Grant GB-5799.

LICEACEAE

Licea parasitica (Zukal) Martin. Four collections, three on oak bark
from Lower Bidwell Park, Chico, Butte Co., K 2342, Jan. 8, 1966,

378 MADRONO [Vol. 20

K 9648, Dec. 22, 1966, K 9739, March 13, 1967, and one on decayed
wood, Sutter Buttes, 700 ft. elev., Sutter Co., K 5739, Feb. 18, 1967.
All four collections were made accidentally, i.e., they were not found in
the field. In each case, the substrate upon which L. parasitica was grow-
ing was originally collected because it had another, larger myxomycetous
species upon it. The minute sporangia of L. parasitica were discovered
later in the laboratory while the substrate was being scanned with a
stereoscopic microscope. There are only two species in the genus Licea
that are sessile and dehisce in a circumscissile manner by a preformed
lid, namely, L. parasitica and L. kleistobolus Martin. The sporangia of
both species are similar in shape and size, being subglobose to discoid
and 0.05—0.2 mm in diameter. They differ, however, in color and texture.
The sporangia of L. parasitica are dark brown or black and have a thick
horny wall at maturity, while those of L. kleistobolus are bright cop-
pery-brown and have a membranous wall. Licea parasitica has been
found as far west as Iowa and Texas and is considered rare. This, how-
ever, may be due to its small size and it probably occurs throughout the
United States.

Tubifera ferruginosa (Batsch) Gmel. Two collections, both on de-
cayed wood, MacKerricher Beach State Park, Mendocino Co., C 428,
Jan. 26, 1967 and C 1138, Jan. 25, 1968. Although the pseudoaethalia
in these collections are smaller than normal, measuring less than 1.0 cm
in diameter, their identity is easily established. There are four known
species in the genus and T. ferruginosa is separated from the other three
by the lack of a pseudocapillitium, the sessile fruiting bodies and by
having spores 6—8 » in diameter. This is a common taxon, being known
throughout the United States.

CRIBRARIACEAE

Cribraria microcarpa (Schrad.) Pers. Two collections, one developing
in a damp chamber from wood collected in Muir Woods, Marin Co., by
Victor Duran on Feb. 8, 1965, and one on decayed wood, 3 miles west
of Paul Dimmick State Park, Mendocino Co., K 8164, April 10, 1968.
The lack of a distinct basal cup, stalks more than six times the diameter
of the sporangia, and the small sporangial size, less than 0.3 mm in
diameter, separate this taxon from other members of the genus. The
sporangia in these collections are, in fact, much smaller. The majority
are about 0.1 mm in diameter. This is a common species, known from
many collections throughout the United States. It has probably been
overlooked in California until now because of its minute size.

Cribraria minutissima Schw. On decayed wood, Juniper Lake, Lassen
Volcanic National Park, 6,700 ft. elev., Lassen Co., K 3874, July 22,
1966. This is another tiny species, the sporangia being 0.1-0.2 mm in
diameter. It can be distinguished from C. microcarpa by the presence
of a distinct basal cup which is often constricted at the apex and by

1970] KOWALSKI & CURTIS: MYXOMYCETES 379

the shorter stalks. As mentioned above, in C. microcarpa the stalks are
usually more than six times the diameter of the sporangia while in
C. minutissima they are usually less than four times the diameter of
the sporangia. This species is also widely distributed in the United
States, but because of its small size, is not often collected.

ECHINOSTELIACEAE

Echinostelium minutum De Bary. Three collections, two on decayed
wood, K 5949, Covered Bridge, Honeyrun Road, Butte Co., April 15,
1967, K 7562, 5 miles east of Mineral, 5,800 ft. elev., Tehama Co., May
15, 1966, and one on bark, Lower Bidwell Park, Chico, Butte Co.,
K 9732, Dec. 27, 1966. As was the case with the Licea parasitica col-
lections mentioned above, these specimens were also found by accident
in the laboratory while viewing the substrate under a stereoscopic micro-
scope. Because of its small size, the sporangia being about 50 » in diam-
eter, most of the known collections of this species were made from moist
chamber culture. Thus, these collections are valuable since they rep-
resent natural fruitings which are rarely made. Of the four known spe-
cies in the genus, only E. minutum and E. cribrarioides Alex. have a
capillitium. Echinostelium minutum can be separated from £. crib-
rarioides by the fact that it has a scanty capillitium with free, hooked
ends and spores 7-8 » in diameter, while FE. cribrarioides has a well-
developed capillitium which forms a complete net and spores 9-10 p
in diameter. Previously, the furthest known western locality of E.
minutum in the United States was Texas. However, this taxon can prob-
ably be found in every state of the union, since any diligent damp cham-
ber work will usually turn up this tiny species.

STEMONITACEAE

Stemonitis webbert Rex. Two collections, both on decayed wood,
K 1679, Patrick’s Point State Park, Humboldt Co., July 3, 1965, and
K 4381, Lower Bidwell Park, Chico, Butte Co., Dec. 22, 1966. This
taxon and S. splendens Rost. are very closely related. In general, the
sporangial and spore characteristics are identical. The two taxa are sep-
arated on capillitial differences. The capillitium of S. splendens is purp-
lish-brown and the meshes of the surface net are 20—50 w in diameter.
The capillitium of S. webberi has red metallic reflections and the meshes
of the surface net are mainly 30-100 uw in diameter. As can be seen, these
differences are slight and it is possible that both of these taxa simply
represent different forms of the same species. Stemonitis webberi is
known throughout the United States and its exclusion from the pub-
lished reports from California may be due to some workers just consid-
ering it asa form of S. splendens.

Comatricha lurida Lister. On decaying leaves, Lower Bidwell Park,
Chico, Butte Co., C 578, Feb. 16, 1967. The sporangia in this collec-
tion have a total height of 1.0-1.5 mm, a capillitium that arises mainly

380 MADRONO [Vol. 20

from the apex of the columella and coarsely warted spores 8-10 p in
diameter. These features all fit the published descriptions for this
taxon. There is one characteristic, however, which is atypical, and that
is, several of the sporangia have pieces of the peridium remaining at-
tached to the ends of the capillitium. In one sporangium the peridium
is almost completely persistent, being fugacious only at the apex. This
unusual feature is not found in any of the published descriptions for this
taxon. It is simply stated that the peridium is fugacious or evanescent.
Because this species can have a partially persistent peridium, but, more
importantly, since most of the capillitium arises from the apex of the
columella, perhaps this species and C. elegans (Racib.) Lister, which
is similar to C. lurida, should be transferred to Lamproderma. This is
the genus in the Stemonitaceae which contains species that have per-
sistent peridia and a capillitium radiating from the apex of the colum-
ella. We believe these species are more closely related to Lamproderma
arcyrionema Rost. and L. biasperosporum WKowalski than to any other
species of Comatricha. However, since we do not have adequate ma-
terial of either species, we do not plan to make the transfers at this
time. According to Martin (1949), C. lurida is a rare species. In the
United States it has only been reported from Iowa and New York.

PHYSARACEAE

Physarum leucopus Link. Five collections, all from Lower Bidwell
Park, Chico, Butte Co., C 562, Feb. 3, 1967, K 4145, Dec. 10, 1966, and
K 5227, Feb. 4, 1967 on decaying bark and C 128, Dec. 7, 1966, and
K 2145, Nov. 21, 1965 on decayed wood. The sporangia in these collec-
tions all have small patches of white lime on the peridium and stalks
that are white and distinctly calcareous. In general, these collections
fit the published descriptions perfectly. While throughout the United
States this is not a particularly common species, it is extremely abundant
in the Sacramento Valley. We have made numerous collections of this
taxon, but for the sake of brevity, we only list five here.

Physarum luteolum Peck. Two collections, both made on decaying
bark, 2 miles west of Child’s Meadows, 4,400 ft. elev., Tehama Co., April
16, 1966, K 2724 and 2748. The sporangia in these collections are ses-
sile, crowded, brilliant yellow in color and have a single-layered peridium.
This species is not common. Except for a listing by Hagelstein (1944)
from Colorado, this species is only known from lower elevations east of
the Mississippi River. It is thus surprising that the only known Califor-
nia collections are from the mountains. Our work has shown that, with
few exceptions, species of Myxomycetes found in the mountains are not
found in the lowland areas and vice-versa. Perhaps the montane Cali-
fornia collections represent a different ecotype from the eastern low-
land collections. On a morphological basis, however, there is no ques-
tion that they represent the same species.

1970] KOWALSKI & CURTIS: MYXOMYCETES 381

DIDYMIACEAE

Diderma umbilicatum Pers. Five collections, three on decayed wood,
Butte Creek and Skyway, Butte Co., K 4469 and 4492, Dec. 27, 1966,
K 5822, March 18, 1967, one on decaying Eucalyptus bark, Point Reyes
Ranger Station, Marin Co., K 5148, Jan. 29, 1967, and one on decayed
leaves, Sutter Buttes, 700 ft. elev., Sutter Co., A 7891, March 2, 1968.
This species is also very common in the lowland areas of California. We
have found it approximately 40 times, but only list five specimens here
to conserve space. The exact taxonomic standing of this taxon varies
with different authors. Both Lister (1925) and Hagelstein (1944) treat
it as a variety of Diderma radiatum (L.) Mrogan. Martin (1949) sim-
ply placed it in synonymy with D. radiatum. We believe, however, that
the differences which it exhibits from D. radiatum are distinct and con-
sistent enough to warrant its retention as a separate species. The major
differences between the two species are as follows: In D. radiatum the
sporangia are gray to, more commonly, brownish or red-brown, the
stalks are ochraceous to reddish brown and the sporangial wall is dis-
tinctly cartilaginous and dehisces in a stellate fashion. In D. umbili-
catum the sporangia are usually white but may be cream-colored, but
are never brown or red, the stalks are white or cream-colored and the
cartilaginous nature of the sporangial wall can only barely be observed
and it never dehisces in a stellate manner. In fact, except for a few
cracks in the wall at the apex of the sporangium, dehiscence must be
by external breakage. Diderma umbilicatum is apparently a rare taxon.

Although Lister (1925) states that it has been found in many of the
United States, we do not have knowledge of any collections other than
our own.

Department of Biology, Chico State College, Chico, California.

LITERATURE CITED

HAGELSTEIN, R. 1944. The Mycetozoa of North America. Mineola.
KowatskI, D. T. 1968a. Three new species of Diderma. Mycologia 60:595-603.
. 1968b. Observations on the genus Lamproderma. Mycologia 60:756—768.
. 1969a. A new coprophilous species of Calonema. (Myxomycetes). Ma-
drono 20:229-231.
—. 1969b. A new coprophilous species of Didymium. Mycologia 61:635-639.
. 1970. Concerning the validity of Lamproderma echinosporum. Madrono
20:323-326.
. and D. H. Curtis. 1968. New records of Myxomycetes from California
III. Madronio 19:246-249.
Lister, A. 1925. A monograph of the Mycetozoa. 3rd ed. by G. Lister, Brit. Mus.
Nat. Hist. London.
Martin, G. W. 1949. North American Flora 1:1—190.

JZ - MADRONO [Vol. 20

NOTES AND NEWS

FASCIATION OF COASTAL REDWoops.—Fasciation involves a flattening of the nor-
mally cylindrical stem. A fasciated stem is usually much heavier than a normal
shoot. The flattened growth is due to the formation of a row of linked meristems,
instead of a single one at the apex. It occurs both on conifers and on hardwoods
as well as on many other plants. Fasciation has not yet been reported on ‘coastal
redwood. Its cause is unknown, and according to my experience of seven years of
redwocd research I have seen only two cases of this curious phenomenon.

Fic. 1. Fasciation in Sequoia sempervirens.

Fasciation is sometimes only of annual duration, some of the terminal buds
resume normal shoot growth again the following growing season, but it may con-
tinue for longer periods. Fasciation can be due to wound stimulation, possibly as a
result of insect attack, or to overnutrition or a disbalance of growth hormones.
There is no evidence to support this in the two cases observed in fasciation of
coastal redwood. In some cases, it is considered to be genetically controlled by a
mutation, which can be propagated vegetatively and which may come true from
seed. However, the above two reported cases indicate that only one or two leaders
developed such fasciation out of a young growth redwood tree. Another possibility
may be to rank it as a pathological curiosity, probably caused by virus infection.
It is very rare which is the reason to report it here. The photographs illustrate
(fig. 1) this phenomenon on a young growth redwood tree collected January 5,
1966, by Robert J. Wright, Utility Tree Service, Inc., Eureka, California on the

1970] REVIEWS 383

Fickle Hill Road above Arcata, California. The specimen has not been preserved
to my knowledge. It is known that the Utility Tree Service, Inc. crews have used
herbicides in their program of brush control on power line right-of-ways.

The author is interested in receiving information about any extraordinary or
abnormal growth features on coastal redwoods. Any such information should in-
clude details about the observed abnormality and should possibly be accompanied
with the abnormal specimen itself or a photograph—Rupo.tF W. BeEcxk1nc, School
of Natural Resources, Humboldt State College, Arcata, California.

Back Issues oF MApRoNo.—Back issues of most numbers of Madrono are still
available. Some numbers are in short supply. Any surplus copies of any issue of
Madrono will be gratefully received by the Corresponding Secretary, Department
of Botany, University of California, Berkeley, California 94720.

REVIEWS

Marin Flora. Manual of the Flowering Plants and Ferns of Marin County, Cali-
fornia. Second edition with supplement. By Joon THomMaAs Howe tv. University of
California Press, Berkeley and Los Angeles, California, vii + 366 pp. 1970. $10.00.

Since 1949, when the first edition of this book appeared, the population of the
ten counties making up the immediate San Francisco Bay Area has approximately
tripled to its present level of about five million people. Within the next decade, if
present trends continue, there will be about as many people in this single metropoli-
tan area as there were in the entire world at the time of Christ. In view of this, it is
extremely fortunate that more than a fifth of the 529 square miles of the lovely
Marin County peninsula have been set aside for public enjoyment, the largest seg-
ment being the Point Reyes National Seashore of about 53,000 acres which was
authorized in 1962. It is likewise fortunate that the University of California Press
has added to their extensive publication list of local natural history guides a new
version of this delightfully written and scientifically critical flora.

For the most part, the 1970 printing exactly duplicates that of 1949, but with
several improvements. The quality of the paper and the binding have been im-
proved greatly and the size of the pages increased slightly, making the type much
easier to read. In addition, a 43-page supplement has been added, demonstrating
that four genera and 26 species have been added to the indigenous flora of the
County, together with 37 genera and 99 species of adventive plants. There are at
present 1023 indigenous species and 408 introduced species recorded from the region.
Among the attractive features of the original work that have been retained are the
25 black-and-white photographs of Charles H. Townsend, perhaps a third of which
(those not on public lands) could not be duplicated today. The maps of localities in
Marin County and of trails and localities on Mt. Tamalpais, being larger, are clearer
than in the original. The price continues to be reasonable, and there is every reason
to expect that Marin Flora will continue to enjoy as much popularity in the years
to come as it has in the past—PrETER H. Raven, Department of Biological Sciences,
Stanford University.

384 MADRONO [Vol. 20

Nightshades, The Paradoxical Plants, by CHar.tes B. HEIsER, JRr., 200 pp. W. H.
Freeman and Company, San Francisco. 1969. $5.95.

Charles Heiser has written an entertaining and informative book about those
members of the Nightshade Family (Solonaceae) that have been important in
human history. The orientation of the book is neither too technical for the lay-
man nor too popular for the professional biologist.

The prologue, written in Heiser’s very personal style, sets the tone for the entire
volume. In it he briefly sketches the pertinent information about the taxonomy,
morphology, and cytology of the Solonaceae. Although the organization of a few
of the chapters suggests that there was some difficulty in integrating material from
diverse sources, the accounts of the various species make for enjoyable reading.
One chapter (“Some Like It Hot’’) discusses the uses and economic history of the
chili pepper, Capsicum, while another chapter (“Love Apples’) chronicles the to-
mato in European culture. Among the other nightshades discussed in the book are
those of medicinal, horticultural, magical and narcotic value. The most interesting
chapter, however, describes the controversy between Luther Burbank and The
Rural New Yorker concerning Burbank’s “Wonderberry.” The Rural New Yorker
claimed Burbank’s creation was a fraud. They believed “Wonderberry” to be noth-
ing more than the common nightshade Solanum nigrum, rather than a hybrid be-
tween S. guineense and S. villosum. Heiser’s own investigation into the matter pro-
vides a satisfying conclusion to the chapter.

One would hope that more books in this vain and of this quality will soon ap-
pear. Nightshades, The Paradoxical Plants, will make an important addition to any
botanical library—DeENNIs R. PARNELL, Department of Biological Science, Califor-
nia State College, Hayward.

Flowers of the Point Reyes National Seashore. By ROXANA S. FERRIS. xi +
119 pp., illustrated. University of California Press, Berkeley. 1970. $2.65, paper,
$7.95 cloth.

The majority of books about the plants of different areas fall into one of two
categories: 1, they are intensely technical and hence difficult for the interested
amateur, or 2, they are too sketchy, too abbreviated and hence satisfy no one,
neither the amateur nor the professional. This very nice book about the plants of
the Point Reyes National Seashore in Marin County, California, strikes a good
balance between the two extremes. The text is accurate, interesting to read, and is
accompanied by nearly 200 line drawings by Jeanne R. Janish. Keys are not used,
but the plants are grouped by flower color. The introductory material contains a
map of the Point Reyes area, some 53,000 acres, and a description of the plant
associations found within the Seashore. An index of both botanical and popular
names is included as well as a bibliography.

We should have many more books like this for different areas so that the public
can gain a better understanding of the world around us.—JoHn H. Tuomas, Depart-
ment of Biological Sciences, Stanford University.

A WEST AMERICAN JOURNAL OF BOTANY

The subscription price of Madrono is $8.00 per year ($4.00 for stu-
dents and $12.00 for institutions). If your institution does not now sub-
scribe to Madrofio, we would be grateful if you would make the neces-
sary request.

INFORMATION FOR CONTRIBUTORS

Shorter items, such as range extensions and other biological notes,
will be published in condensed form with a suitable title under the general
heading, “‘Notes and News.”

Abbreviations of botanical journals should follow those in Botanico-
Periodicum-Huntianum (Hunt Botanical Library, Carnegie-Mellon Uni-
versity, Pittsburgh, Pennsylvania, 1968).

Footnotes should be avoided whenever possible.

Membership in the California Botanical Society is normally considered
a requisite for publication in MADRONO.

STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION
(Act of Oct. 23, 1962; Section 4369, Title 39, United States Code)

Maprono, A West American Journal of Botany, is published quarterly at Berke-
ley, California.

The publisher is the California Botanical Society, Inc., Life Sciences Building, Uni-
versity of California, Berkeley, California 94720.

The editor is John H. Thomas, Division of Systematic Biology, Stanford Univer-
sity, Stanford, California 94305.

The owner is the California Botanical Society, Inc., Life Sciences Building, Uni-
versity of California, Berkeley, California 94720. There are no bond holders, mort-
gagees, or other security holders.

The average number of copies distributed of each issue during the preceding 12
months is 650; the number of copies of the single issue closest to the filing date is
650.

I certify that the statements made by me above are correct and complete.

Joun H. THomas, Editor
Nov. 1, 1970.

QK
\
Migs
Pa oT.

VOLUME 20, NUMBER 8 OCTOBER 1970

Contents

An EcoLocicaL CONTRIBUTION TO THE TAXONOMY

oF ArTEMIsIA, A. A. Beetle 385
C. Leo HitcHcock 387
IRIS PSEUDACORUS IN WESTERN NortH AMERICA,

Peter H. Raven and John H. Thomas 390
A New SPECIES OF PROBOSCIDEA (MARTYNIACEAE) FROM

BAJA CALIFORNIA, MEXICO, Richard H. Hevly 392

A New AstTRAGALUS (FABACEAE) FROM Nevapa, R. C. Barneby 395

Notes on LoEFLinctaA (CARYOPHYLLACEAE), R. C. Barneby
and Ernest C. Twisselmann 398

Unvusvuat Factors CoNTRIBUTING TO THE DESTRUCTION OF
Younc Giant Sequoras, Howard S. Shellhammer,
Ronald E. Strecker, H. Thomas Harvey, and

Richard J. Hartesveldt 408
Lrvear MIcrosPoRE TETRADS IN THE GRASS STIPA ICHU,

Frank W. Gould 411
EXTENSION OF THE RANGE OF ABIES LASIOCARPA INTO CALIFORNIA,

J.O. Sawyer, D. A. Thornburgh, and F. F. Bowman 413

Reviews: Tyler Whittle, The Plant Hunters (John H. Thomas) ;
D. Briggs and S. M. Walters, Plant Variation and Evolution
(Dennis R. Parnell) 415

Notes AND News: MaproNo 391

RIBES MALVACEUM IN THE FOOTHILLS OF CALAVERAS COUNTY,
CatrrorniA, Dean William Taylor 416

A WEST AMERICAN JOURNAL OF BOTANY > SSP

BLISHED QUARTERLY BY THE CALIFORNIA eect RONEN
| OM tnd Ni

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MADRONO
A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established
1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional
subscription price $12.00 per year. Published quarterly in January, April, July, and
October by the California Botanical Society, Inc., and issued from the office of
Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley,
California. Orders for subscriptions, changes in address, and undelivered copies
should be sent to the Corresponding Secretary, California Botanical Society, Depart-
ment of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS
CLASS OF:

1970—Lyman Benson, Pomona College, Claremont, California
Mrirprep E. Maruias, University of California, Los Angeles

1971—Marion OwnBeEy, Washington State University, Pullman
Joun F. Davinson, University of Nebraska, Lincoln

1972—Ira L. Wiccins, Stanford University, Stanford, California
REED C. Rotiins, Harvard University, Cambridge, Massachusetts

1973—WattacE R. Ernst, Smithsonian Institution, Washington, D.C.
Roy L. Taytor, University of British Columbia, Vancouver

1974—KenTon L. CHAMBERS, Oregon State University, Corvallis
EMLEN T. LITTEL, Simon Frazer University, Burnaby, British Columbia

1975—ArTURO GoMEz Pompa, Universidad Nacional Autonoma de México
Duncan M. Porter, Missouri Botanical Garden, St. Louis

Editor — Joun H. THomas
Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JUNE McCasKILL
P.O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

The Council of the California Botanical Society consists of the officers listed
above plus the immediate past President, Harry Thiers, Departments of Ecology and
Systematic Biology, San Francisco State College; the Editor of Madrofio; and
three elected Council Members: Robert Ornduff, Department of Botany, University
of California, Berkeley; Malcolm Nobs, Carnegie Institution of Washington, Stan-
ford; and Elizabeth McClintock, Department of Botany, California Academy of
Sciences, San Francisco.

AN ECOLOGICAL CONTRIBUTION TO THE
TAXONOMY OF ARTEMISIA

A. A. BEETLE

Beetle (1960) recognized three species of Artemisia (A. arbuscula,
A. nova, and A. longiloba) as distinct because they were “‘distinguish-
able on the basis of morphology, distribution, and ecology in addition
to being reasonably disjunct.”’ Holmgren and Reveal (1966) in a check-
list recognized only one species, Artemisia arbuscula. This they divided
into ssp. arbuscula (with A. longiloba as a synonym) and ssp. nova
implying that these taxa are not as distinct as maintained by Beetle.

In recent years the woody species of Artemisia have been studied in-
tensively both in the field and in the herbarium. Most of the field work
has resulted from the emergence of a new field of science, range man-
agement. Woody species of sagebrush are unproductive in terms of do-
mestic animal carrying capacity, and because various taxa respond
differently to chemical spraying, to protection, to burning, to various
degrees of ungulate grazing and to mechanical treatments they have
received much attention.

Range managers have been able to describe differences in phenology,
and in edaphic distributions. This remarkably productive area of field
observation, carried out in at least five different states and by inde-
pendent workers representing different institutions and using varying
research techniques, has resulted in a western concensus about the num-
ber of species and the degree of subspecific variation.

Passey and Hugie (1962) recognized A. arbuscula (low sagebrush),
A. nova (black sagebrush) and A. Jongiloba (alkali sagebrush) as spe-
cies. They described the different soils on which each occurs ‘‘on the
foothills and plains of the Great Basin.”

Robertson, et al. (1966) found that “in North Park, Colorado, the
alkali sagebrush (A. longiloba) plant community stands out in sharp
contrast from adjacent sagebrush range.” In a similar study of sites in
Wyoming, Thatcher (1959) found A. nova equally distinct on its own
site. In Nevada Zamora (1968) found A. arbuscula, A. longiloba and
A. nova distinct.

In Oregon, Gates (1964) recognized leaf defoliating moths as occur-
ring on both A. nova and A. arbuscula.

Young, et al. (1963) studied chemically the three species in question
as they occurred in Wyoming and recognized all three as distinct. A
similar study in Nevada (Holbo and Mozingo, 1965) achieved similar
results. More recently, in Idaho, Winward and Tisdale (1969) have
agreed with both Young and Holbo.

Maprono, Vol. 20, No. 8, pp. 385-416, August 6, 1971

385

386 MADRONO [Vol. 20

There may be an explanation for the fact that floral lists usually dis-
agree with the conclusions of field workers. Often field identification of
sagebrush is definitive, but the same plant on a pressed specimen in
the herbarium may be quite confusing. Field students are largely con-
cerned with mapable units of vegetation and study pure stands. Collec-
tors who contribute to herbaria are more likely to be concerned with
variation. Variation is easy to find since all species hybridize when given
the opportunity. These hybrids have a longer survival value in the
herbarium than they do in the field. Herbarium material of the woody
species of Artemisia does not reflect accurately the field situation. In
the herbarium the percentage of sheets representing hybrid variants is
much higher and relatively more significance is placed on them because
of the taxonomic difficulty of pidgonholing such specimens.

While most of the species in this section of Artemisia are old, con-
servative, and derived from diploid populations (e.g., A. mova and A.
longiloba), some of the entities (e.g., 4. arbuscula) are synthetic. Un-
less these differences are understood the taxonomic treatment may fail
to reflect the true situation.

Eventually the two groups (field ecologists and herbarium taxono-
mists) will find a common meeting ground but for the present, it may be
expected that a difference in taxonomic treatment of the same group of
plants will continue.

Range Management, University of Wyoming

LITERATURE CITED

BEETLE, A. A. 1960. A study of sagebrush. Wyoming Agric. Expt. Sta. Bull.
368:1-83.

Gates, D. H. 1964. Sagebrush infested by leaf defoliation moth. J. Range Managem.
17:209-210.

Horso, H. R., and H. N. Moztnco. 1965. The chromotographic characterization
Artmisia, Section Tridentatae. Amer. J. Bot. 52:970-978.

Hormcren, A. H., and J. L. REvEaAL. 1966. Checklist of the vascular plants of the
Intermountain Region. Intermountain Forest Range Exp. Sta. Res. Pap. INT-32.

Passey, H. B., and V. K. Hucuie. 1962. Sagebrush on relic ranges in the Snake
River Plains and northern Great Basin. J. Range Managem. 15:274-278.

Rosertson, D. R., J. L. NIErsen, and N. H. Bare. 1966.. Vegetation and soils of
alkali sagebrush and adacent big sagebrush ranges in North Park, Colorado.
J. Range Managem. 19:17-20.

TuHatTcHer, A. P. 1959. Distribution of sagebrush as related to site differences in
Albany County, Wyoming. J. Range Managem. 12:55-61.

Winwarp, A. H., and E. W. TispaAte. 1969. A simplified chemical method for sage-
brush identification. Univ. Idaho Forest, Wildlife, Range Expt. Sta. Note 11:1-2.

Younc, A. L., et al. 1963. A taxonomic system by chemical differentiation of the
genus Artemisia. Wyoming Range Managem. 172:1-3.

ZAMORA, B. 1968. Artemesia arbuscula, A. longiloba, A. nova plant associations in
central and northern Nevada. M.S. thesis, Univ. Nevada, Reno.

C. LEO HITCHCOCK

A TRIBUTE FROM THE CALIFORNIA BOTANICAL SOCIETY

To friends, colleagues and students he is known simply as “Hitchy.”
This informal nickname epitomizes Dr. C. Leo Hitchcock the man, and
conjures up warm memories for the many who have known him over the
years. He who tries to classify Hitchy encounters a tough taxonomic
problem. The attributes of the taxon run the gamut from thorough
monographer of several genera of flowering plants, writer of floristics,
and peerless teacher, to long-distance backpacker, keen horticulturalist,
avid ‘birder,’ terrific bridge-player, and formidable touch-footballer.

C. Leo Hitchcock was born (April 23, 1902) and grew up in Cali-
fornia and so came to know that flora intimately under the tutelage and
companionship of such eminent California botanists as Philip Munz,
Edmund Jaeger, and Marcus E. Jones, and as well as through his fre-
quent field trips with fellow students David Keck, George Goodman and
others. Hitchy’s undergraduate training was at Pomona College where
he stayed on to do a Master’s thesis with Dr. Munz on Clarkia (then,
Godetia). He allows that he was ultimately scooped years later by Har-
lan Lewis.

During the time Hitchy was in college he worked in the nearby oil
fields. We suspect that his earthy camaraderie may have been nurtured
by association there with fellow crewmen. From Pomona, Hitchy went
on to Washington University and the Missouri Botanical Garden, to
study with J. M. Greenman; Edgar Anderson’s influence on Hitchcock
must be acknowledged too. His Ph.D. thesis (1931) was on the solana-
ceous genus Lycium; he is still called on to arbitrate problems in the
group.

Hitchy’s long and colorful teaching career started at Pomona College
(1931-32); he soon moved to the University of Montana where he
taught general botany and taxonomy. We still encounter former students
(many now in forestry and range management) who vividly remember
the demanding pace (pedagogical and physical) that Hitchy set for
them, yet always generously laced with fun and games. It was at Mon-
tana that he produced a field manual on grasses and grass-like plants
which showed early in his career his dual skills—as a meticulous tax-
onomist and as a consummate artist-draftsman. The Montana epoch
also saw the beginning of the unique Summer Field Courses which
Hitchy continued to run when he came to Seattle in 1937. Beginning
then in the Department of Botany at the University of Washington
where he soon became chairman, the legend of the man continued to
grow. The remarkable ‘espirit’ that has made the department such a
friendly, livable habitat for botanists is surely traceable to the Hitch-

387

[Vol. 20

MADRONO

388

Leo HIitTcHCOCK

1970] C. LEO HITCHCOCK 389

cock inoculum both while he was chairman and in more recent years
when he returned to full-time teaching and research. For the many
graduate students in taxonomy who began their professional careers with
Hitchy, the frequent gay soirees hosted at their home by Evelyn and
her man make memorable embellishments on the fabric of an educa-
tional experience with Hitchcock.

For “years and years and years,” Hitchy has taught general botany
to foresters, elementary taxonomy (local flora, a course open to and
eagerly sought by all), as well as ornamental plants and advanced tax-
onomy. Not only have generations of young students benefited from
the experience of the local flora course from Hitchy, but repeatedly
through the years have adults in the continuing education program come
to count on his evening and weekend offerings. And, of course, the
famous Summer Field Trips continued for many years. Who could forget
a typical day for the field party, camped out miles away from the nearest
town? Up at 5:00 a.m. to the din of kitchen-pot cymbals and a hearty
breakfast of Hitchcock pancakes or biscuits. Then with hardly a breather,
off into the field they would go, students trying hard to keep up with
the master who fired off binomial salvos and sundry anecdotal items
about the plants in the area. All the while, his assistants were ‘making
hay,’ putting up herbarium material in substantial replicate for the UW
Herbarium and for exchange. To finish the day in camp, some violent
sport, like touch football took over with Hitchy sparking the play. The
years that Draba was being monographed, each Summer Field Class
literally ran up and down most of the high peaks in the West.

During the war years when there was little demand for classes in tax-
onomy, Hitchy and his close friend and field companion, Clarence V.
Muhlick used their rationed gas to put up thousands of plants from the
most inaccessible places throughout the Pacific Northwest. These col-
lections have formed the solid nucleus for floristic study of the region.
Nearly every major herbarium in the U. S. and elsewhere surely have
“Hitchcock and Muhlick” sheets—always skillfully prepared (including
pressed mosquitos) and amply documented.

In the early 1950’s, Hitchcock joined his taxonomic talents with those
of Arthur Cronquist, Marion Ownbey, and J. W. Thompson to begin the
projected five-volume illustrated flora of the Pacific Northwest. One by
one the volumes have appeared, beginning with Volume 5 (the Com-
positae by Cronquist), and culminating in 1969 with Volume 1. Modesty
would get in the way of Hitchy’s accounts of his contribution to this
monumental work, but we all well know that he has been the chief and
most persistent organizer-catalyst-editor-caretaker of the project as well
as author of many substantial families in the flora. The botanical public
will be pleased to know that a one-volume abridgement is nearing com-
pletion; Hitchy, again in collaboration with Cronquist, has devoted his
energies to all phases of this long-awaited condensation.

390 MADRONO [Vol. 20

So we salute you, C. Leo Hitchcock. We wish you many more years
of productive taxonomic output, and hope the years ahead also give you
ample time for gardening, for responding to the seasonal call of the
game birds, and for the perpetual rejuvenation that you may deservedly
derive from continued contact with your many, many friends—students,
fellow botanists, neighbors, and all others who have drawn from your
well of friendship. — A. R. K.

IRIS PSEUDACORUS IN WESTERN NORTH AMERICA

PETER H. RAVEN and JOHN H. THOMAS

Iris pseudacorus L. has been well established in swamps and other wet
habitats in eastern North America for nearly a century (Cody, 1961).
In western North America this showy, yellow-flowered /ris is of more
recent introduction, and as in the east seems destined to spread even
farther.

Preece (1964) has reported it from several localities in western Mon-
tana and it has since become increasingly more common in glacial pot-
holes, along ditches, and marshy areas in the Mission Valley in Lake
Co. (Thomas 11020, DS, US; Woodland 319, DS). In British Columbia
it is known from Lulu I., near Vancouver (Beamish & Vrugtman 60540,
OSC). In Oregon it has been found in Columbia Co. (Wolrod s. n., OSC)
and in Benton Co. (Merkle & Merkle s. n., OSC).

Hitchcock, et al. (1969), in summary, recorded this species in this
Pacific Northwest as “‘well established in our area in many lakes and
ponds and along rocky stream banks.”

In California the first record was probably that of Mason (1957),
who reported /. pseudacorus from Merced Co. Subsequently Rubtzoff
(1959) recorded it from Forestville, Sonoma Co. (Rubtzoff 1836, CAS,
RSA; 1258, CAS; 1813, CAS; 1549, CAS; 1946, CAS). Other California
localities are: Searsville, San Mateo Co. (Rubtzoff, 1959; Thomas, 1961)
(Thomas 7165, CAS, DS; 9221, DS); Mettlers Station, Kern Co.
(Munz, 1969; Twisselmann, 1963; 1967) (Twisselmann 8028, CAS) ;
near Yountville, Napa Co. (Thomas 15027, DS); Santa Cruz, Santa
Cruz Co. (Rubtzoff, 1959; Thomas, 1961) (Hesse 2764, DS) near
Montague, Siskiyou Co. (Rubtzoff, 1959) (Howell 28360, CAS); and
Lyons Springs, Ventura Co. (Rubtzoff, 1959) (Pollard s. n., CAS).

Mason (1959) remarked that /. pseudacorus “is apparently moving
down the watercourses.” This prediction has been fulfilled. T. C. Fuller,
of the California Department of Agriculture, has told us that there are
dense colonies of this species all along the Merced River in Merced Co.
Along Dana Slough west of Snelling, Fuller noted that this /vis was the

1970] RAVEN & THOMAS: IRIS 391

most common species of marsh plants, growing to the complete exclusion
of Typha and other characteristic California marsh plants.

In the Delta Region, 7. pseudacorus still grows as relatively small
isolated clumps and local populations. During April 27-28, 1969, one
of us (PHR) observed it at the following points near and just east of
the Franks Tract in the delta of the San Joaquin River, Contra Costa
Co.: Sand Mound Slough, about 1.3 miles southwest of Franks Tract;
Rock Slough, about 1.2 miles east of junction with Sand Mound Slough;
southwest corner of Quimby I.; and two clumps about 0.4 miles apart
at the southeast end of Mandeville I. It is probably much more widely
distributed in the Delta Region than these sight observations would
indicate.

It seems worthwhile to record these occurrences as there is every indi-
cation that this /ris will spread and displace many native plants. It is,
of course, regrettable to see the populations of native species declining
in the face of this new alien, which apparently spread from moist gar-
dens, but one can at least be grateful that they are losing ground to
such an attractive plant.

Kenton L. Chambers, Reid V. Moran, and Robert R. Thorne have
kindly supplied information about specimens of /. pseudacorus.

Department of Biological Sciences, Stanford University, Stanford, California

LITERATURE CITED

Copy, W. J. 1961. Iris pseudacorus L. escaped from cultivation in Canada. Canad.
Field-Naturalist 75:139-142.

Hitcucock, C. L. et al. 1969. Vascular plants of the Pacific Northwest. Vol. 1. Univ.
Washington Press.

Mason, H. L. 1957. A flora of the marshes of California. Univ. Calif. Press, Berkeley.

Mouwz, P. A. 1968. Supplement to a California flora. Univ. Calif. Press, Berkeley.

PREECE, S. J. 1964. Iris pseudacorus in Montana. Proc. Montana Acad. Sci. 24:1-4.

Rustzorr, P. 1959. Iris pseudacorus and Caltha palustris in California. Leafl. W.
Bot. 9:31-32.

Tuomas, J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford
Univ. Press.

TWISSELMANN, E. 1963. Plant records in Kern County, California. Leafl. W. Bot.
10: 46-64.

—. 1967. A flora of Kern County, California. Wasmann J. Biol. 25:1-395.

NOTES AND NEWS

MaproNo.—Please send all manuscripts intended for publication in Madrofio to
Dr. Robert Ornduff, Department of Botany, University of California, Berkeley,
California 94720.

The titlepage and index for Volume 20 of Madrono will be mailed with an early
number of Volume 21.

Numbers 1, 2, and 3 of Volume 21, will appear very shortly and publication will
then be on schedule.

A NEW SPECIES OF PROBOSCIDEA (MARTYNIACEAE)
FROM BAJA CALIFORNIA, MEXICO

RICHARD H. HEVLY

Annetta Carter recently asked me to annotate some specimens of
Proboscidea which she had collected in Baja California. Besides P. al-
theifolia (Benth.) Decne., a yellow-flowered, tuberous-rooted perennial,
these included a purple-flowered annual superficially resembling P. par-
viflora (Woot.) Woot. & Standl. Careful study, including comparison
with the type specimens of all described species of the genus, convinced
me that the latter specimens represent a new species endemic to the Sierra
de la Giganta of Baja California. This new species is described below
and compared with the other annual species from the Sonoran Desert

(fig. 1).

Proboscidea gracillima Hevly, sp. nov. Herba annua, glandulosa ad
viscido-pubescens, ramosissma, ad circa 3 dm lata et 4.5 dm alta; folia
opposita vel alterna, ovata vel deltoida, basibus cordatis, marginibus
integeris vel inaequaliter dentatis, petiolis 5-10 cm longis; racemi ter-
minales, 3—5—floribus, pedicellis per anthesin tenuissimis, circa 2.5—3.5
cm longis, sed maturitate crescentibus ad circa 4 cm longitudine; calyx
0.9-1.4 cm longus, 5—lobatus, ad basim ventraliter fissus, basi bracteo-
lis duabus, ovatis vel oblongis; corolla oblique infundibularis, 2.4—3.5
cm longa, limbo 5 lobato, 1.6-3.0 cm lato, luteo-violaceo; stamina fer-
tilia 4, didynama, quinto rudimentario; stylus stamina superans; stig-
mata duo; fructus ovatus, circa 9 cm longus.

Viscid to glandular pubescent annual herb arising from a well-devel-
oped tap root with fibrous secondaries; stems 30-45 cm in height, the
branches and leaves opposite or subopposite; petioles 5-10 cm long,
glandular-pubescent on the nerves below; basal leaves 3.75-6.75 cm
long and 3.25—6.50 cm wide, broadly ovate to deltoid, entire or with a
very shallowly undulate (sinuate) margin; inflorescence racemose, 10-18
cm long, only one- or two-flowered at any one time but ultimately pro-
ducing about 20 flowers, pedicels in anthesis erect to ascending, 2.5—3.5
cm long, 1 mm thick but becoming reflexed, thicker (2-3) mm, and
longer (up to 4 cm) in fruit; bracts 2, broadly ovate, oblong or falcate,
4—5 mm long and 2—3 mm broad; calyx thin, papery 0.9 to 1.4 cm long
and 1.0 cm broad, 5 lobed, the terminal lobe extended, the lateral lobes
3-5 mm wide, the intermediate and basal lobes 9 to 1.3 mm long, the
sinus 4 to % the length of the calyx; corolla 2.4-3.5 cm long, strong-
ly ventricose, (dorsal and ventral measurements differing by as much as
1 cm), the narrow portion of the tube 0.5—1.0 cm long, the flaring por-
tion 1.5—2.5 cm long and 1.1 to 2.0 wide at the mouth, reddish purple

392

1970] HEVLE: PROBOSCIDEA 393

Fic. 1. Holotype of P. gracillima. A young specimen with immature fruit.

externally. Yellowish purple within, with conspicuous purple dots, a pro-
nounced yellow band extending somewhat over the lower lobe, the limb
1.6-3.0 cm broad, reddish purple, the upper lobes maroon, the lobes
0.5-1.0 cm long and 0.7—1.2 cm broad; stamens 4, didynamous; ovary
1 celled with two parietal placentae, style exceeding the stamens, stig-
mas 2, ovate and sensitive; fruit an ovate to elliptical ligneous drupe
4—5 cm long with a prominent dorsal crest and horns 4—5 cm long; seeds
numerous (fig. 1).

[Vol. 20

MADRONO

394

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1970] BARNEBY: ASTRAGALUS 395

Type. Mexico: Baja California, Mesa del Potrero de San Javier (north-
east of Mission San Javier), Carter 4993 (MEX, UC-holotype), Sept.
20, 1965.

Other collections. Sierra de la Giganta, Carter 3137 (UC), Carter
4478 (UC), Carter 5289 (UC). These and the type came from eleva-
tions between 500 and 700 m. North of Comondu, Hammerly 179 (DS,
We):

Proboscidea gracillima belongs to section Proboscidea (=Sect. Eupro-
boscidea Stapf) (Stapf, 1895), characterized by annual habit and purp-
fish, reddish, pinkish, or whitish flowers. It is most similar to P. parvi-
flora vegetatively and keys to that species in the most recent revision
of the genus (Van Eseltine, 1929). In internal throat ornamentation and
in infloresence, however, it shows some affinity to P. sinaloensis Van
Eselt. All three species have small calyces and may be distinguished by
leaf shape, infloresence characteristics, corolla size and color, and fila-
ment pubescence, as well as by geographical distribution (table 1).

Northern Arizona University, Flagstaff

LITERATURE CITED

Stapf, O. 1895. Martyniaceae. Jn Engler & Prantl, Naturlichen Pflanzenfamilien
4(3b) :265-269.

Van ESELTINE, G. P. H. 1929. A preliminary revision of the unicorn plants (Mar-
tyniaceae). New York Agric. Exp. Sta. Bull. 129.

A NEW ASTRAGALUS (FABACEAE) FROM NEVADA
R. C. BARNEBY

Astragalus phoenix Barneby, sp. nov., in sect. Argophyllis juxta A.
newberryi Gray a quo habitu multicipiti late pulviniformi nec simplici-
us caespitoso, pube crasse tomentoso-pilosa candidissima, racemisque
brevissime pedunculatis 1—2 (nec 3—8)-floris absimilis inserenda.

Diu perennis subacaulescens e radice perpendiculari valida, caudicis
iteratim ramosi ramulis superne stipulis petiolisque marcidis crebre ob-
sitis columnaribus, demum pulvinos hemisphaericos vel depresse-con-
vexos ad 4—5 dm usque latos efformantibus, tota pilis patulis rigidius-
culis (minime gossypinis) ad 0.8—1.3 mm longis piloso-tomentosa cana;
stipulae crebre imbricatae ovatae acutae vel breviter acuminatae 2—3
mm longae, extus tomentosae, intus glabrae venosae; foliorum 1.5—
3.5(4) cm longorum petiolus rigidus marcescens, foliola 1—4-, saepis-
sime 2 vel 3-juga ovata vel obovata (2)3—6(7) mm longa secus rachin
2—10(15) mm longum conferta, mox decidua; pedunculus utriusque
ramuli unicus erectus brevissime 1—2-florus 2—5 mm longus stipulis
fulcrantibus ad maximum duplo longiribus; calycis laxe pilosuli 12.5—

396 MADRONO [Vol. 20

Fic. 1. A mature plant ef Astragalus phoenix, which furnished the type speci-
men. The cushion of foliage is approximately 3.2 dm in diameter. Photograph by
A. Cronquist.

15 mm longi tubus cylindricus subtumescens 9.5—11 mm longus, 4—4.6
mm diametro, dentes subulati 3—4 mm longi; petala pallide lilacina, vex-
illo pallidori, omnia sicca straminea, quoad formam illis A. newberrw
simillima; vexillum 24—25 mm longum, 9.5—11 mm latum; alae = 20.5
mm longae; carinae 19—21 mm longae unguiculi = 11.5 mm, laminae
lunatim semi-ellipticae 8—8.5 mm longae obtusae; antherae 0.7—0.8
mm longae; legumen eum A. newberryi exacte simulans, ovoideo-acu-
minatum ultra medium incurvum + 1.8 cm longum, 1 cm diametro,
valvulis coriaceis simul tomentulosis ac pilosis, pilis brevioribus densis
longioribus patulis nitidis ad 2 mm usque longis; ovula + 32.

Type. Nevada: Nye Co., on barren, alkaline, white clay slopes over-
looking a dry wash at the east end of Ash Meadows, elevation 2300 feet,
Township 18 S, Range 50 E, section 1 or 12, Cronquist 10657 (BRY,
NY-holotype, RSA, UTC), April 21, 1966.

Additional specimens. Nevada: Nye Co., with Enceliopsis nudicaulis
and Distichlis stricta, on dry, hard, alkaline flats, Ash Meadows between
Big Spring and Point of Rocks, elevation 2280 feet, Roos & Roos 6143
(NY), June 13, 1954; Ash Meadows, Purpus 6034 (POM). The three
localities are probably all close together, possibly all the same, lying in
the southern angle of Nye Co. close to the California boundary.

The Ash Meadows Milk-vetch, A. phoenix, was first collected, in
fragmentary specimens, as long ago as the summer of 1898 by Carl

1970] BARNEBY: ASTRAGALUS 397

Anton Purpus, who crossed Pahrump Valley on his adventurous journey
across the then pathless Mohave Desert into the botanically unknown
mountains of southern Nevada. More complete material was gathered
by Roos and Roos in 1954, but again too late in the year (mid-June) to
show more than withered flowers and dehiscent pods. I have already
referred to theses two collections (Barneby, 1964) as representing a
species related to A. newberryi Gray but probably undescribed. The fine
flowering specimens now before me, complemented by field-notes and
the photograph reproduced herewith, confirm this conjecture.

Detached from the plant, the individual flower and pod of A. phoenix
(born of ashes) cannot be distinguished from those of typical large-
flowered A. newberryi. The average mature plant of the latter consists
of some one to five, exceptionally a dozen scarcely elongating rosettes
of leaves gathered into a tuft sessile or nearly so on the root-crown; if
caudex-branches develop, they remain short, always shorter than the
longest leaves, and are simple or little ramified. The pubescence of the
foliage is variable in quality and orientation, but the young, newly ex-
panded leaves are always silvery-silky with shining hairs. The flowers
are only exceptionally less than three to the raceme and are elevated on a
scapelike peduncle seldom less than 1 cm long. As the flowers fade the
peduncle bends outward, and the pods ripen in contact with the ground.
By contrast the mature plant of A. phoenix forms a dense hemisphere
or depressed mound of foliage that reaches a diameter of 4—5 dm and
is composed of several score, perhaps over 100 rosettes of leaves. The
caudex is repeatedly branched, becoming several times longer than the
longest leaf. Deep within the cushion, impacted with white clay, the
older branches are brown and woody, clothed in a flaking bark, but
distally become columnar from the thatch of tomentose stipules and
stout persistent leaf-stalks. Already at early anthesis the pubescence is
composed of relatively coarse, spreading hairs, the general effect of which
is white-tomentose rather than silky. The flowers, pink-purple with a
paler banner, followed by the pods, sit apparently stemless, one or two
together, among the leaves. The permanently erect peduncle is at most
5 mm long and often scarcely surpasses the subtending stipules.

It seems probable that A. phoenix is derived by specialization from
A. newberryi. The species is adapted and very likely confined to a pe-
culiar habitat of calcareous flats and knolls on the valley floor, a habitat
that provides a home for some other pulvinate species of the Nevadan
deserts such as Lepidium nanum Wats., Eriogonum shockleyi Wats., as
well as for some pulvinate ecotypes of ordinarily cespitose Astragalus
calycosus Torr. ex Wats. and Oxytro pis oreo phila Gray. The elevation of
Ash Meadows is near 2280 ft (685 m). In the mountains of the eastern
Mohave Desert and the Death Valley region A. newberryi is not un-
common on limestone formations in the pinyon-belt, but has not been
collected and cannot be expected below an elevation of about 5000 ft

398 MADRONO [Vol. 20

(1500 m). Three other densely pubescent Argophylli occur near A.
phoenix: A. coccineus Brandg., A. funereus Jones, and A. purshii var.
tinctus Jones. The first of these has in common with A. phoenix per-
sistent petioles and coarse pubescence, but has more numerous and
longer red flowers elevated on long peduncles. The other two have finer,
cottony pubescence, soft petioles, and three or more flowers borne to-
gether, again on developed scapes. The pod of the nearly sympatric A.
funerus is much larger, 3—5 not 2 cm long; that of A. purshii var. tinc-
tus is in the same size-range as that of A. phoenix, but the whole ap-

pearance of the plant is quite different.
New York Botanical Garden, New York

LITERATURE CITED

Barnesy, R. C. 1964. Atlas of North American Astragalus. Mem. New York Bot.
Gard. 13:1-1188.

NOTES ON LOEFLINGIA (CAROPHYLLACEAE)
R. C. BARNEBY and ERNEST C. TWISSELMANN

INTRODUCTION

The small caryophyllaceous genus Loeflingia is of interest to plant
geographers because of its bicentric dispersal. In the Old World its
center of abundance coincides with the western end of the Mediterranean
basin, with greatest concentration of variability and of numbers in the
southern and eastern quarters of the Iberian Peninsula and in northern
Morocco and Algeria. In Africa it extends south into the Sahara, but
from the Mediterranean coast eastward from the longitude of Malta
there are only a few scattered records of the common species, L. /is-
panica L. In the Old World, Loeflingia is clearly a west-Mediterranean
type. The range of the genus in North America is less extensive but
more discontinuous. Representatives occupy four well defined floristic
provinces, one east and three west of the Continental Divide: 1, east-
central Texas north, interruptedly, to western Nebraska; 2) floor of the
Sonoran Desert in southern Arizona and northern Sonora; 3, the Basin
and Range sagebrush deserts of northeastern California, southeastern
Oregon, and southwestern Wyoming; and 4, cismontane California
southward from Santa Cruz and Stanislaus counties into northern Baja
California. Wherever they occur, the loeflingias are associated with light,
often disturbed or wind-modified, commonly sandy soils, and show
marked tolerance or even preference for genuine dune habitats. They
appear intolerant of competition and tend to occupy microhabitats in
which most plants have difficulty in taking foothold.

It was early suggested by Hooker (1840) and by Brandegee (1890)
that Loeflingia might not be native to America, but this view is un-
tenable. Variation in our plants is plainly correlated with familiar dis-

1970] BARNEBY & TWISSELMANN: LOEFLINGIA 399

persal patterns, a situation that could not be expected of immigrant
weeds. Furthermore the Old World loeflingias differ from ours in hav-
ing at once smaller capsules, larger petals, and longer styles; and
although the mode of branching is alike everywhere in basic design
there are differences in ratio of the main internodes to length (and
density) of the monochasial cymes beyond the initial dichotomies of
the stem that give the North American and Mediterranean plants a
subtly different aspect. However, as Hooker remarked (1840), when he
described L. texana, the species are so similar in general organization
that a case could be made for treating all as races of the original L.
his panica.

While the generic range of Leoflingia is probably now well worked
out in broad outline (if not yet in fine detail), the taxonomy remains
in a fluid state. Comparison of two recent accounts of the Old World
species (Maire, 1963; Heywood, 1964) and of the modern floras cov-
ering the Intermountain United States and Sonoran Desert in America
make this very clear. Our interest in the Mediterranean species is
aroused by the similarity of the problems presented by Leoflingia in its
two main areas of dispersal, but we lack the material to pursue it. The
objectives of this study are to determine: a, the nature, and the rela-
tionship to LZ. squarrosa Nutt., of the long known but mysteriously local
L. pusilla Curran, reported only from Kern Co., California; and b) the
identity of the loeflingias found in the Sonoran Desert, and of similar
plants encountered in widely scattered stations in the Intermountain
region northward. Along the way we have been obliged to reexamine
all the North American species.

HIsToRY OF LOEFLINGIA IN AMERICA

The Loe flingia first described from America was L. squarrosa (Torrey
and Gray, 1838-1840), based on a plant collected by Nuttall near San
Diego, California, in 1836. The genus had actually been discovered
slightly earlier, in 1833 or 1834, by Drummond in Texas. Hooker (1840)
described and figured Drummond’s plant as L. texana, a proposition
promptly reduced by Torrey and Gray (1838-1840) to ZL. squarrosa.
Matters rested there until Mary K. Curran (Katharine Brandegee)
(1885) described LZ. pusilla from plants collected the previous summer
along the railroad west of Tehachapi. This was said to differ from ZL.
squarrosa in being ‘““much more delicate” and inferentially in its short,
straight, entire sepals; it was further noted as pentandrous and apeta-
lous. The only other described American species, L. verna Nelson, is
Arenaria pusilla Wats. (Loeflingia verna Nelson, Bot. Gaz. (Crawfords-
ville) 54:138. 1912, “Secured by Macbride . . . near New Plymouth,
[Idaho], April 24, 1911, no. 773.”, DS!, RM!—hototype).

Systematic literature dealing with the genus in America is meager.
It appeared in Gray’s Genera (1849) where L. texana is illustrated

400 MADRONO [Vol. 20

under the name L. squarrosa. Robinson (1893) furnished a key to the
three described species, distinguishing L. pusilla from the rest by its
toothless outer sepals and separating L. squarrosa from L. texana by
supposedly smaller stature, less secund branching, recurving sepals, and
oblong-elliptic rather than obovate seeds. According to Robinson all
Loeflingia in the New World should be triandrous, in contradiction to
the protologues of ZL. squarrosa and L. pusilla and to Gray’s account of
the genus just mentioned. This point has long been in dispute. Bran-
degee (1890) had already remarked that the flower of L. pusilla was
triandrous, although this is untrue of the one individual plant known
to survive out of the type-collection. We now know of apparently con-
stant pentandrous populations, of pentandrous individuals in largely
triandrous populations, and of individual flowers varying in number of
stamens on a single plant.

During the present century it has been customary to follow Robin-
son in maintaining L. squarrosa and L. texana as distinct species, but
their ranges are so well separated that no occasion has arisen to com-
pare them critically. With the few exceptions mentioned directly, L.
squarrosa has been treated as endemic to cismontane Califorina or at
least to the California floristic province. Peck (1941) extended the
range of L. squarrosa to interior Oregon in Harney Co. Its presence in
Arizona was confirmed by Kearney and Peebles (1942). However, Shreve
and Wiggins (1964) subsequently referred the Arizona plant to ZL.
texana, a species not recorded otherwise from west of the Divide. In
floras that cover all of California, Jepson (1914), Abrams (1944), and
Munz (1959) agree that L. pusilla is known only from Tehachapi, and
all published accounts of this species, up to a recent report by Twissel-
mann (1967) from the western Mojave Desert in Kern Co., go back to
the type collection, for which no exact match has ever been encount-
ered. Before presenting our views on the taxonomy, we propose to dis-
cuss briefly the comparatively few phenotypic characters that have been
used or promise to prove useful in delimiting taxa.

HABIT AND MODE OF BRANCHING

Branching in Loeflingia is of two types, one preceding the other:
strictly dichotomous, when the primary axis divides into two branches
of equal vigor, usually containing a sessile flower in the fork; and
monochasial, as one branch of the dichotomy becomes reduced or obso-
lete at several successive nodes. Dichotomy may start directly from the
axils of the cotyledons, or beyond several internodes of a simple axis,
and distally gives way, either abruptly or gradually, to a more or less
pronounced monochasial mode. When the distal monochasia are well
developed, they form fan-shaped sprays; when poorly developed, each
may be reduced to a single flower and the inflorescence comes to sim-
ulate a spike. The position of the first dichotomy and the relative size

1970] BARNEBY & TWISSELMANN: LOEFLINGIA 401

of the monochasia together determine the aspect of the plant, an aspect
which is often characteristic of all members of a population and also
tends to be dominant over large areas. We have developed no objective
formula for describing the permutations of branching, but believe that
the intangible quality of habit permits intuitive sorting of material into
categories that coincide with comprehensible dispersal patterns. Stature
of the individual plant is governed to some extent by variations in rain-
fall from year to year, as observed at several stations in Kern Co. On
the other hand maximum or potential stature is to some degree genet-
ically controlled, for none of the desert loeflingias, however favorable
the season, seem ever to surpass three centimeters in height.
FLOWERS

The flowers of Loeflingia, at least in America, are cleistogamous. The
five sepals, the two outermost of which are often leaflike in form and
are perceived as sepals through their position and (occasionally) by being
accompanied by an opposed stamen, connive over the ovary throughout
anthesis. The three, four, or five filaments are closely contained between
the sepals and the ovary and the filaments are exactly long enough to
elevate the minute anthers to the level of the (in America subsessile)
stigmas. Pollination is automatic, and ordinarily (disregarding some
terminal buds which never reach maturity) is 100% effective, for it is
rare indeed to find on any plant a single infertile flower. Obligate autog-
amy gives rise to internal homogeneous populations which, immune to
the leveling effects of outcrossing, hand on intact an indefinitely re-
duplicated genetic structure. The resulting phenetic uniformity of pop-
ulations has tended to elevate the apparent taxonomic value of minor
characters and permits the elaborate but, we suspect, artificial heir-
archy of subspecies and formae worked out by Maire (1963) for L. his-
panica sens. lat. in North Africa. In America few such characters find
uniform expression over any considerable land area, and there is only
limited correlation between any pair of them. The reduced flower of
Leoflingia, commonly apetalous in the American forms or with petals
represented by vestigial and somewhat amorphous scales, presents few
taxonomic features: length and curvature of sepals; presence of lateral
teeth on 2 or more sepals; and stamen number.

Within populations as represented by herbarium specimens we have
seen little variation in length of the outer sepals, which are, however,
normally longer than the three inner ones. The variation between popu-
lations is marked. In cismontane California the longest sepal of L. squar-
rosa sens. str. varies from approximately 3 to 6 mm in length, that is
from a trifle longer to twice as long as the capsule. The curvature of the
sepal seems largely a function of its length, modified by age. Sepals
nearly straight in bud become squarrose as the fruit ripens, but curva-
ture occurs mainly in the part projecting beyond the capsule. It fol-

402 MADRONO [Vol. 20

lows that the longer the sepal the greater the curvature. Length and
curvature of the sepals seem to bear some relation to geographic dis-
persal but are poorly linked with other morphological characters and
we believe that they have been overestimated as taxonomic criteria.

A filamentous or setiform tooth or spur arising on each side of at least
the two outermost sepals is a generic character of Loeflingia. The tooth
appears homologous with what is generally interpreted, in the cauline
leaves, as the free tip of a partially adnate stipule. Rare individuals
that lack all such appendages become technically indistinguishable
from Minuartia. The sepal teeth vary considerably in length and stout-
ness, and the stipules vary with them, suggesting that they are under
the control of the same gene or genes. In America the outer sepals are
nearly always appendaged, the three inner ones very rarely so. Usually
the toothed outer pair do not subtend a functional stamen and then,
because they are not only longer than the three inner but also entirely
leaflike in shape and texture, they simulate bracts enclosing a trimerous
flower. Occasionally, however, the presence of an opposed stamen reveals
the sepaloid nature of the outer pair. In those rare instances where the
sepal teeth are obsolete, and the outer pair of sepals become obviously
sepaloid and unleaflike, the flower becomes pentandrous. Thus the pas-
sage from true leaf into segment of the perianth is not marked by the
usual discontinuity. We have learned to regard the presence of teeth on
2,3, or 5 sepals and the occasional absence of teeth from all sepals as in
the nature of individual variation, linked neither with geography nor
with other characters. On the other hand the longer and stiffer setae on
both leaves and sepals of the Texan loeflingias contribute materially
toward the distinct facies of the populations east of the Continental
Divide.

We have noted the correlation between a pentandrous flower and
sepaloid sepals and between a triandrous flower and a pair of foliaceous,
toothed exterior sepals. The latter combination is far the commoner
everywhere in America, universal (so far as we have observed) east of
the Divide. In cismontane California we find only sporadic instances
of more than three stamens, and these sometimes occur on the same
plant with triandrous flowers, for example, Los Angeles Co., Newhall,
Pringle, NY; San Diego, Orcutt, NY; the condition described by Torrey
and Gray for the original L. squarrosa though since denied. In the Inter-
mountain region there are populations fully pentandrous (Ripley &
Barneby 7938, CAS, NY), partly triandrous and partly pentandrous
(Honey Lake, Brandegee, UC), and wholly, so far as sampled, tri-
androus, but no correlation with other phenetic characters has been dis-
covered. Probably all loeflngias are primarily triandrous but retain a
latent potentiality for return to what is presumably the primitive pent-
androus condition.

1970] BARNEBY & TWISSELMANN: LOEFLINGIA 403

CAPSULE AND SEED

West of the Sierra crest in California the capsule of Loeflingia is
narrowly ovate to lanceolate in profile, with width-length ratio of
approximately 1:3—4. All capsules of drought-inhibited individuals and
late capsules of vigorous plants tend to be smaller than average but no
wider proportionately. On the deserts and east of the Divide the capsule
is ovate in profile, with width-length ratio of 1:2—2.5. In both areas
some capsules lie inconveniently in the ratio of 1:2.5—3, but these are
generally longer absolutely if from west of the Sierra, absolutely shorter
if from the east. There is no abrupt discontinuity, but nevertheless we
have found the capsule-outline more helpful than any other character
in delimiting typical L. squarrosa. As shown below, it was crucial in our
disposition of the litigious L. pusilla.

Robinson (1893) was the first to notice differences in size and out-
line of the seeds of ZL. texana and L. squarrosa, the former being shorter
and plumper. Within a given capsule the seeds are virtually uniform in
size, and within the population the difference, if any, is barely percept-
ible. Seeds of cismontane California Loeflingia are 0.4—0.5 mm, rarely
up to 0.55 mm long; of ZL. texana 0.3-0.4 mm long. In the Sonoran
Desert the seeds fall within the size-range of L. texana; in the Inter-
mountain region within that of Z. squarrosa. In conjunction, the capsule
and seeds, as so often in Caryophyllaceae, furnish useful, even though
not infallible taxonomic criteria.

CONCLUSIONS

When Robinson contrasted a Californian ZL. squarrosa characterized
by long, recurved sepals and relatively long seeds with a Texan L. texana
differing in its relatively straight sepals and short, plump seeds, he had
for comparison no material at all from intervening territories and only
a few specimens even from California. Plants collected since his day,
especially on the deserts, have effectively blurred the supposed morpho-
logical discontinuity between the loeflingias of the Pacific and Atlantic
slopes. The Arizona plants, which have been referred because of their
recurving sepals to L. squarrosa and alternatively because of their small
seeds to L. texana, are neatly intermediate in terms of Robinson’s cri-
teria. They lean somewhat to L. texana in their relatively stiff and long
sepal teeth, but differ greatly in their diminutive stature and their habit
of dense dichotomous branching which starts from the cotyledons or
from the first node. This growth habit is equally alien to L. squarrosa
and L. texana, but is nearly duplicated in the loeflingias found in the
Intermountain region to the north. The latter, now known from five
mutually remote areas of small extent, three in transmontane California,
one in southeastern Oregon, and one in southwestern Wyoming, have
characteristically short, straight sepals combined with the seeds of

404 MADRONO [Vol. 20

L. squarrosa in the capsule of L. texana. Because of overlapping and
recombination of the available criteria in these four geographic prov-
inces we believe the American loeflingias are reasonably interpreted as a
single species, L. sguarrosa, composed of four subspecies.

Key TO THE SUBSPECIES OF L. SQUARROSA NUTT. IN Torr. & GRAY

Capsule narrowly ovate to lanceolate in profile, (2.5)2.7-3.7 mm long, 0.8-1 mm
in diameter, the width-length ratio mostly 1:3-4; stems usually 3-10 cm long,
shorter only in depauperate individuals or in populations dwarfed by drought;
primary stem axis usually simple through 2-3 internodes, the first flower borne
at a point 5 mm or more distant from the cotyledons (this character fallible
especially in drought inhibited plants); distal monochasia mostly 1-flowered;
stipules and sepal teeth weak, short, filamentous, usually less than 1 mm long;
sepals variable in length, the longest 3-6 mm long, slightly longer to twice as
long as the capsule, when long becoming squarrose in age; cismontane Califor-
nia southward from Stanislaus and Santa Cruz counties to northern Baja
California, and one locality on the western Mojave Desert in Kern Co., where
sympatric but not intergrading with ssp. artemisiarum. . . . ssp. squarrosa

Capsule ovate in profile, (1.5)1.8-2.5(2.7) mm long, (0.7)0.8-1.2 mm in diameter,
the width-length ratio mostly 1:2—2.7(3); east of the crests of the Sierra
Nevada in California, to Wyoming, Nebraska, Texas, and Sonora.

Diminutive plants, stems to 3 cm long, often less, mostly dichotomous from the
cotyledons or from the first succeeding node, the first flower borne at a
point only 1—4(5) mm distant from the cotyledons; monochasia distal to
the dichotomies all reduced to one flower; seeds, sepals, and stipular setae
various; Intermountain states and Sonoran Desert.

Seeds 0.4-0.5 mm long; sepals always short, 1.8-4 mm long, not or little
recurved at tip; stipular setae and sepal teeth weak, filamentous, less
than 1 mm long (as in ssp. squarrosa); Intermountain United States,
southwestern Wyoming to southeastern Oregon and northeastern Califor-
nia, and on the western Mojave Desert in Kern and Inyo counties, Cali-

fornia... . . ssp. artemisiarum

Seeds 0.3- 0.41 mm © loner eae oct co 3.5- 5 mm “Jong, squarrose; stipular
setae and sepal teeth intermediate in length and rigidity between those of
ssp. squarrosa and ssp. texana; Sonoran Desert, from south central Ari-
zona to northern Sonora. . . . . . SSp. cactorum

Taller plants, the stems, unless deerme es 4-15 cm lone bearing the first
flower at points at least 5 mm from the cotyledons, following several simple
internodes; monochasia distal to the dichotomies more than 1-flowered,
tending to form scorpioid cymes; seeds small, 0.3-0.4 cm long; stipular setae
and sepal teeth stiff, subspinose, 1-1.5 mm long; east central Texas north-
ward interruptedly to western Nebraska... . . . . . . ssp. texana

LOEFLINGA SQUARROSA Nutt. in Torr. & Gray ssp. sQuarRosa, FI.
N. Amer. 1:174. 1838, “Sandy plains, St. Diego, California... Nuttall.’”,
NY!—isotype. L. Loic Curran, Bull. Calif. Acad. Sci. 12152. 1885,
“Tehachapi, Alt. 4,000 feet, May.’, UC!—isotype labelled: ‘Bet.
Tehachapi and Girard Station (now Marcel), along the railway, May,
1884. Pt of type, K.B.” The holotype collected in 1884 by M. K. Curran
was not found at CAS and was probably destroyed.

Thin sandy and gravelly soils, mostly below 2000 ft, sometimes in dry
stream-beds, in abandoned fields, waysides, and dunes, South Coast

1970]. BARNEBY & TWISSELMANN: LOEFLINGIA 405

Ranges of California inland from the ocean from Santa Cruz to Santa
Barbara Co.; margins of San Joaquin Valley and Sierra foothills from
Stanislaus to Kern Co., there ascending through the Digger Pine belt
to about 3900 ft; thence south through coastal and interior southern
California to San Bernardino Valley and western San Diego Co.; and
reportedly (Brandegee, 1890) to lat N. 28° in Baja California; and in
Kern Co. at 2450 ft on stabilized dunes around Buckhorn Dry Lake in
the western Mojave Desert, there sympatric with ssp. artemisiarum.

Representative specimens. California: Hoover 5130, NY, UC; Keck
& Stockwell 3354, DS; Twisselmann 2008, 2856, 8489, 13016, all CAS;
Brandegee, in 1909, CAS, DS, UC; Howell 5814, 29243, all CAS; How-
ell & Barneby 29424, CAS; Parish 4158, NY, UC; Brandegee, in 1898,
NYS, UC,

DISPOSITION OF L. PUSILLA

We have already noted that until 1967 all records in the literature of
L. pusilla are based on the type collection, which to date has never been
precisely matched. The holotype was presumably part of the California
Academy collection lost in the San Francisco fire in 1906. A single in-
dividual that survived labelled as authentic by K. Brandegee (UC) and
agrees perfectly with the protologue, up to the last detail of toothless
sepals and pentandrous flowers. We surmise that this specimen and T. S.
Brandegee s.n., 23 June 1892 (UC), from Honey Lake, California fur-
nished the model for Abrams’s Fig. 1718 (1944), of which the only
fault is that the capsule is enlarged twice as much as that of L. squarrosa
in Fig. 1717, thereby giving a false impression of the proportionate dif-
ferences. Repeated search around Tehachapi has yielded nothing quite
like L. pusilla, although ssp. squarrosa was collected as early as 1909 by
Brandegee at Keene, close to the type locality. Because the type of
L. pusilla was collected along the railroad we cannot discount the possi-
bility that it was a waif: but if waif it was, its origin will remain obscure
until it can be matched with some naturally occurring population. We
can affirm, in any case, that L. pusilla is not an introduced form or race
of L. hispanica. It can be accommodated without severe strain in our
concept of L. squarrosa because, although it possesses a unique combina-
tion of characters, some of them uncommon in the species, it has no
character unique to itself. The only outstanding question is whether to
refer it to the cismontane or transmontane Californian subspecies of
L. squarrosa.

The unique character combinations in L. pusilla are: a, small stature,
to 5 cm; b, dichotomies and flowers starting only 2 mm from the coty-
ledons; c, distal monochasia more than 1-flowered; d, obsolete or ves-
tigial stipule tips and sepal teeth; e, sepals all short, 3-3.5 mm; f, 5
stamens; g, slenderly ovoid capsule (+ 3 X 1 mm), and h, relatively
large seeds, between 0.5 and 0.55 mm. Of these characters a is easily

406 MADRONO [Vol. 20

matched in cismontane ssp. squarrosa, especially in dry years; the plant
is not “much more delicate” than many modern collections from the
Coast Ranges, but is taller than any known individual of ssp. artemisi-
arum. Characters c and g are normal for ssp. squarrosa, alien to ssp.
artemisiarum, and g (slender capsule) is in our opinion one of the best
diagnostic features of the former. Characters b, e, and f, are all known
to occur in ssp. squarrosa, but only exceptionally, and elsewhere not
together; only b and e are characteristic of ssp. artemisiarum; { is
sporadic in both. Character h, common to ssp. artemisiarum and ssp.
squarrosa serves only to exclude L. pusilla decisively from the more east-
ern races of the species (either of which could have been adventive at
Tehachapi). Character d has been seen in only one other Loeflingia,
Brandegee’s collection of ssp. artemisiarum from Honey Lake. We have
already suggested that loss of setae is an individual variation of no sys-
tematic importance. Thus we are led to believe that the type of L. pusilla
was an unusual variant of ssp. sqguarrosa in which several rare features
are combined. The plants cited as L. pusilla by Twisselmann (1967)
from the western Mojave Desert in Kern Co. are now interpreted as
L. squarrosa ssp. artemisiarum. The dunes around Buckhorn Lake are
the only station in which two subspecies of L. squarrosa are known to
grow together. Twisselmann (1967) has already observed that in this
locality the two seem to have slightly different ecological preferences,
the ssp. artemisiarum favoring the stiffer, more alkaline soils.

LOEFLINGIA SQUARROSA Nutt. in Torr. & Gray ssp. artemisiarum
Barneby & Twisselmann, ssp. nov. Habitu pumilo ssp. cactorum simu-
lans sed imprimis seminibus majusculis (0.4—0.5 mm longis) iis ssp.
squarrosae aequimagnis sepalis sesmper abbreviatis rectis capsulam
paullo superantibus absimilis, a ssp. squarrosa fere toto allopatrica prae-
ertim statura semper minima atque capsula latius ovoidea_ breviori
recedens.

Type. Oregon: Harney Co., sandy flats 3 miles south of Wright’s
Point, June 24, 1942, Morton E. Peck 21370, NY!, CAS!—holotype.

Dunes and sandy flats, often among sagebrush, mostly between 4000
and 7000 feet, northeastern California (Lassen and Plumas counties)
and southeastern Oregon (Harney Co.), to be sought in northern Neva-
da; southwestern Wyoming (upper Green River Valley in Sweetwater
Co.); also in Owens Valley in Inyo Co. and at approximately 2450 ft.
around Buckhorn, Rogers, and Rosamond dry lakes on the western Mo-
jave Desert in southeastern Kern and adjacent Los Angeles counties,
California (there sometimes associated with ssp. squarrosa).

Specimens examined. Wyoming: Sweetwater Co., 26 miles east of
Farson, Ripley & Barneby 7938, CAS, NY. Oregon: Harney Co., French
Glen, Peck 21419, CAS; 8 miles north of Narrows, Ripley & Barneby
6086, CAS. California: Lassen Co., Honey Lake, Brandegee, UC; Plumas

1970] BARNEBY & TWISSELMANN: LOEFLINGIA 407

Co., 5.8 miles east of Beckwourth, Howell, CAS; Inyo Co., near Bigpine,
Twisselmann 15537, CAS, NY, RSA; Kern Co., Buckhorn Dry Lake,
Twisselmann 10777, 10838, both CAS; between Old Pancho Barnes
place and Buckhorn Dry Lake, Twisselmann, McMillan, & Smith 14227,
CAS, NY; south end of Rogers Dry Lake, Twisselmann 10714, CAS;
Los Angeles Co., 5 miles north of Lancaster, Hoffmann, SBM.

LOEFLINGIA SQUARROSA Nutt. in Torr. & Gray ssp. cactorum Barn-
eby & Twisselmann, ssp. nov. Habitu deminuto, caulibus ex ipso basi
simul dichotomis ac florigeris ssp. artemisiarum proxima sed _ sepalis
elongatis demum recurvis et praesertim seminibus minoribus 0.3-0.4 mm
longis lis ssp. texanae aequimagnis absimilis. A ssp. squarrosa necnon
ssp. texana statura minima, habitu, monochasiis superioribus |-floris,
ulterius ab illa seminibus parvis recedens, ab omnibus affinibus deserto
Sonorensi incola allopatrica.

Type. Arizona: Pima Co., Sabino Canyon, Santa Catalina Mountains,
J. J. Thornber 5340, Mar. 26, 1905, CAS!, DS!, NY!—holotype.

Sandy and gravelly desert flats, sometimes in hard-packed soil of
ridges and adobe flood plains, below 3300 ft; south central Arizona in
Pima Co., and in Pinal and Maricopa counties according to Kearney and
Peebles (1951); and in the districts of Altar, Magdalena, and Hermo-
sillo in Sonora.

Specimens examined. Arizona: Pima Co., 16 miles north of Tucson,
Abrams 13100, DS. Sonora: 15 miles north of Magdalena, Fosberg 7468,
DS; 7 miles south of Sasabe, Keck 3970, DS; 10 km northeast of San
Pedro, east of Hermosillo, Ripley 14333, CAS, NY; north of Cumerol,
Abrams 13170, DS.

LOEFLINGIA SQUARROSA Nutt. in Torr. & Gray ssp. texana (Hook.)
Barneby & Twisselmann, comb. nov. L. texana Hook., Icones plantarum
3: tab. 275. 1840, “Interior of Texas. Drummond (3d Coll. n. 464)”,
presumed isotype, Drummond 464 (but “Coll. IT’), NY!

Sandy soils below 2000 ft; east central Texas, the lower Colorado
and Brazos valleys in Travis, Waller, and Colorado counties, and
around Dallas and Fort Worth, Dallas and Tarrant counties; north
central Oklahoma, the Cimarron Valley in Payne Co.; and greatly dis-
junct at about 3400 ft near the headwaters of the Niobrara River in
Dawes Co., Nebraska.

Representative specimens. Texas: Hall 480, NY; Wright 25, NY;
Reverchon distrib. Curtiss 346, NY; Lundell 14031; UC; Shinners
14650, CAS. Oklahoma: Waterfall 13170, CAS. Nebraska: Webber, in
1889, NY.

New York Botanical Garden, New York
Cholame, California

408 MADRONO [Vol. 20

LITERATURE CITED

ABRAMS, L. R. 1944. Illustrated flora of the Pacific States. Vol. 2. Stanford Univ.
Press.

BRANDEGEE, T. S. 1890. Loeflingia squarrosa Nutt. Zoe 1:219-220.

Curran, M. K. 1885. Descriptions of some Californian plants collected by the
writer in 1884. Bull. Calif. Acad. Sci. 1:151-155.

Gray, A. 1849. The genera cf the plants of the United States. Vol. 2. Putnam, New
York.

Heywoop, V H. 1964. In T. G. Tutin, et al. (editors), Flora Europaea. Vol. 1.
Cambridge Univ. Press.

Hooker, W. J. 1840. Icones plantarum. Vol. 3. Longman, et al., London.

Jepson, W. L. 1914. A flora of California. Vol. 1. Associated Students Store, Berkeley.

KearneEY, T. H., and R. H. PEEBiEs. 1942. Flowering plants and ferns of Arizona.
Government Printing Office, Washington.

. 1951. Arizona flora. Univ. California Press, Berkeley.

Marre, R. 1963. Flore de ]’Afrique du nord. Vol. 9. Paul Lechevalier, Paris.

Mouwz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

Peck, M. E. 1941. A manual of the higher plants of Oregon. Binfords and Mort,
Portland.

Rosinson, B. L. 1893. The North American Sileneae and Polycarpeae. Proc. Amer.
Acad. Arts 28:124-155.

Torrey, J., and A. Gray. 1838-1840. A flora of North America. Vol. 1. Wiley and
Putnam, New York.

TWISSELMANN, E. C. 1967. A flora of Kern County, California. Wasmann J. Biol.
25:1-395.

SHREVE, F., and I. L. Wiccrins. 1964. Vegetation and flora of the Sonoran Desert.
Vol. 1. Stanford Univ. Press.

UNUSUAL FACTORS CONTRIBUTING TO THE
DESTRUCTION OF YOUNG GIANT SEQUOIAS

Howarp S. SHELLHAMMER, RONALD E. STECKER, H. THomAs HARVEY,
and RICHARD J. HARTESVELDT

During the summer of 1966, a stand of dead and dying 10- and
11-year-old giant sequoias, Sequoiadendron giganteum, was discovered
in the Abbot Creek drainage of the Cherry Gap Grove of Sequoias in
Sequoia National Forest. This grove is located immediately south of
Converse Basin at 36°46’5” N lat., and 118°56’49’’W long.

This area was originally logged along with Converse Basin during the
latter part of the last century. Although evidence is lacking, it is felt
that the parent trees of the young sequoias in question were seeded at
the time of logging. In 1955, this entire area was consumed by an in-
tensely hot fire known locally as the “McGee Burn.” The parent trees
were killed by the fire, but disseminated the seeds which had apparently
remained viable in the green cones after the fire.

1970] SHELLHAMMER, ET AL.: SEQUOIA 409

Fic. 1. Stump of a 10 year old giant sequoia showing the base girdled by rodents
and the fruiting bodies of Hyphloma sp.

410 MADRONO [Vol. 20

Approximately 150 saplings were examined along a tributary of Abbot
Creek where soil moisture conditions were suitable for germination and
rapid seedling growth. With almost full sunlight, the young trees had
grown to ene varying from 5 to 15 feet.

Of the 150 saplings, 54 had been damaged by small rodents. Of these,
27 had been completely girdled and were dead, and the remainding 17
were partially chewed and displayed varying degrees of browning foliage.
There was no apparent selectivity for trees of any size class or crown
condition.

Judging from the tooth marks on the xylem, the rodents were small,
probably meadow mice, Microtus sp, or possibly gophers, Tomomys sp.
Bark from two of the trees was removed to a height of about 1% feet
above the ground, indicating that the feeding may have occurred during
the winter on the surface of the snow. Similar microtine damage has
been observed by the authors on Abies concolor nearby in Kings Canyon
National Park,, but it has not been previously recorded for the giant se-
quoia.

Nearly all of the dead saplings were being attacked by a fungus whose
bright yellow fruiting bodies clustered densely at the base of each tree
(fig. 1). Robert Bega of the Pacific Southwest Forest and Range Experi-
ment Station, and Lee Bonar of the University of California identified
the fungus as Hyphloma, possibly H. fasciculare, a known saprophyte.
This genus is previously unrecorded in association with sequoia. It seems
noteworthy that a species of tree whose remnants have lain undecayed
for as long as 2,000 years is here found in a state of decay within two or
three years after death. The wood of these stems was, however, largely
sapwood which lacked heavy deposition of tannin characteristic of old
heartwood.

Several of the dead trunks which were examined for insect activity
showed considerable working of Cerambycid beetles, Semanotus ligneus
am plus, to a depth of 3 or 4 inches in the xylem tissue. These activities
occurred after the death of the trees as far as could be ascertained.

These unique interrelationships are probably due in part to the
usually wet soil habitat and lack of over story vegetation in which
these trees are now growing. Such conditions are conducive both to micro-
tine populations and to the growth of the fungus. Within the range of
the giant sequoias, these conditions are uncommon at this early stage of
plant succession. It is felt that an unusually high population density of
rodents developed and, under winter conditions, they resorted to the
sequoia saplings which provided the most abundant source of food. The
damage inflicted by the rodents was followed by fungus and beetle at-
tacks upon the weakened or killed trees.

Department of Biological Sciences, San Jose State College, San Jose

LINEAR MICROSPORE TETRADS IN THE
GRASS STIPA ICHU

FRANK W. GOULD

Microspores of angiosperms commonly are in tetrahedal, isobilateral,
or decussate tetrads. T-shaped tetrads have been reported for Avisto-
lochia (Samuelsson, 1914) and Butomopsis (Johri, 1936), and linear
tetrads are known to occur in Halophila of the Hydrocharitaceae (Kau-
sik and Rao, 1942) Zostera of the Zosteraceae (Rosenberg, 1901), and
some genera of the Asclepidaceae (Gager, 1902). Two or three types of
microspore tetrads have been reported for a number of genera including
the monocots Musa, Agave, and Habeneria (Maheshwari, 1950).

In the Gramineae, division of the microspore mother cells is of the
successive type. A cell plate is laid down immediately after the first
meiotic division (Maheshwari, 1950). Characteristically and with great
regularity the microspore tetrads are isobilateral. Davis (1966) noted
that grass microspore tetrads are “occasionally T-shaped or linear’ but
cited no specific references. It is to be assumed that the extensive lit-
erature citation of Davis does include reports of T-shaped and linear
grass microspore tetrads but such types certainly are rare.

The purpose of the present paper is to report the occurrence of linear
and T-shaped microspore tetrads (figs. 1, 2) in addition to the usual
isobilateral tetrads (fig. 3) in Stipa ichu (Ruiz & Pav.) Kunth and also
the occasional development of linear groups of five microspores (fig. 4)
in the same species. Linear and T-shaped microspore tetrads were ob-
served in bud material of S. ichu from two widely separated localities
in Mexico. One collection (Gould 11622, TAES) was from 20 miles east
of Mexico City and the other (Gould 11681, TAES) was from near San
Cristobal de las Casas, near the Guatemalan border. Chromosome num-
bers were determined to be 2n = 40 in both collections. Pollen mother
cell divisions in the San Cristobal material, however, were not entirely
regular and the count was reported as ca. 2n = 40 (Gould, 1966). Of
possible significance was the collection of Stipa virescens H. B. K. at the
same locality with a chromosome number of 2n — 60 and very irregular
meiotic divisions.

About 70% of the microspore tetrads of the San Cristobal Stipa ichu
plant were linear and the remainder were T-shaped and isobilateral in
about equal proportions. A few linear groups of five microspores (fig. 4)
were also present. The percentage of linear tetrads in the Mexico City
collection was somewhat less than in the San Cristobal material but
linear and T-shaped tetrads were numerous.

Maheshwari (1950) noted that the formation of groups of more than
four microspores usually results from lagging chromosomes which organ-
ize into micronuclei. He further stated that in general such abnormal-

411

412 MADRONO [Vol. 20

35.

Fics. 1-4. Microspores of Stipa ichu: 1, linear tetrad; 2 linear tetrad and
T-shaped tetrad; 3, isobilateral tetrad; 4, linear group of five microspores. Magnifi-
cations not uniform.

ities are found only in hybrids characterized by a high degree of steril-
ity. As can be observed in Fig. 4, the five spores of the Stipa ichu “pen-
tad” are fairly uniform in size and all but the apical cell have well-
developed, apparently functional nuclei. It is possible that the fifth cell
has been organized about a micronucleus, but in that case the terminal
position of the extra cell is difficult to explain.
This paper is technical article no. TA 7470, Texas Agricultural Experi-
ment Station.
Department of Range Science, Texas A & M University

1970] GOULD: STIPA 413

LITERATURE CITED

Davis, G. L. 1966. Systematic embryology of the angiosperms. Wiley, New York.

Jouri, B. M. 1936. The life history of Butomopsis lanceolata Kunth. Proc. Indian
Acad. Sci. 5:139-162.

Kausik, S. B., and P. V. K. Rao. 1942. The male gametophyte of Halophila ovata
Gaudich. Half-yearly J. Mysore Univ. 3:43-49,

Gacer, C. S. 1902. The development of the pollinium and sperm cells in Asclepias
cornuti Decne. Ann. Bot. (London) 16:123-148.

Goutp, F. W. 1966. Chromosome numbers of some Mexican grasses. Canad. J. Bot.
44:1683-1696.

Maneswuarl, P. 1950. An introduction to the embryology of angiosperms. McGraw-
Hill, New York.

RosENBERG, O. 1901. Uber die Embryologie von Zostera marina L. Hih. Kongl.
Svenska Vetensk.-Akad. Handl. 28:1-24.

SAMUELSSON, G. 1914. Uber die Pollenentwicklung von Anona and Aristolochia und
hire systematische Bedeutung. Svensk Bot. Tidskr. 8:181-189.

EXTENSION OF THE RANGE OF ABIES LASIOCARPA
INTO CALIFORNIA

J. O. Sawyer, D. A. THoRNBURGH, and W. F. BowMAN

Abies lasiocarpa (Hook.) Nutt. has been located in two drainages
near Russian Peak in the Salmon Mountains of Siskiyou Co. During the
summer of 1968, while making a vegetational reconnaissance of this
area, A. lasiocarpa was first located in forests surrounding the meadows
west of Little Duck Lake (Etna quadrangle, T. 40N. R. 9W. Sec. 19,
elev. 6400 ft). Here it forms a forest with Tsuga mertensiana (Bong.)
Carr., Pinus monticola Dougl., and Abies magnifica A. Murr. var. shas-
tensis Lemmon. The trees are healthy; reproduction is plentiful with
some advancing into the wet meadows.

Later a more extensive forest was found along South Sugar Creek
(T. 40N. R. 9W. Sec. 30, 31). In this area the trees not only occur
around the wet meadows near South Sugar Lake at 6800 ft, but descend
along the creek terraces to 5800 ft. Below 6400 ft A. dastocarpa is mixed
with Picea engelmanni Parry ex Engelm. Both species are vigorous, and
are reproducing well.

It is surprising that this species has not been reported previously.
Munz (1959) reports P. engelmanni along Sugar Creek, tributary of
the Scott River. South Sugar Creek is a branch of this stream. Only a
rough and apparently recent trail has been constructed by fishermen to
South Sugar Lake, so accessibility is recent. An established Forest Serv-

414 MADRONO [Vol. 20

ice trail parallels Sugar Creek through stands of P. engelmanni. This
might explain the oversight.

Some older literature incorrectly reports the occurrence of A. lasio-
carpa in the “Salmon Mountains.” The references are traceable to speci-
mens of Abies amabilis (Dougl.) Forbes collected by Gillespie in 1928
near Hancock Lake in the Marble Mountain Wilderness Area (Gilles-
pie, 1931). Haddock (1961) discusses the history of the confusion
noting that A. amabilis was first correctly recognized in that location in
1932 by Crebbin, a forester on the staff of the Klamath National For-
est.

These new records do not greatly extend the range of A. lasiocarpa.
Fowells (1965) follows earlier general references in showing the south-
ern range of A. lasiocarpa to be in the Cascades in the vicinity of Crater
Lake. The first discovery of A. lasiocarpa in the Klamath Region was
made by Dennis (1959) on the slopes of Mt. Ashland, Jackson Co., thus
extending the range further south. He found one small group, possibly
a single clone due to layering. Our findings continue the range about 50
miles south of the Mt. Ashland location, and into a more central part
of the Klamath Region.

Its presence, here, accentuates the region’s central refugial nature as
discussed by Whittaker (1960; 1961). What is more noteworthy is that
A. lasiocarpa and P. engelmannii are only two of 17 conifers found above
5000 ft in the Sugar Creek drainage. These are as follows: Abies con-
color (Gord. & Glend.) Lindl., A. lastocarpa (Hook.) Nutt., A. mag-
nifica A.Murr. var. shastensis Lemmon, Juniperus communis L. var.
saxatilis Pall., Libocedrus decurrens Torr., P. breweriana Wats., P.
engelmannii Parry ex Engelm., Pinus albicaulis Engelm., P. balfouriana
Grev. & Balf., P. contorta Dougl. var. murrayana (Grev. & Balf.)
Engelm., P. jeffreyi Grev. & Balf., P. lambertiana Dougl., P. monticola
Dougl., P. ponderosa Dougl. ex P. & C. Lawson, Pseudotsuga menziesu
(Mirb.) Franco, Taxus brevifolia Nutt., and Tsuga mertensiana (Bong)
Carr.

A notable species is Pinus balfouriana Grev. & Balf. on a ridge over-
looking Little Duck Lake and Sugar Lake stands of A. lasiocarpa and
P. engelmannii. Possibly nowhere else in California is such a complete
representation of northwestern and Serrian conifers present in a single
square mile. We also doubt that any region can match the number of
conifers found in one small area.

Initial studies have begun in this area on the other plant groups. The
flowering plants are now incompletely collected due to time limitations.
The collected material, though, shows a similar pattern of mixing of
northwestern and Serrain floras. For example, the northwestern Phyllo-
doce empetriformis (Sm.) D. Don is present rather than Phvllodoce
breweri (Gray) Hell. Other northwestern shrubs which are rather com-
mon in the area include Vaccinium membranaceum Dougl., Vaccinium

1970] REVIEWS 415

scoparium Leib., and the sub-shrub Leutkea pectinata (Pursh) Kuntze.
Mixed with the Vaccinium is the Serrian Leucothoe davisiae Torr. The
rare to California Gaultheria humifusa (Graham) Rydb. was found near
the Little Duck Lake stand of A. lastocarpa. Interesting herbs include the
“uncommon” Mitella pentandra Hook. and Cypripedium fasciculatum
Kell. A complete study of the vascular plant flora of this area is planned
by us for the 1969 field season.

Specimens of A. dastocarpa are in the following herbaria: HSC, JEPS,
and the Klamath National Forest Herbarium, Yreka.

This study is in cooperation wtih and partially financed by the
Pacific Southwest Forest and Range Experiment Station, Berkeley.

Departments of Botany and Forestry, Humboldt State College, Arcata, California

LITERATURE CITED

FoweE tts, H. A. 1965. Silvics of forest trees of the United States. U.S.D.A. Agric.
Handb. 271.

GittesPiz, D. K. 1931. Records of plants new to California. Madronio 2:35-36.

Happock, P. G. 1961. New data on distribution of some true firs on the Pacific
Coast. Forest Sci. 7:349-351.

Dennis, L. R. 1959. A taxonomic study of the vascular flora of Ashland Peak,
Jackson County, Oregon. M.A. thesis, Oregon State Univ., Corvallis.

Muowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and Califor-
nia. Ecol. Monogr. 30:279 338.

. 1961. Vegetation history of the Pacific Coast States and the “central”

significance of the Klamath Region. Madrono 16:5-23.

REVIEWS

The Plant Hunters. By TyLerR WHITTLE. xii + 283 pp., illustrated. Chilton
Book Co., 401 Walnut St., Philadelphia, Penn. 19106. 1970. $8.95.

The subtitle of this book sums up its contents very well: “being an examination
of collecting with an account of the careers & the methods of a number of those
who have searched the world for wild plants.” Starting with Queen Hatshepsut
and continuing into the 20th Century the author of this very interesting and emi-
nently readable book has detailed the story of the men and women who went
around the world looking for plants, sometimes for medical reasons, sometimes in
search of ornamentals, sometimes to find spices, and sometimes to advance the
knowledge of the kinds of plants. The whole world is covered, hence no one
geographical area has received exhaustive treatment: nor in a book of this kind
would one want it.

The vistas and unspoiled plant communities of the past are often no longer
with us. It makes one rather depressed to realize what man has done and is con-
tinuing to do to his surroundings, especially in decreasing the diversity of living
things. Perhaps one of the great services of this book, especially to young readers,
will be to install in them a sense of curiosity about and interest in plants, for after-
all, there is little motivation to preserve that about which one knows nothing or in
which one has no interest——Joun H. THomas, Department of Biological Sciences,
Stanford University.

416 MADRONO [Vol. 20

Plant Variation and Evolution. By D. Briccs and S. M. WALTERS. 256 pp.,
illustrated. McGraw-Hill Book Company, New York, 1969. $2.45

This book is an introduction to plant biosystematics. The initial chapters sketch
briefly the historical context in which modern plant systematics developed. The
remainder and major portion of the book is written essentially as a discussion of
speciation and includes the following topis: changes in population, breeding sys-
tems, hybridization, polyploidy, and patterns of evolution. The many well illus-
trated examples which accompany the text are drawn primarily from studies of the
European Flora.

Many sections within the various chapters are exceptionally well written. Of
particular note is the treatment of apomixis and hybridization. Although consid-
erable space is devoted to the factors involved in species formation, the discussion
of the “species problem” is most unsatisfactory. Rather than questioning if a
single species definition is possible or desirable the authors state that “the ‘ideal’
‘biological’, or ‘evolutionary species’ of the experamentalist is the hologamodeme,
defined as composed of individuals which ‘are believed to interbreed with a high
level of freedom under a specified set of conditions, and separated from other
hologamodemes by at least partial sterility’”. Such a definition is clearly non-
operational for most plant groups and therefore cannot be used as an absolute
guideline by plant systematists.

The recently published Plant Biosystematics by Otto T. Solbrig is the only book
comparable in its scope to Plant Variation and Evolution. However, in contrast,
the Solbrig book appears to offer a more complete and accurate treatment of many
topics.

Plant Variation and Evolution should best be regarded as a reference work for
students——DeEnnis R. PARNELL, Department of Biological Science, California State
College, Hayward, California.

NOTES AND NEWS

RIBES MALVACEUM IN THE FOOTHILLS OF CALAVERAS COUNTY, CALIFORNIA.—
In March, 1967, I observed a small population of Ribes malvaceum Sm. in the
foothills of the Sierra Nevada near Valley Springs along the access road to the
south arm of New Hogan Reservoir. Subsequent search revealed a considerable
number of stations scattered a few miles apart in the chaparral at elevations ranging
from 450 to 1000 feet. One large group of about 200 shrubs is located at the junc-
tion of the Southern Pacific Railroad with State Highway 12, 2 miles west of
Valley Springs (Taylor 11, CAS, Fresno State College Herbarium, GH, MO). The
plants at this locality are well spaced on a northwest exposure in association with a
number of chaparral species including Quercus wislizenii, Arctostaphylos mariposa,
Baccharis pilularis var. consanguinea, Adenostoma fasciculatum, and Fremontoden-
dron californicum. Other localities in the area include the spillway at New Hogan
Reservoir and along the south arm of the Reservoir (Taylor 12, Fresno State College
Herbarium). All localities show uniformity as to slope, soil, and composition of the
vegetation. This population extends the range southward in the Sierra Nevada
from E] Dorado Co.—DEAN WILLIAM Taytor, Biology Department, Fresno State
College, Fresno, California.

1970]

INDEX 417

INDEX TO VOLUME XX

Classified entries: authors; book reviews; botanical names (excluding taxa in
floristic lists) ; major subject headings; new combinations and taxa (in boldface) ;

and titles.

Abbott, I. A.: Some new species, new
combinations, and new records of red
algae from the Pacific Coast, 42

Abies lasiocarpa into California, Exten-
sion of the range of, 413; concolor,
13; c. lowiana, 13; grandis, 15, 68;
lasiocarpa, 16; lowiana, 13; sonomen-
sis, 23

Acalypha adamsii, 263; albermarlensis,
262; baurrii, 261; chatamensis, 263;
cordifolia, 262; diffusa, 262; flaccida,
261; parvula chathamensis, 263; p.
cordifolia, 262; p. flaccida, 261; p. p.,
261; p. procumbens, 263; p. pubescens,
260; p. reniformis, 263; p. strobilifera,
263; p. velutina, 261; reniformis, 263;
sericea baurrii, 261; s. indefessus,
261; s. s., 260; spicata, 262; strobili-
fera, 263; velutina, 261; v. minor,
261; wigginsii, 261

Acanthospermum lecocarpoides, 250; lep-
tolobum, 250

Achyranthes glaucescens, 264; nudicau-
lis, 264; strictiuscula, 264

Agrostis perennans (Poaceae), a remote
disjunction in the Pacific Northwest,
220

Aletes acaulis, 215; filifolius, 214

Alteranthera filifolia glaucescens, 214;
f. microcephala, 264; f. nudicaulis,
264; f. pintensis, 264; f. rabidensis,
265; glauscenscens, 264; g. strictiuscu-
la, 264; nudicaulis, 264

Anderson, D.: Review, Grass Systemat-
ics, 288

Anderson, E. F., and D. L. Walkington:
New combinations and taxa in the
Cactaceae of the Galapagos Islands,
256

Anderson, L. C.: Embryology of Chry-
sothamnus (Astereae, Compositae),
337

Anredera ramosa, 266

Arctostaphylos columbiana, 63;
mularia, 63

Arcyria versicolor, 280

Arenaria pusilla, 399

Artemisia, An ecological contribution to
the taxonomy of, 385; arbuscula, 385;
longiloba, 385; nova, 385

Astragalus (Fabaceae) from Nevada, A
new, 395; Astragalus lentiginosus and
Tridens pulchellus in relation to rain-
fall, Perennation in, 326; coccineus,
398; funereus, 398; lentiginosus fre-
montii, 326; |. variabilis, 329; new-

num-

berryi, 395; phoenix, 395; purshii,
395; p. tinctus, 398; tidestromii, 329
Atlas, Proposed California tree, 236

Banwar, S. C.: Fossil leaves of Lyono-
thamnus, 359

Barbeyella minutissima, 75, 280

Barbour, M. G.: The flora and plant
communities of Bodega Head, Califor-
nia, 289

Barneby, R. C.: A new Astragalus (Fa-
baceae) from Nevada, 395; and E. C.
Twisselmann: Notes on _ Loeflingia
(Caryophyllaceae), 398

Beattle, A. A.: An ecological contribu-
tion to the taxonomy of Artemisia,
385

Beattley, J. C.: Perennation in Astragal-
us lentiginosus and Tridens pulchellus
in relation to rainfall, 326

Becking, W. P., Fasciation of coastal
redwoods, 382

Bemis, W. P., and T. W. Whitacker:
The xerophytic cucurbita of north-
western Mexico and _ southwestern
United States, 33

Bodega Head, California, The flora and
plant communities of, 289

Bonar, L.: Harold Ernest Parks, 373

Boussingaultia baselloides, 266; ramosa,
266

Bryophytes and vascular plants, Phyto-
geography of northwestern North
America, 155

Bucholtzia glaucescens, 264; nudicaulis,
264

Burch, D.: A new combination in Cha-
maesyce from the Galapagos Islands,
209

Cactaceae of the Galapagos Islands, New
combinations and taxa in the, 256

Calonema (Myxomycetes), A new co-
phrophilous species of, 299; aureum,
229, luteolum, 229

Calylophus (Onagreaceae), A new spe-
cies and some new combinations in,
241; australis, 243; drummondianus,
243; d. berlandieri, 243; hartwegii,
243; h. fendleri, 243; h. filifolius,
243; h. lavandulifolius, 243; h. mac-
cartii, 243; h. pubescens, 243; h.
toumeyi, 243; serrulatus, 243; tubicu-
la, 243

Campanula from northern California, A
new, 231; piperi, 234; scabrella, 231;

418 MADRONO

shetleri, 231; wilkinsinana, 234

Carlbom, C. G.: Agrostis perennans (Po-
aceae), a remote disjunction in the
Pacific Northwest, 220

Castanopsis chrysophylla, 68

Castilleja from the southern Sierra Ne-
vada, A new species of, 209; cinerea,
213; culbertsonii, 213; pilosa, 213;
praeterita, 209

Centrospermae of the Galapagos Islands,
Nomenclatural changes and new sub-
species in the, 264

Ceratiomyxa fructiculosa, 279

Cereus sclerocarpus, 256

Chaetospira funckii, 255

Chamaecyparis nootkatensis, 9

Chamaesyce from the Galapagos Islands,
A new combination in, 253; nummu-
laria glabra, 253

Chambers, K. L.: Review, Flora of
Alaska and neighboring territories, 78

Chondrus ocellatus, 51

Chremosome numbers in some North
American species of the genus Cirsium.
II., 225; Chromosome studies in Mel-
ampodium (Compositae, Heliantheae),
365; Chromosome numbers and a
proposal for classification in Sisyrin-
chium (Iridaceae), 269

Chrysothamnus (Astereae, Compositae),
Embryology of, 337; albidus, 340;
greenei, 340; linifolius, 337; nauseosus
albicaulis, 337; paniculatus, 341; par-
ryi, 341; teretifolius, 341; vicidiflorus
humilis, 339; v. lanceolatus, 339; v.
puberulus, 339

Cirsium. II. Western United States,
Chromosome numbers in some North
American species of the genus, 225;
acanthodontum, 226; arizonicum, 228;
brevistylum, 226; californicum, 227;
canescens, 227; coloradense, 226;
foliolosum, 227; occidentale, 227; och-
recentrum, 227; pastoris, 228; ryd-
bergii, 226; scariosum, 227; scopulo-
rum, 226; subniveum, 227; tiogonum,
227; tweedyi, 226; undulatum, 227;
utahense, 227; wallowense, 227

Clarkina jolonensis (Onagraceae), a
new species from the Inner Coast
Range of California, 321; deflexa, 321

Classification Society, The, 236

Claytonia nevadensis from northwestern
California, New distributional record
fors31

Comatricha elegans, 379; fusiforme, 280;
lurida, 379; nigra, 280; pacifica, 280;
suksdorfii, 280; typhoides, 281

Compositae of the Galapagos Islands,
New combinations in the, 255

Comptonia parvifolia, 361; peregrina,
362

[Vol. 20

Constance, L.: Review, The evolution
and classification of flowering plants,
77

Coolidge, J.: A new species of Polygo-
num (Polygonaceae), 266

Cribraria argillacea, 279; microcarpa,
378; minutissima, 378; rufa, 279

Critchfield, W. B., and G. A. Allenbaugh:
The distribution of Pinaceae in and
near northern Nevada, 12

Croton albescens, 258; brevifolius, 259;
incanus, 258; macraei, 258; rivinifo-
lius, 257; scouleri brevifolius, 259; s.
castelllanus, 258; s. darwinii, 259; s.
glabriusculus, 259; s. grandifolius,
259; s. macrael, 258

Cronquist, A.: New combinations in the
Compositae of the Galapagos Islands,
255

Cryptopleura brevis, 45; dichotoma, 45;
lobulifera, 45; rosacae, 45

Cucurbita of northwestern Mexico and
southwestern United States, The xero-
phytic, 33; cordata, 33; cylindrata,
33; digitata, 33; foetidissima, 33;
moschata, 36; palmata, 33; pedatifolia,
33

Cupressus pygmaea, 60

Curtis, D. H.: New records of Myxomy-
cetes from Oregon. I., 75; A prelim-
inary report of the Myxomycetes of
Crater Lake National Park, Oregon,
378

Cyperaceae of the Galapagos Islands,
New combinations in the, 253

Cyperus drummondii, 254; gatesii, 253;
inconspicuous, 253; liebmanii, 257; mi-
crodontus, 253; m. texensis, 253; poly-
stachyos fugax, 254; p. holosericeus,
253; p. inconspicuous, 253; p. laxiflor-
us, 253; p. leptostachyus, 253; p. tex-
ensis, 254; surinamensis, 254; texensis
253, drummondii, 254

Daubenmire, R.: Ecological plant geog-
raphy of the Pacific Northwest, 111
Dicentra (Fumariaceae), Pollen aperture
variation and phylogeny in, 354; subg.
Hedycapnos, 356; subg. Macranthos,
358; burmanica, 355; canadensis, 355;
chrysantha, 355; cucullaria, 355;
eximia, 355; formosa, 355; grandi-
foliolata, 355; lichiagensis, 355; ma-
crantha, 355; macrocapnos, 355; ne-
vadensis, 355; ochroleuca, 355; pauci-
flora, 355; paucinervia, 355; peregrina,
356; roylei, 356; scandens, 356; spec-
tabilis, 356; torulosa, 356; uniflora,
356

Dichomena ciliata, 254

Diderma deplanatum 281; nigrum, 281;
niveum, 281; radiatum, 381; subcae-

1970]

ruleum, 281; umbilicatum, 377, 381
Dianema andersonii, 76, 279; corticatum,
279; deplanatum 76; nigrum, 76
Darwiniothamnus lancifolius, 255; tenui-
folius glabriusculus, 255; t. glandulo-
sus, 255
Echinostelium cibraroides, 379; minu-
tum, 379
Eliason, U.: Nomenclatural changes and
new subspecies in the Centrospermae
of the Galapagos Islands, 264
Embryology of Chrysothamnus
tereae, Compositae), 337
Enerthenema melanospermum, 281
Enteridium olivaceum, 279
Erigeron lancifolius glabriusculus, 255;
tenuifolium lancifolius, 255
Eriogonum apricum (Polygonaceae), A
new prostrate variety of, 320; a. pro-
stratum, 320
Erioxylum aridum, 346, palmeri, 347
Ernst, W. R.: Review, Rocky Mountain
flora, 29; Review, Flora Europaea,
Vol. 2, 237; and M. F. Baad: Two
new species of Lamourouxia (Scro-
phulariaceae) in Mexico, 342
Euphorbiaceae, Notes on Galapagos, 257

(As-

Fasciation of coastal redwoods, 382

Ferlatte, W. J.: New distributional rec-
ord for Claytonia nevadensis from
northwestern California, 31

Ferris, R. S.: Laura M. Lorraine, 1904—
1968, 26

Flora and plant communities of Bodega
Head, California, The, 289

Floristic and vegetational history of the
Pacific Northwest, Neogene, 83

Forest-podsol ecosystem and its dune
association of the Mendocino Coast,
The pygmy, 60

Froelichia lanata, 265; lanigera, 265; 1.
scoparia, 265; nudicaulis lanigera,
265; spocaria, 265

Fryxell, P. A.: Notes on some Mexican
species of Gossypium (Malvaceae),
347

Fuligo septica, 281

Galapagos Islands. I. New species and
combinations, Studies on plants of the,
250

Galium ferrugineum, 250; galapagoense,
Z50

Galpinsia toumeyi, 243

Gaultheria shallon, 63

Gigartina, 52

Gilbertson, R. L.: A new Vararaia from
western North America, 282

Gossypium (Malvaceae), Notes on some
Mexican species of, 347; aridum, 347;

INDEX 419

gossypioides, 348; trilo-
bum, 348

Gould, F. W.: Linear microspore tetrads
in the grass Stipa ichu, 441

Griffin, F. R.: Proposed California tree
atlas, 236

Grindelia, 340

Gutierrezia, 340

rosel, 347;

Haplopappus macronema, 341; propin-
quus, 341, trianthus, 341

Hartmannia domingensis, 248

Heckard, L. R.: A new Campanula from
northern California, 231; and R. Baci-
galupi: A new species of Castilleja
from the southern Sierra Nevada, 209

Hemitrichia clavata, 76; karstenii, 280;
montana, 76, 280

Heterosiphonia asymmetria and H. den-
siuscula and their life histories in cul-
ture, The conspecificity of, 313; erec-
ta, 313; laxa, 313; wurdenmannii, 313

Hevly, R. H.: A new species of Probos-
cidea (Martiniaceae) from Baja Cali-
fornia, 392

Hickman, J. C., and M. P. Johnson: An
analysis of geographical variation in
western North American Menziesia
(Ericaceae), 1

Hitchcock, C. L., A tribute from the
California Botanical Society, 387

Hymenena setchellii, 45

Hyphloma fasciculare, 410

Iridaea, 52
Iris pseudacorus in western North Amer-
ica, 391

delicatus,
250) le

Jasminocereus howellii var.
256; thoursii delicatus,
sclerocarpus, 256

Jenny, H., R. J. Arkley, and A. M.
Schultz: The pygmy forest-podsol
ecosystem and its dune associates of
the Mendocino coast, 60

Kowalski, D. T.: A new coprophilous
species of Calonema (Myxomycetes),
229; Concerning the validity of Lam-
proderma echinosporum, 323; and D.
H. Curtis: New records of Myxomy-
cetes from California, IV, 377

Koyama, T.: New combinations in the
Cyperaceae of the Galapagos Islands,
256

Kruckeberg, A. R.: Soil diversity and
distribution of plants, with examples
from western North America, 129; C.
Leo Hitchcock, 387

Lamourouxia (Scrophulariaceae) in
Mexico, Two new species of, 342;

420

colimae, 342; gracilis, 344; gutierrezii,
344; jaliscana, 344; lanceolata, 344;
longiflora, 346; viscosa, 346

Lamproderma echinosporum, Concern-
ing the validity of, 323; arcyrioides,
281; arcyrionema, 380; atrosporum,
325; biasperosporum, 281, 380; cares-
tiae, 281; echinulatum, 323; gulielmae,
325; sauteri, 281, 325

Lang, F. A.: A new name for a species
of Polypodium from northwestern
North America, 53

Ledum glandulosum, 63

Lepidoderma chailletii, 281

Leptochloa albermarlensis, 253

Leucocarpus lecocarpoides, 256; lepto-
lobus, 256

Licea kleitobolus, 378; minima, 279;
parasitica, 377; pusilla, 75, 279

Lindbladia effusa, 279

Lithocarpus densiflorus, A mutant of,
221; d. attenuato-dentatus, 224

Loeflingia (Caryophyllaceae), Notes on,
398; hispanica, 398; pusilla, 399;
squarrosa, 399; s. artemisiarum, 406;
s. cactorum, 407; s. texana, 407; tex-
ana, 399; verna, 399

Lorraine, 1904-1968, Laura M., 26

Lundh, J.: A new variety of Opuntia
megasperma from the Galapagos Is-
lands, 254

Lycogala epidendrum, 279; flavofuscum,
75, 279

Lyonothamnoxylon nevadensis, 361

Lyonothamnus, Fossil, leaves of, 359;
floribundus asplenifolius, 359; f. f.,
361; mohavensis, 361; parvifolia, 361

Madrono, 213, 391; back issues of, 383;
editorship of, 364; special issue of, for
the XI International Botanical Con-
gress, 81

Mathias, M. E., L. Constance, and W.
L. Theobald: Two new species of Um-
belliferae from southwestern United
States, 214

Melampodium (Compositae, Helian-
theae), Chromosome studies in, 365;
americanum, 369; aureum, 369; cine-
reum, 369; cupulatum, 369; dicoelo-
carpum, 369; diffusum, 370; divarica-
tum, 370; glabrum, 370; gracile, 370;
hispidum, 370; leucanthum, 370; lin-
earilobum, 371; longifolium, 371; lon-
gipes, 371; longipilum, 371; micro-
cephalum, 371; montanum, 371;
paniculatum, 371; perfoliatum, 371;
rosel, 372; sericeum, 372; tenellum,
B12

Menziesia (Ericaceae), An analysis of
geographic variation in western North
America, 1; ferruginea, 1; glabella, 1;

MADRONO

[Vol. 20

g. ferruginea, 1

Minuartia, 402

Mollugo flavescens angustifolia, 265; f.
gracillima, 265; f. insularis, 265; f.
intermedia, 265; f. striata, 265; flori-
ana santacruziana, 265; gracillima
latifolia, 265; gracilis, 265; insularis,
265; snodgrassii santacruziana, 265;
striata, 265

Mosquin, T.: Chromosome number and
a proposal for classification in Sisy-
rinchium (Iridaceae) , 269

Myatt, R.: A new prostrate variety of
Eriogonum apricum (Polygonaceae),
320

Myrica californica, 68

Myriogramme, hollenbergii, 42

Myxymycetes from California. IV., New
records of, 377; A new coprophilous
species of Calonema, 229; of Crater
Lake National Park, Oregon, A pre-
liminary report of the, 278; from Ore-
gon. I., New records of, 75

Nemophila, 276

New publications, 32, 336

Nitrophyllum, cincinnatum, 43; dotyi,
44; hollenbergii, 42; mirabile, 43;
northii, 43

Notes and news, 31, 213, 236, 336, 349,
364, 382, 391, 416

Oenothera brandegeei from Baja Califor-
nia, Mexico, and a revision of subge-
nus Pachylophus (Onagraceae), 350;
Oenothera subg. Hartmannia (Ona-
graceae), Two new species and some
nomenclatural changes in, 246; albi-
caulis, 353; brandegeei, 352; caespi-
tosa, 350; c. brandegeii, 352; cavernae,
351; coronopifolia, 353; delessertiana,
249; domingensis, 248; epilobiifolia
cuprea, 248; e. e., 248; fendleri, 234;
greggii pubescens, 234; hartweggii,
234; kunthiana, 248; lavandulaefolia,
234; macrosceles, 352; muelleri, 352;
multicaulis, 248; platanorum, 246;
primiveris, 350; rosea, 246; speciosa,
249; s. childsii, 249; tarquensis, 248;
texensis, 247; tubicula filifolia, 234;
tubifera, 352; xylocarpa, 352

Oligonema flavidum, 229; schweinitzii,
280

Opuntia megasperma from the Galapa-
gos Islands, A new variety of, 254;
echios zacana, 256; galapagela zacana,
256; g. profusa, 256; megasperma
mesophytica, 254

Ornduff, R.: Review, Handbook of
Northwestern plants, 31; Review, Vas-
cular plants of the Pacific Northwest,
Part 1, 238; Review, Plants of Oregon

1970]

coastal dunes, 287

Ownbey, G. B., and Y. Hsi: Chromo-
some numbers in some North Ameri-
can species of the genus Cirsium. II.
Western United States, 225

Oxalis laxa in California, Records and
observations on a rare plant, 349

Ozophora californica, 48; clevelandii,
48; latifolia, 49; norrissii, 50

Paratimia conicola, 63

Parks, Harold Ernest, 373

Parnell, D. R.: Review, Plant taxonomy,
236; Clarkia jolonensis (Onagraceae),
a new species from the Inner Coast
ranges of California, 321; Review,
Modern methods in plant taxonomy,
335; Review, Nightshades, the para-
doxical plants, 384; Review, Plant
variation and evolution, 416

Passiflora colinvauxii, 251

Peponapis, 37

Perichaena, 230

Petradoria, 341

Petroglossum pacificum, 51

Philbrick, R. N.: Review, The native
cacti of California, 333

Pholistoma (Hydrophyllaceae), Com-
parative natural history of two sym-
patric populations of, 276; auritum,
276; a. arizonicum, 271; membranace-
um, 276; racemosum 276

Phyllophora californica, 48; clevelandii,
48; submaritimus, 48

Physarum albescens, 281; auripigmen-
tum, 76, 281; decipiens, 281; leucopus,
380; luteolum, 380

Phytogeography of northwestern North
America: Bryophytes and_ vascular
plants, 155

Picea engelmannii, 23, 413; lahontense,
ae

Pinaceae in and near northern Nevada,
The distribution of, 12; albicaulis, 18;
aristata, 21; attenuata, 63; balfouri-
ana, 414; contorta bolanderi, 60; c.
latifolia, 21; c. murrayana, 21; crosii,
23; engelmannii, 16; flexilis, 20; har-
neyana, 23; jeffreyi, 22; monophylla,
21; monticola, 18; muricata, 60; pon-
derosa, 22; p. scopulorum, 22; washo-
ensis, 22

Plantago paralias pumila, 252; tomento-
Sa.02572

Plant geography of the Pacific North-
west, Ecological, 111

Pollen aperture variation and phylogeny
in Dicentra (Fumariaceae), 354

Polygonum (Polygonaceae), A new spe-
cies of, 266; cascadense, 268; mini-
mum, 268; sawatchense, 266; tenue,
268; triandrous, 266

INDEX 421

Polypodium from northwestern North
America, A new name for a species of,
53; amorphum, 55; australe, 56; cam-
bricum, 56; columbianum, 56; gly-
cyrrhiza 53; hesperium, 53; montense,
57; scouleri, 54; vulgare, 53; v. co-
lumbianum, 56

Porophyllum ruderale macrocephalum,
255

Porter, D. M.: A new Tetragastris (Bur-
seraceae) from Panama, 346

Prionitis andersonil, 48

Proboscidea (Martyniaceae) from Baja
California, Mexico, A new species of,
382; altheifolia, 392; gracillima, 392;
parviflora, 392; sinaloensis, 395

Prototrichia metallica, 279

Pseudocymopterus longiradiatus, 217;
montanus, 214

Pseudolephantopus funckii, 255; spiralis,
250

Pseudotsuga menziesii, 17, 60; m. glauca,
17; m. m., 17; sonomensis, 23

Pteroglossum, 47

Raven, P. A.: Review, Supplement to a
California flora, 239; Oenothera bran-
degii from Baja California, Mexico,
and a review of subgenus Pachylophus
(Onagraceae), 350; Review, Marin
flora, 383; and D. R. Parnell: Two
new species and some nomenclatural
changes in Oenothera subg. Hartman-
nia (Onagraceae), 246; and J. H.
Thomas: Iris pseudacorus in western
North America, 390

Red algae from the Pacific Coast, Some
new species, new combinations, and
new records of, 42

Redwoods, Fasciation of coastal, 382

Reeder, J. R., and C. G. Reeder: A new
combination in Trichoneura from the
Galapagos Islands, 253

Reviews: Ehrlich, P. R., The population
bomb, 28; Weber, W. A., Rocky
Mountain flora, 29; Porter, C. L., Tax-
onomy of flowering plants, 30; Gilkey,
H. M., and L. R. M. Dennis, Hand-
book of northwestern plants, 31;
Cronquist, A., The evolution and
classification of flowering plants, 77;
Hulten, E., Flora of Alaska and neigh-
boring territories, 78; Tutin, T. G.,
et al., Flora Europaea, Vol. 2, 237;
Heywood, V. H., Plant taxonomy, 237;
Hitchcock, C. L., et al., Vascular
plants of the Pacific Northwest, Part
1, 238; Munz, P. A., Supplement to a
California flora, 239; Wiedemann, A.
M., L. R. J. Dennis, and F. H. Smith,
Plants of the Oregon coastal dunes,
287; Gould, F. W., Grass systematics,

422 MADRONO

288; Solbrig, O. T., Principles and
methods of plant biosystematics, 332;
Benson, L., The native cacti of Cali-
fornia, 333; Heywood, V. H., Modern
methods in plant taxonomy, 335;
Taylor, T. M. C., Pacific northwest
ferns and their allies, 335; Howell, J.
T., Marin flora, 383; Heiser, C. B.,
Nightshades, the paradoxical plants,
384; Ferris, R. S., Flowers of the Point
Reyes National Sea Shore, 384; Whit-
tle, T., The plant hunters, 415; Briggs,
D., and S. M. Walters, Plant variation
and evolution, 416

Rhododendron macrophyllum, 63

Rhodoglossum, 52

Rhodymenia pacifica, 49

Rhynchospora ciliata, 254; nervosa cili-
ata, 254

Ribes malvaceum in the foothills of
Calaveras County, California, 416

Sawyer, J. O., D. A. Thornburgh, and
W.F. Bowman: Extension of the range
of Abies lasiocarpa into California, 413

Shofield, W. B.: Phytogeography of
northwestern North America: bryo-
phytes and vascular plants, 155

Scytinostroma praestans, 284

Searcy, K. B.: Comparative natural his-
tory of two sympatric populations of
Pholistoma (Hydrophyllaceae) , 276

Selera gossypioides, 348

Sequoia sempervirens, 60, 382

Sequoias, Unusual factors contributing
to the destruction of young giant, 408

Sequoiadendron giganteum, 408

Shellhammer, H. S., R. E. Stecher, H. T.
Harvey, and R. J. Hartesveldt: Un-
usual factors contributing to the de-
struction of young giant sequoias, 408

Sicyocaulis pentagonus, 252

Sisyrinchium (Iridaceae), Chromosome
numbers and a proposal for classifica-
tion in, 269; albidum, 272; atlanticum,
272; angustifolium, 271; arenicola,
272; bellum, 271; bermudiana, 271;
capillare, 272; campestre, 272; grami-
noides, 272; halophilum, 272; langloi-
sil, 272; littorale, 272; montanum,
271; mucronatum, 272; sagittiferum,
272; sarmentosum, 272

Stemonitis, splendens, 379; webberi, 379

Stern, K. R.: Review, Taxonomy of
flowering plants, 30; Pollen aperture
variation and phylogeny in Dicentra
(Fumariaceae), 354

Stipa ichu, Linear microspore tetrads in
the grass, 411; virescens, 411

Strother, J. L.: Review, Principles and
methods of plant biosystematics, 332

Stuessy, T. F.: Chromosome studies in

[Vol. 20

Melampodium (Compositae, Helian-
theae), 365

Smith, A. R.: Review, Pacific northwest
ferns and their allies, 335

Soil diversity and the distribution of
plants, with examples from western
North America, 129

Solidago, 341

Spirochaeta funckii, 255

Tandonia ramosa, 266

Taylor, D. W.: Ribes malvaceum in the
foothills of Calaveras County, Cali-
fornia, 416

Telanthera angustata, 264; glaucescens,
264; nudicaulis, 264

Tetragastris (Burseraceae) from Pana-
ma, A new, 346; panamensis, 346;
tomentosa, 346

Thomas, J. H.: Review, Flowers of the
Point Reyes National Seashore, 384;
Review, The plant hunters, 415

Towner, H. F., and P. H. Raven: A
new species and some new combina-
tions in Calylophus (Onagraceae), 241

Tree atlas, Proposed California 236

Trichia affinis, 76, 280; contorta, 280;
favoginea, 280; lutescens, 280; pusilla,
280; varia, 280

Trichoneura from the Galapagcs Islands,
A new combination in, 253; lindley-
ana albermalensis, 253

Tridens pulchellus in relation to rainfall,
Perennation in Astragalus lentigino-
sus and, 326

Tsuga heterophylla, 68

Tubifera ferruginosa, 378

Tucker, J. M., W. E. Sundahl, and D.
O. Hale: A mutant of Lithocarpus
densiflorus, 221

Umbelliferae from the southwestern
United States, Two new species of,
214

Vaccinium ovatum, 63

Vallesia glabra pubescens, 252

Vararia from western North America, A
new, 282; athabascensis, 283; granu-
losa, 282; investiens, 283; ramosa, 283

Vegetational history of the Pacific
Northwest, Neogene floristic and, 83

Watt, K. F.: Review, The population
bomb, 28

Webster, G. L.: Notes on Galapagos
Euphorbiaceae, 257

Weiler, J.: Records and observations on
a rare plant, Oxalis laxa, in California,
349

West, J. A.: The conspecificity of Heter-
osiphonia asymmetrica and H. densi-

1970] INDEX 423

uscula and their life histories in cul-
buce, 383

Wiggins, I. L.: Studies on plants of the
Galapagos Islands. I. New species and
combinations, 250

Wolfe, J. A.: Neogene floristic and vege-
tational history of the Pacific North-
west, 83

Xenoglossa, 37

Zoe, 31

To you “Bill,” Witt1AM McKIn_LEy HIesey, the California Botanical
Society takes sincere pride and pleasure in dedicating Volume Twenty of
Madrono. Your sound and constructive guidance during a long and active
membership; as Business Manager, President, and Member of the Board
of Directors has set a standard that has, and will benefit the society for
years in the future.

As the plant physiologist and “‘anchor-man” on the Carnegie Institu-
tion of Washington’s Experimental Taxonomy team of Clausen, Keck
and Hiesey, your imaginative transplant studies have fused the diverse
disciplines of genetics, cytology, taxonomy and environmental distribu-
tion with basic physiology, giving a lucid picture of the genecological
nature of plant species and an insight into the processes of natural selec-
tion which have molded these entities. As a pioneer in physiological ecolo-
gy your careful, precise research has established a firm foundation which
will endure in all future studies.

Quiet and modest, your primary energy is devoted to research and
post-doctoral teaching; however, students of biology at all levels of
training throughout the world seek your council and all receive gener-
ously of your enthusiastic guidance and warm friendship. In sincere
appreciation we thank you and look forward to many future years of
your warm human guidance.

CONTENTS

Page
Frontispiece: William McKinley Hiesey . . iv
An analysis of geographical variation in Reser North Annet ee Weenie
(Ericaceae), James C. Hickman and Michael P. Johnson . . . 1
The distribution of Pinaceae in and near northern Nevada, William B. Crick
eld and Gordon L. Allenbaugh . . Se dttedl 3yston ee ere ane i
Laura M. Lorraine, 1904-1968, Rocaias Fertis Ge sd = 20
IROVAG WS 35 cee te ae ee ete ee -. «28, 77, 237, 287, 332, 383, 418
Notes and News .. . oe 31, 213, 236, 336, 349, 364, 382, 391, 416
The xerophytic Cucurbita OF nor heen Mexico and southwestern nied
States, W. P. Bemis and Thomas W. Whitaker . . . 33
Some new species, new combinations, and new records ai ail alee con the
Pacific Coast, Isabella A. Abbott . . . . 42
A new name for a species of Polypodium from Moninestemn North meric!
Frank A. Lang .. . 5S
The pygmy forest-podsol SOR aad hs ie ACEC GS of the Mendocino
Coast, H. Jenny, R.J. Arkley,and A.M. Schultz . . . . Se 9 4s o> 100
New recone of Myxomycetes from Oregon. I., Dwayne H. Cus aot tet ih)
Neogene floristic and vegetational history of the Pacific Northwest, ieee A.
Wolfe .. ef in ela BOS
Ecologic plant nari a the Dacia N orth R. Denne Are 111
Soil diversity and the distribution of plants, with examples from western North
America, A. R. Kruckeberg . . ee ar ee ee ere 2)
Phytogeography of northwestern Noth aneren: hierar & and vascular
plants, W. B. Schofield... . 155
A new species of Castilleja from the eouthers Siow Needs Paibrence R.
Heckard and Rimo Bacigalupi. . . 209
Two new species of Umbelliferae from the eoutbtgestenn United SES, M deed
E. Mathias, Lincoln Constance, and William L. Theobald . . . . 214
Agrostis perennans (Poaceae), a remote disjunction in the Pacific Norinmeet
Curt G. Carlbom .. . 220
A mutant of Lithocarpus sencionae: John M. averer Wallan E. Sata
and Dale O. Hall . .. . 27)
Chromosome numbers in some North Remain) soreaties ae the ome Gacinm
II. Western United States, Gerald B. Ownbey and Yu-tseng Hst . . , £225
A new coprophilous species of Calonema (Myxomycetes), Donald T. Kowalski 229
A new Campanula from northern California, Lawrence R. Heckard . . . . 231
A new species and some new combinations in fiat ak (Onagraceae),
Howard F. Towner and Peter H. Raven .. . 241
Two new species and some nomenclatural changes in Oonceher, are eane!
mannia (Onagraceae), Peter H. Raven and Dennis R. Parnell. . . . . 246
Studies on plants of the Galapagos Islands. I. New species and combiner!
Ira L. Wiggins . . . 250
A new combination in mMrienonedta foun ite Galparee Teens oun 'R.
Reeder and Charlotte G. Reeder . .. . 253

A new combination in Chamaesyce from the Galaparoe Tstands, Derek Burch 253
New combinations in the Cyperaceae of the Galapagos Islands, Tetsuo Koyama 253
A new variety of Opunita megasperma from the Galapagos Islands, J. Lundh 254
New combinations in the Compositae of the Galapagos Islands, Arthur

Cronquist . . . 255
New combinations Aa oa | in ihe Cae nee ot me CRIES Tetands ina

F. Anderson and David L. Walkington. . . er ee ee es See 4) )
Notes on Galapagos Euphorbiaceae, Grady L. Webster a 25
Nomenclatural changes and new subspecies in the Ganiaananine ion the Gas

lapagos Islands, Uno Eliasson . .... . . . 2. 4 ee a ws 6264

A new species of Polygonum (Polygonaceae), Jerrold Coolidge

Chromosome numbers and a proposal for classification in Sisyrinchium Grades
ceae), Theodore Mosquin . .

Comparative natural history of two aerate popultions i Pholisconra: ( Hy-
drophyllaceae), Karen B. Searcy

A preliminary report of the Vijaomnycetes” oi Gates ake Nawonal Parke
Oregon, Dwayne H. Curtis : a

A new Vararia from western North Aunties, Ieee L. Giberccn :

The flora and plant communities of Bodega Head, California, M. G. Baroour

The conspecificity of Heterosiphonia asymmetria and H. densiuscula and their
life histories in culture, John A. West

A new prostrate variety of Eriogonium apricum (eel eons Roancy Moraes

Clarkia jolonensis (Onagraceae), a new species from the inner coast ranges
of California, Dennis R. Parnell

Concerning the validity of Lamproderma Reina coenurn Donald T. Kowalski

Perennation in Astragalus lentiginosus and Tridens pulchellus in relation to
rainfall, Janice C. Beatley

Bebe ology of Chrysothamnus Gaetereaes Gamporeon over C. Anderson

Two new species of Lamourouxia (Scrophulariaceae) in Mexico, Wallace R.
Ernst and Michael F. Baad . Lae

A new Tetragastris (Burseraceae) from Bara Duncan M. Pore

Notes on some Mexican species of Gossypium (Malvaceae), Paul L. Fryxell

Oenothera brandegeei from Baja See a Mexico, and a review of subgenus
Pachylophus, Peter H. Raven .

Pollen aperture variation and phylogeny in iieeniess (Enmenceesoy ees
R. Stern.

Fossil leaves of Teouernenme Satish C. Sana

Chromosome studies in Melampodium (Compositae, Richantheaey) Tod F.
Stuessy ; :

Harold Ernest Parks, Tee Bonar .

New records of Myxomycetes from inne Ke Donne T. Neotees and
Dwayne H. Curtis .

An ecological contribution to fie (Pesan a Niuean, A. A, Bees

C. Leo Hitchcock oe

Iris pseudacorus in western North enece Peter ‘H. Raven. Arg John H.
Thomas . :

A new species of Beobocuiic (CUE FEp ae) ao iene Cain, Wiexieo,
Richard H. Hevly .

A new Astragalus (iabaccae) icon Neveds R. C. Bapnepy : :

Notes on Loeflingia (Caryophyllaceae), R. C. Barneby and Ernest C. Gece
Mann. . . « :

Unusual factors Sonbntine iD he Geoecion Si Sonne ae eeqiore
Howard S. Shellhammer, Ronald E. Strecker, H. Thomas gate and
Richard J. Hartesveldt

Linear microspore tetrads in the grass Hae atte Frank W. Childe bs

Extension of the range of Abies lasiocarpa into Galiornia, J.O. Sawyer, D. A.
ee ae and F. F. Bowman .

Index

vii

266
269
276
278
282
289

313
320

SYA
323

326
337,

342
346
347
350

354
359

365
S43

347
385
387
390

392
395

398

408

411

413
417

Dates of publication of Madrono, Volume 20:

No. 1, Feb. 20, 1969
No. 2, Aug. 11, 1969
No. 3, Aug. 20, 1969
No. 4, Mar. 25, 1970

viii

No. 5, Sept. 9, 1970
No. 6, Oct. 29,1970
No. 7, Mar. 17,1971
No. 8, Aug. 6, 1971

A WEST AMERICAN JOURNAL OF BOTANY

The subscription price of Madrofio is $8.00 per year ($4.00 for stu-
dents and $12.00 for institutions). If your institution does not now sub-
scribe to Madrono, we would be grateful if you would make the neces-
sary request.

INFORMATION FOR CONTRIBUTORS

Shorter items, such as range extensions and other biological notes,
will be published in condensed form with a suitable title under the gen-
eral heading, ““Notes and News.”

Abbreviations of botanical journals should follow those in Botanico-
Periodicum-Huntianum (Hunt Botanical Library, Carnegie-Mellon Uni-
versity, Pittsburgh, Pennsylvania, 1968).

Footnotes should be avoided whenever possible.

Membership in the California Botanical Society is normally considered
a requisite for publication in MapRoNo.

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