Auf, Hh 6,0! Pa aah 
PON BES a 
a: 


Md i ie 
4 


iy ' 


<A if | 


‘i aa ? 
“oO 
ee 


oa\ ; 
aS | 
344 ? 
be 
evil et 
ioe ast? | : 

a i 


fh 


‘ul bP a) 

Ne Bets pA ee 
4 Ee i 
al 


é3 
Ty 
ror 


=a | 
a —- en 
3 ad rc) 

é Sioa “et ea ‘ 


ha 
ed 
a Lana 
ey 
ares 
y 
AME 


e 

Sic 

an - 
Sp, 

= 


“eo ‘ 

2 N Ye 
“age iiss os, 

Seite 


Steerer 


| | ie f & 
eee 

eset sate, | a 
? Sees crows Q 


nora, EY 


e 4 1 | 


ray 


as 


Fs 


rye, 
Ice 
‘o, 
es 
0 
an 
LF 
“e 
i) 


AES 
Bie 
EEE 
i : 
Ng 


a 
hs 
“ob 
yam 
w 
a 
fi 


Pe: 
fas 
Si 

aes 


alessio 


1m a 


| . We 


<p %, 


a 


“fg ws 
= 4) 


x 2) ) 
G 
| xe Say 
er a ay : 
~, Ne ce ee ad 


or a i 7. 1% f4 | en 


24N, 


a? a! _ 


"i 


Al | 
oe oad a j 3 ‘A 


a 


= 


9 parce 7 SE 
- wa” 7 »# oe 
XO PA 
ei | 


he 


= 
“ot 


Ss, ‘ T =. or ae 
é Li Me @ cca iA 


EE 
A | 4 i 


= as, 


oe 


Agere 


. & Geeta’ ty Y A 
“¥ wy ae a ete ay ° 8 % 
a4 - i, “i S 4 & ni a 
ts, Py c: a i ca Vs, nad g 5 
ae 
e ‘jl Ne - Sh, ; 
(s fisties 
t i Aj \ i ia , ge % 
is ant “ey 
Vy 


ah, 
nor 


Ree 
ye 


. 


ae | 


i 


er 1 


iM aS ‘ 25, os 
cs . Me 


i 
i 


‘ ceety 
<— H eos alk’ 
WY if Ga _%/ ae |! 
‘“ A eet 2a 


. CBs AN ape 
#: S bs Jg nae 

oe, cr Dee J 4) 
Ss Vy, i soak eee a) 


| 


= 
fig gas teas Pe 
es ae eis, 

Age 


4 ie 
ye 


“a i yy “% 


EY i | 
a a? Sy cy 


ns oe 


Ih 
Se “yy, h i re 


is me 


if, ote Mi - eat ; 


oo : i A - th mee ry : 
4 : _" = OT ry, it > i 


7 


* 
y fr 
' i} 
’ . + : af 
= : 
+ phy 
r i th “ 
ih A ! 7 
ron 
" me J) 4 ) : 
a i 7 
it : 7 ' i - ity) 
ii hy “a O : 
ee Jobe, S ae & 
yoo Ad tf 
a 1 is Uy ja a) 


4 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 
VOLUME XVII 


1963-1964 


BOARD OF EDITORS 


EpGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 
HERBERT F. COPELAND, Sacramento College, California 
JouN F. Davipson, University of Nebraksa, Lincoln 
WALLACE R. ErRNsT, Smithsonian Institution, Washington, D.C. 
HERBERT L. Mason, University of California, Berkeley 
Mitprep E. Martuias, University of California, Los Angeles 
ROBERT ORNDUFF, University of California, Berkeley 
MARION OWNBEY, Washington State University, Pullman 
REED C. ROLLINS, Gray Herbarium, Harvard University 
Tra L. WiccINs, Stanford University, Stanford, California 


Editor—JoHn H. THOMAS 


Dudley Herbarium, Stanford University, Stanford, California 


BUSINESS MANAGER AND TREASURER 
Douglas M. Post 
Biology Department, San Francisco State College 


1600 Holloway Avenue, San Francisco 27, California 


Published quarterly by the 
California Botanical Society, Inc. 


2016 Life Sciences Building, University of California, Berkeley 


Printed by Gillick Printing, Inc., Berkeley, California 


To IRA LOREN WIGGINS. It is with great pleasure and satisfaction 
that the California Botanical Society dedicates this seventeenth volume 
of MaproNo to you. We have long benefited by your wise counsel, both 
during your presidency of the Society and subsequently during your many 
years as a member of the Council. Your studies of plants of western 
North America have spanned the latitudes from the Arctic to the Tropics; 
from these studies have come two impressive works: A Flora of the Alas- 
kan Arctic Slope and the monumental portion, Flora of the Sonoran 
Desert, of the Vegetation and Flora of the Sonoran Desert. And it is 
not only botanists who have been influenced by you. As Director of 
The Arctic Research Laboratory at Point Barrow for four years, fellow 
scientists of all kinds—zoologists, meteorologists, ecologists—benefited 
from their association with you. Too, you have been instrumental in the 
activities of the California Academy of Sciences, having served as its 
president, as a member of its board of trustees, and you have taken an 
active part in its expeditions. Last but not least, there is that host of 
students, many of whom under your versatile guidance really saw for 
the first time the flowers, the birds, the beasts and the rocks of the out-of- 
doors. They return through the years to Stanford University, and go to 
your office door hopefully to have a visit with their favorite professor. 
They are always welcome and go away content, their affection renewed. 


ni ue a 


— ee | 


CONTENTS 


PAGE 


Frontispiece: Ira Loren Wiggins 
Juglans hindsii, the central California black walnut, native or introduced? 


EOE TC VUCMIL GRHLOTIUS CMU cacti ame Re he wn gn dcs acct Se uaa tees ioueieacb te Oe 1 
A controlled hybrid between Sitanion hystrix and Agropyron trachycau- 

JR) aes a 2 10) (2 aaa Ate ek nla nso Ween aes ca Me eens. a ee 10 
Cytotaxonomic observations on Mentzelia, sect. Bartonia (Loasaceae), 

CHIN ee LILO P SOM ete ares escheat Ae ee eae re et, en OL en yh. abc ce 16 
A revision of the genus Thaxterogaster, Rolf Singer and 

PAVE IU Cheol Ls ONUUL IE tee ets ae cet oc Nes R ee ale SS Ose RO 22 
Cytophyletic analysis of Hymenoxys anthemoides, Bernice M. Speese and 

OLAS (ULC TOLILE Jig eae 2 iran td Neon 2h RnB aula co 1 See ieee 35 1 OY ale a a7 
] SRE SRE i a eB tr oe Oe ere ee 30, 66, 89, 140, 170, 233 
IN@teS- amGdeING WS: 22. 2). eee ence rid cet S25 055 Oily ds a7. 14s oe 


197, 203, 235, 268, 280, 294 
Quaternary closed-cone pine flora from travertine near Little Sur, Califor- 


nia, Jean H. Langenheim and J. Wyatt Durham ............22...0....22220000--+- 33 
Chromosome numbers of some phytogeographically interesting Chilean 
POLS VL IV O07 Cnt sera. tan vetoes Aas, tence id it aledone seees tot dast ouch a2 


Chromosome counts in section Erythranthe of the genus Mimulus 
(Scrophulariaceae). II., Robert K. Vickery, Jr., Barid B. Mukherjee, 


AMC CV OCT C WY CGIIS texte est ec Aetna oe aster caren Bont eee. Meee ee eee 6) 
An analysis of variation in Viola nephrophylla, Norman H. Russell and 
TET S) @ OU OSS 10 IUUCR Peace cg eee eh hes tee sage ees 56 


A new species of Isopyrum endemic to the Queen Charlotte Islands of 
British Columbia and its relation to other species in the genus, 


erAt eC GIO CraaTN Gales 1a Ie ON VO) ce coat secu seery sean ee cass nee eee 69 
Cytophyletic analysis of Hymenoxys odorata: A recapitulation, B. L. 

LUE SARA LO ae A NORAD MOET PIE CORI TRRE SDE PENA? OR SNE SES A PE TED OTS Vd 
A note on taxonomic characters in Lolium, Frank C. Vasek and J. Kirk 

Gg CLUS 1 pane Ocak Pir matin set oe etal an P ce ree Ja Oh ger ge Recess ea 79 
Cleistogamy in the Malvaceae, Paul A. Fryxell .........-22....ccc-2ecceneecennnceennees 83 
Swallenia, a new name for the California genus Ectosperma (Gramineae), 

Thomas R. Soderstrom and Henry F. Decker ............22222222.2000000000000---- 88 
Some cordilleran plant species new for the Sierra Nevada of California, 

i IVUIGHOTAANG (Se AN DOMLDETS x0 eee ee eee 2 ee 93 
Natural and artificial hybrids of Besseya and Synthyris (Scrophularia- 

ceae), A. R. Kruckeberg and F. L. Hed glin .....0....2.0.222cccceceecceeeeee eee 109 
Documented chromosome numbers of plants........00...22000.220--e2e-eee ee 116, 266 
Artificial intergeneric hybrids of Helianthus and Viguiera, Charles B. 

15 OSGI P=R | Ran See iy BPC are 2 eee Or Ze RPP eM IR See ca ee tT aR 118 


Chromosome numbers in the Compositae. VII. Additional species from 
the southwestern United States and Mexico, A. M. Powell and B. L. 


BEIRUT get ane Pa ee Oe ach eta ec kT ses hese pes toe eee se ae ae 128 
lantscollection in Nepal, D:D: BHGUt cc... ccc eee eek ee et 145 
Cytological studies in the genus Ficus. III. Chromosome numbers in sixty- 

ENV ONSDECICSNI 1.0.) SCONAIE 2 oe. acs. cccstee teeta Been oe ae 153 


Chromosome counts in the section Simiolus of the genus Mimulus 
(Scrophulariaceae). VI. New Numbers in M. guttatus, M. tigrinus, 
and M. glabratus, M. M. Mia, B. B. Mukherjee, and R. K. Vickery, 


By tage ee a Bo Sr ON a ene eh 208d ce ren esl ewan cscs. eee eels 156 
Extended dormancy of chaparral shrubs during severe drought, R. A. 
ENOTD CAIN EL Ae VE OON CY ace 80 avon oct scat soos a0ha Bel Maazel 161 


Cytological observations on some genera of the Agavaceae, Marion S. 
Oi) ge ae ey SRE ar Sg ee, NON ts 8 an cue PA oe 163 


Cytotaxonomy and distributional ecology of western North American vio- 

lets; Jems Clausen. 053 oe ee eee er ED ee 173 
Nomenclatural problems in the Acacia cornigera complex, Velua E. Rudd 198 
Notes on the leaf epidermis and chromosome number of Swallenia (Gra- 


mineae) @Dennts Andersons 2: | ee eee eee ee ee 201 
The pollen grain morphology of Collomia as a taxonomic tool, Alfred R. 

Loeolich, LL 255 ee ee eee era) oe ote Re ee 205 
A™new species of pine irom Mexico; son Larsen 5 ee 217, 
Two new species related to Clarkia unguiculata, Frank C. Vase ................ 219 
Taxonomic notes on the Chrysothamnus viscidiflorus complex (Astereae, 

Compositae),, ordan- ©: Ard CrsOn = rst, eee ee 222 
David Douglas and the digger pine: Some Questions, James R. Griffin...... 227), 
Bark photosynthesis in ocotillo, H. A. Mooney and B. R. Strain ................ 230 


The genus Xerocomus Quelet in northern California, Harry D. Thiers..... 237 
Survival of transplanted Cupressus and Pinus after thirteen years in Men- 


docino County, California, Calvin McMillan ..0..22...22....22.cc22cccecceeeenees 250 
A peculiar case of hemlock mistletoe parasitic on larch, Job Kuajt ............ 254 
Lyonothamnoxylon from the lower Pliocene of western Nevada, Virginia 

VE TPO SC: tote. Becht u Paces Pade Sa eet os Nines eae ieee eet 251 
The Hordeum jubatum — caespitosum — brachyantherum complex in 

Alaska, WoW. Maichell anda CO Wilione. 22 ee 269 
The genus Eschscholzia in the South Coast Ranges of California, Wallace 

| AGS Dy 2X) aaa BE AER OO OE De PR PPee od tet Mninty Seceey ete ear ge ee iy og iar rit 281 


WiGlex: cos oe ee ey ec ee ge a ee 296 


Borie | 


ew 
VOLUME 17, NUMBER 1 JANUARY, 1963 


Contents 


PAGE 
JUGLANS HINDSII, THE CENTRAL CALIFORNIA BLACK 
WALNUT, NATIVE OR INTRODUCED? 
Harriette H. Thomsen ae 1 


A CONTROLLED HYBRID BETWEEN SITANION HYSTRIX 
AND AGROPYRON TRACHYCAULUM, W.S. Boyle 10 


CYTOTAXONOMIC OBSERVATIONS ON MENTZELIA, SECT. 
BaARTONIA (LOASACEAE), Henry J. Thompson 16 


A REVISION OF THE GENUS THAXTEROGASTER, 
Rolf Singer and Alexander H. Smith 22 


CYTOPHYLETIC ANALYSIS OF HYMENOXYS ANTHEMOIDES, 
Bernice M. Speese and J. T. Baldwin, Jr. 27 


Reviews: Richard Evans Schultes, Native orchids of 
Trinidad and Tobago (Myron Kimnach); Albert N. 
Steward, La Rea Dennis, and Helen M. Gilkey, 

Aquatic plants of the Pacific Northwest with vegeta- 
tive keys (Robert Ornduff); Ira L. Wiggins and John 
Hunter Thomas, A Flora of the Alaskan Arctic Slope 
(S. Galen Smith); Margaret McKenny, The savory 
wild mushroom. A Pacific Northwest guide (Isabelle 
Tavares) 30 


NoTES AND NEws 32 


A WEST AMERICAN JOURNAL OF BOTANY 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


HERBERT L. MAson, University of California, Berkeley, Chairman 
EpcAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 

HERBERT F.. COPELAND, Sacramento College, Sacramento, California 

Joun F. DaAvipson, University of Nebraska, Lincoln 
MItpreD E. MaArTurias, University of California, Los Angeles 24 
MaArIon OWNBEY, State College of Washington, Pullman 
REED C. Rotiins, Gray Herbarium, Harvard University 
Ira L. Wicctns, Stanford University, Stanford, California 


Secretary, Editorial Board—ANNETTA CARTER 
Department of Botany, University of California, Berkeley 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Herbert L. Mason, Department of Botany, University of California, 
Berkeley. First Vice-President: Paul C. Silva, Department of Botany, University of 
California, Berkeley. Second Vice-President: Robert F. Hoover, California State 
Polytechnic College, San Luis Obispo. Recording Secretary: Mary L. Bowerman, 
Department of Botany, University of California, Berkeley. Corresponding Secretary, 
Margaret Bergseng, Department of Botany, University of California, Berkeley. 
Treasurer: Douglas M. Post, Biology Department, San Francisco State College, San 


Francisco, California. 


JUGLANS HINDSII, THE CENTRAL CALIFORNIA BLACK 
WALNUT, NATIVE OR INTRODUCED? 


HARRIETTE H. THOMSEN 


Many anthropologists have noted that a distinctive vegetation tends 
to appear on man’s habitation sites. Hrdlicka (1945) quoted Eyerdam 
as stating that the site of stone age villages in the Aleutians could gener- 
ally be recognized during spring and summer by the predominance of 
two perennial plants, Heracleum lanatum Michx. (wild rhubarb or cow 
parsnip), and Aconitum kamtschaticum Rchb. (monkshood). Lillard, 
Heizer and Fenenga (1939) state that nettles and thistles were often as- 
sociated with village sites in the Sacramento Valley. A more recent obser- 
vation was made by Elsasser (1960) that ‘““mule-ears” (Wyethia mollis 
Gray) sharply defined the extent of a California Indian habitation site 
in the Sierra foothills. 

The association of certain plants and man has an obvious practical use 
in site reconnaisance; a two-year study made by Zeiner (1946) at the 
Angel Mounds in Indiana demonstrated that the location of buried walls 
and earthworks could be traced by the distribution of certain species of 
plants. Although of less immediate application, it would seem that con- 
sideration should also be given to the possibility that a useful chronology 
might be established on the basis of plant progression. In either case, such 
study would necessarily involve the differentiation of the plant cover into 
categories of indigenous and introduced flora. 

The investigator into the relationship of vegetation to man’s occupa- 
tion in California would pretty surely turn to the publications of Willis 
Linn Jepson, whose ‘“‘A Flora of California,” “The Silva of California,” 
and “A Manual of the Flowering Plants of California,’ have been the 
primers for several generations of botanists. Jepson (1909, p. 365; 1910, 
p. 194) made the observation that the central California black walnut 
was to be found near ancient Indian village sites, although little signifi- 
cance was attached to his observation at the time. 

An anthropological study of known Indian sites in the Bay Area coun- 
ties was initiated in an effort to ascertain if the black walnut is, in fact, 
a plant associated with pre-contact habitations of the California Indian. 
In the course of the investigation it became evident that on the basis of 
present-day evidence, definite proof is lacking as to whether or not the 
black walnut of central California is indigenous or introduced. The wide- 
spread use of black walnut as a rootstock for the commercial propagation 
of the English walnut (Juglans regia L.) has proliferated the occurrence 
of the black walnut to such an extent that no Indian association with a 


given stand can be verified or disproved without a determination of its 
origin. 


Maprono, Vol. 17, No. 1, pp. 1-32. January 30, 1963. 


2 MADRONO [Vol. 17 


The anthropological study is continuing; questions which should still 
be answered include: were the riparian locations that were so often occu- 
pied advantageous to both the trees and to the indigenous peoples, hence 
the shade and the fruits were utilized fortuitously? Since so good and 
portable a source of food must have been transported during the seasonal 
migrations of the Indians, is the presence of walnuts at non-riparian loca- 
tions the result of accidental seeding by man or by rodents? Or is their 
location in sites well above permanent water courses the result of natural 
distribution and an indigenous origin, the present isolated groves of trees 
representing relictual stands? 

In order to attempt an answer to the possibility of natural origin as 
contrasted to introduction by Indians, the writer was led into a study of 
the fossil record, with the result that a case is made herein for the distri- 
bution of the central California black walnut, Juglans hindsii (Jeps.) 
Jeps., as an indigenous plant of central California. 

Six to nine genera are recognized by taxonomists dealing with Jug- 
landaceae; the present investigation will be concerned with three of these: 
Carya, Pterocarya, and Juglans. Of the three sections in the genus Juglans 
(sections Rkysocaryon, Cardiocaryon and Dioscaryon), the discussion 
will deal primarily with section Rhysocaryon, since all of the western 
North American species of Juglans can be assigned to this section. Wolfe 
(1959, p. 13) considers that morphologically, 

“There are two distinct groups of species in R/ysocaryon. One of these 
groups, including J. nigra and the Central and South American species 
tends to have leaflets which have a broad base and a broadly triangular 
(ovate) shape. ... The other group of species have teeth which are broad- 
ly triangular and dentate, or narrowly conical and serrate. The shape is 
narrowly triangular although a narrowly quadrate condition prevails in 
J. californica’. Wolfe’s work is mainly concerned with foliar character- 
istics, and minor attention is given to fossil fruits. An essential basis for 
differentiation between the extant species of Juglans which are either in- 
troduced or occur naturally in California lies in the fruits. Unlike its 
eastern cousin, the rugose J. nigra L., which has a strongly grooved black 
shell, or the important commercial so-called English walnut, J. regia L., 
the central California black walnut, J. Aindsi (Jeps.) Jeps., has a char- 
acteristically smooth or very lightly-grooved nut, light brown in color. 
In shape, the fruits of both J. migra and J. regia are characteristically 
longer and more pointed than in J. Aindsii. Specimens of J. Aindsu fruits 
taken in widely separated parts of the central California habitat show a 
consistent roundness, with a flattened stem end, reminiscent of the tiny 
Eocene J. clarnensis Scott. Foliar characters differ among the three, 
especially in the number of leaflets: J. Aindsiz has as many as nineteen 
leaflets and seldom fewer than fifteen; J. migra may have as many as 
twenty-seven leaflets, while J. vegza usually has seven, but sometimes five. 

There are two centers of distribution for California black walnuts, one 


1963 | THOMSEN: JUGLANS 3 


in the southern part of the state and one in the north-central part, the 
trees occurring without connecting localities. The northern California 
black walnut was discovered along the lower Sacramento River area by 
Richard Brinsley Hinds of the Sulphur Expedition in 1837. By an his- 
torical accident, however, the black walnut of southern California was 
described first as J. californica by Watson (1875), who included within 
his concept J. rupestris Engl. var. major Torr., the walnut of Arizona and 
Texas. Jepson (1908) named the northern California trees as a variety 
of the southern ones, J. californica var. hinds, honoring their discoverer 
and saying that the trees of the two regions “‘differ somewhat.” 

In his Flora, Jepson (1909) still contented himself with this varietal 
concept, averring that the northern trees were introduced from southern 
California by the trading of the native Indian tribes. In his Silva (1910), 
he continued the same concept but pointed out certain anomalies, namely 
that (1) Watson’s original description of J. californica did not cite a type 
and included as a synonym the walnut of Arizona and Texas, J. rupestris 
var. major, and (2) the southern, rather than the northern California 
walnuts must bear the name J. californica Wats. for several reasons: Wat- 
son had no northern specimen before him, his northern locality was vague, 
his description better fitted the southern form. It was not so easy, how- 
ever, for Jepson to ignore a further anomaly, the peculiar gap in distri- 
bution between southern and northern California (275 miles between 
stations in Ventura County and near Mount Diablo), and he concluded 
again that the introduction from the south by native tribes was the only 
plausible explanation for the existence of the northern trees. Later, how- 
ever, in his Manual (1923) he elevated the northern trees to the rank of 
a species, without further discussion, as J. hindsii (Jeps.) Jeps., a tacit 
admission that he must have come to the point of view that the original 
black walnuts of central California, present before the white man arrived, 
were indigenous and were not introduced from those of southern Califor- 
nia. His morphological treatment bears this out. 

A review of the paleontological literature of the past half century may 
give perspective to the facts of distribution of California Juglans. Since 
the fossil beds are distributed in space as well as in time, the discussion 
of these deposits and their relation to Juglans can be more easily followed 
if they are examined by geographical groupings as well as by geological 
time periods. A glance at a relief map of the western United States dis- 
closes that territorially the area is broken up into clearly defined regions 
by the Coast, the Cascade and the Sierra Nevada mountain ranges; these 
ranges run roughly parallel longitudinally. Minor ranges, the Klamath to 
the northwest, and the Warner to the northeast, complete the California 
picture. One major transmontane range, the Tehachapi Mountains, shuts 
off northern from southern California, forming the southern rim of the 
great Central Valley, whose rivers converge from north, east and south 
towards San Francisco Bay. 


4 MADRONO [Vol. 17 


Some paleontological literature has been available since Jepson first 
became interested in the problem of Juglans and its distribution. Sud- 
worth (1908) remarked on the presence of fossil walnut remains in Cre- 
taceous and Tertiary formations, noting that “. ... in the northern Pa- 
cific coast region signs of ancient walnuts have been obtained from the 
Eocene formation, as well as from the gold-bearing gravel beds of the 
California Sierra’’. There follows a brief and very simplified discussion of 
the subsequent paleontological literature which bears on the subject. 

The fossil floras of California, Oregon and Washington indicate tropi- 
cal conditions for the Eocene, although changes in physiography due to 
volcanic action and mountain uplift were making for localized climates. 
Scott (1954) described the fossil fruits and seeds from the Eocene Clarno 
formation of north-central Oregon. He discussed the similarity of the nuts 
of Carya and Juglans, designating the fruits of the Clarno species, Jug- 
lans clarnensis, and describing them as having features intermediate be- 
tween those of sections Cardiocaryon and Rhysocaryon. He says: “. . This 
is the first certain walnut described from rocks as old as Eocene. This 
occurrence of Juglans demonstrates that the fruits of the walnut and the 
hickory (Carya) were generically distinct during Eocene time’”’. It should 
be remarked here, however, that Wolfe (1959, p. 43) assigns J. clarnensis 
Scott to the Oligocene. 

In the Oligocene, tropical fauna were still ranging up to Puget Sound, 
and the fossil flora of southern Alaska indicates a warm, temperate zone 
at that latitude (Smith, 1919). In other parts of North America, such as 
in North Dakota and Colorado, fossil beds containing Juglans material 
have been related to Oligocene time, and Wolfe (1959, p. 46) places 
J. kentensis, from a bed in northwest Oregon, in that time period. 

MacGinitie (1937, p. 112) did an extensive study of the Weaverville 
beds, which he assigned to the Oligocene. The beds lie in the Klamath 
Range of northwestern California, clustered near the Trinity River, 
hemmed in between ocean and the rising mountains to the east. The dif- 
ficulties inherent in the identification of fossil material are well illustrated 
in the case of the Weaverville Juglans. Diller (1911) quotes Knowlton 
as having made determination of Juglans schimperi Lx. [reinterpreted by 
MacGinitie (1937) to Inga lancifolia MacG.]|, Juglans egregia Lx. and 
J. oregoniana Lx. from the Weaverville flora. MacGinitie reassigned both 
of the latter specimens to Juglans orientalis. He characterized his ex- 
panded interpretation of J. orientalis MacG. as having a much closer 
resemblance to various living species of Juglans in eastern Asia than to 
living forms in North America. LaMotte (1936) working in the upper 
Cedarville flora of the northern Great Basin (east of the Cascades and 
north of the Sierra Nevada), transferred Juglans oregoniana Lx. to Carya 
with what appears to be fairly cogent argument. 

During an investigation of the Pliocene San Pablo beds, Condit (1938) 
identified fossil leaves from those beds as J. oregoniana Lx., defending 


1963] THOMSEN: JUGLANS 5 


his identification vigorously in opposition to LaMotte’s reassignment of 
this species to Carya. As far as the records of these four investigations 
show, the workers had leaves or leaflets only, and no fruits. But whether 
Juglans or Carya, the species described by MacGinitie as J. orientalis 
MacG. disappears from the North American scene, apparently unable to 
cope with the environmental changes of succeeding ages. No descendants 
are recognized in California today. 

An obvious geographical unit in the western United States is that which 
centers in the Cascades. This mountain range, extending north-south 
from the Canadian border across Washington and Oregon, penetrates 
northern California for a brief distance, curving off toward the east and 
culminating at Mount Lassen, thus lying entirely north and west of the 
Sierra Nevada. This is an area in which extensive paleontological studies 
have been made. Chaney carried out investigations of the fossil floras of 
three Pliocene sites in the Cascades: Deschutes (1938), Dalles and Trout- 
dale (1944 a,b); he reported no Juglans from his studies of the Troutdale 
and Dalles flora, but described a species in another genus of the same 
family, Pterocarya oregoniana Chaney. He placed it in the East Asian 
Element, and listed P. stenoptera DC. as its nearest living equivalent 
species. The extant P. stenoptera, common in parts of China, is not gen- 
erically represented in North America today. Chaney’s study of the 
Deschutes flora (1938, p. 203) disclosed no evidence of either Juglans 
or Pterocarya. 

Axelrod first studied the Sonoma flora in 1944 and later (1950a) he 
issued a report on “A Sonoma Florule from Napa, California”’, in which 
he identified Pterocarya oregoniana Chaney. In his digest of geological 
history which is included in Munz and Keck’s ‘“‘California Flora” (1959, 
p. 6) Axelrod stated that Pterocarya, together with a few other East 
American and East Asian species persisted in the mild coastal strip from 
central California northward, becoming extinct in the early part of the 
Pleistocene. 

The wide area east of the Cascades is rather better documented than is 
the area to the west. This eastern region, tenanted by plant immigrants 
from the holarctic forest, provided a favorable environment for the de- 
velopment of a significant Miocene flora. LaMotte (1936, p. 90) believed 
that the general outlines of North America in upper Miocene time may 
be considered to have been much the same as today, the physiography 
and the inferred climate providing conditions favorable to an extremely 
uniform fossil flora. He reports the finding of numerous leaflets (1936, 
p. 116), with at least two specimens of leaves showing leaflets in place, 
but identifies the entire group as Carya egregia Lx., feeling that the char- 
acteristics resembling hickory outweighed those resembling Juglans. 
What seems important here is not that the botanists disagree on nomen- 
clature, but rather that the evidence for a dominant broad-leafed decid- 
uous forest of oak-hickory-beech association thrived east of the Cascades 


6 MADRONO [Volal7 


in Miocene time, an environment that would have been favorable to the 
persistence of the many species of Juglans found in Washington, Oregon 
and Idaho in preceding and contemporary fossil beds. Berry’s work 
(1927) on the flora of the Esmeralda beds convinced him that all evi- 
dence pointed to the presence of a permanent body of water in western 
Nevada in the Miocene. 

Wolfe (1959, pp. 46, 47, 48)! describes two fossil Juglans from the 
Miocene: J. hesperia from northwestern Oregon and J. fryi from the 
northeastern part of the state. Both of these fossil forms had resemblances 
to J. kentensis referred to in the discussion of Oligocene specimens, but 
also exhibited what he considered to be significant morphological differ- 
ences. Chaney (1927, p. 74) identified materials from eastern Oregon as 
J. oregoniana Lx., and MacGinitie (1933, p. 50) identified material from 
southern Oregon as the same species, but Chaney considered the nearest 
living equivalent to be J. californica Wats., whereas MacGinitie consid- 
ered his specimen to be closer to J. migra L. 

Two more references to Miocene Juglans appear in the literature. The 
first is J. nevadensis Berry (1928), exact provenience unknown, but found 
in the desert east of Truckee. Berry believed it to have a possible affinity 
to J. regia or J. sieboldiana Maxim of Japan, basing such evaluation on 
the lack of corrugation in the shell surface. The second is from Axelrod 
(1950b) who in his restudy of some of the Middle Pliocene Mount Eden 
flora, amended his description of J. beaumontiu Axelrod to include two 
species: a) J. beaumont Axelrod emend. with an affinity to J. rupestris 
Engelm. of the southwest United States and b) J. nevadensis Berry, de- 
scribed as having a smooth outer shell-coat with minor surface irregu- 
larities and greatest affinity to “J. californica of southern California”’ 
(p. 102). 

Dorf (1936) investigated the Weiser beds of southwestern Idaho. This 
site, east of the rising Cascades, is placed in the Upper Miocene or Lower 
Pliocene time. Dorf imputed a more xeric environment with a rainfall 
of probably less than twenty-five inches. He described J. hesperia Knowl- 
ton which he related to J. nigra L., pointing out the similarity of the in- 
ferred Weiser climate to the climate of today in the eastern United States. 

It would appear, therefore, that already in Miocene time, some Juglans 
species were established on the east side of the Sierra Nevada. Chaney 
(1938) concluded that the northern Sierra was sufficiently high at the 
beginning of the Miocene to have eliminated most of the genera requiring 
summer rainfall. Isolated there by the rising range and cut off from ocean 
moisture, the conditions became increasingly xeric, and the genus faced 
the inevitable choices which changing environments present to the plant 
world: adapt to change, migrate, or become extinct. 

We come now to a consideration of the third geographical unit, the 


1Juglans hesperia Knowlton has been transferred to Salix; Juglans hesperia Wolfe 
and J. fryi Wolfe (as well as J. kentensis Wolfe) are unpublished. 


1963 | THOMSEN: JUGLANS 7 


great Central Valley which lies south of the Cascades, east of the Coast 
Range, and west of the Sierra Nevada, closed off at the south by the 
Tehachapi Mountains. A number of fossil localities have been studied in 
the Coast Range and in the western foothills of the Sierras (see distribu- 
tion map, Chaney, 1944c). The Chalk Bluffs, La Porte and Oakdale beds 
have shown no material identified as Juglans, although the Chalk Bluffs 
flora did include specimens identified by MacGinitie (1941, p. 101) as 
Carya sessilis. 

With Condit’s studies of the Remington Hill and Table Mountain 
floras (1944a, b), we reach the southern limit of fossil Juglans reported 
within the Central Valley. The Remington Hill and Table Mountain 
floras occur in, to quote a favorite phrase of the early paleontologists, “‘the 
auriferous gravels of the Sierras”. The Sierras of the Pliocene were yet 
of moderate height and the Table Mountain beds, located in a region 
drained by the Tertiary equivalents of the Merced and Stanislaus Rivers, 
may have been laid down at an elevation not more than five hundred feet 
above the level of a sea which lapped at the foothills—a sea which was 
a hundred miles or so farther inland than it is today. Condit reports that 
the Remington Hill beds are at an altitude of 3840 feet today, which 
would imp'y that they were laid down at a greater elevation than the beds 
of Table Mountain. 

Condit (1944a, p. 42) described a fruit from the Remington Hill beds 
which he named Juglans pseudomorpha and thought might be related to 
J. nigra, but the association in the fossil flora suggested that it was inter- 
mediate between the typical eastern black walnut and the living Cali- 
fornia black walnuts, “J. californica of Southern California as well as 
J. hindsii of the inner north coast range bordering on the Sacramento 
Valley” (p. 28). Wolfe (1959, p.49) was of the opinion that Condit’s 
specimen is undoubtedly a Juglans, but that it is too badly crushed to 
permit comparisons. 

Condit’s study of the Table Mountain flora (1944b) included a re- 
naming of Lesquereux’s Rhus typhinoides to Carya tvphinoides. He dis- 
cussed the similarities of Carya and Juglans at length, and considered 
the possibility that on ecological grounds C. typhinoides should be as- 
signed to Juglans rather than to Carya. 

Wolfe (1959, p. 47) reassigned the Table Mountain leaflets to Juglans, 
giving them the name J. tuolumnensis. He was of the opinion that this 
fossil might have been derived through J. kesperia Wolfe, mentioned in 
the discussion of Miocene flora of northwestern Oregon, but he saw a 
more obvious relationship with J. californica Wats. 

Fossil records for the Pleistocene in western North America are so 
sparse as to be almost non-existent. One allusion in the literature is made 
by Axelrod (1944, p. 118), wherein he refers to “..... the occurrence 
of a characteristic fruit of /uglans californica Watson in the Pleistocene 
of the San Joaquin Valley (H. L. Mason, oral communication, July, 


8 MADRONO LNVoleaty/ 


1940)”. Verification of Pleistocene remains, perhaps through fossil walnut 
pollens from the San Joaquin Delta peats, would greatly reinforce our 
understanding of pre-Holocene distribution. 

To summarize the fossil record: 

(1) Members of the Juglandaceae, such as Juglans orientalis MacG. 
and Pterocarva oregoniana Chaney, which flourished in the western parts 
of California in the Oligocene and Pliocene, respectively, appear to have 
become extinct not later than early Pleistocene; as far as is known the 
only living equivalents are now found in Asia. 

(2) The rise of the Cascades and of the Sierra Nevada in the Miocene 
isolated species of Juglans into westward and eastward slope environ- 
ments; across Oregon and Idaho and down the eastward side of the Sier- 
ras there appears to have been a migration of Juglans tolerating more 
xeric conditions, the plants developing a low, bushy conformity with 
foliage and fruit showing tendencies toward desert-shrub. 

(3) By the Pliocene there were on the west side of the Sierra Nevada 
several species of Juglans that had made the adaptation from the sub- 
tropical environments of the Eocene and were flourishing in the cooler, 
summer-dry climate of the Pliocene at elevations now comparable to the 
lower Sierra Nevada. They occurred in the vicinity of the Tertiary rivers, 
probably not too high above the then great inland sea which reached to 
the foothills of the Sierra. It does not seem to be merely a coincidence 
that one finds today isolated stands of great trees at an altitude of around 
1500 feet, on both sides of the Central Valley. 

(4) As the Pleistocene ice advanced and the temperature lowered, the 
waters receded and the outline of the Central Valley was indicated. Three 
courses were open to the flora: it could stand steadfast and become ex- 
tinct, it could adapt itself to the environmental change, or it could mi- 
grate toward the south or toward the coast. It has been generally accepted 
that portions of the California flora migrated southward, advancing and 
retreating with the deterioration of the Tertiary climate. This assumption 
should be generally valid. Specifically, however, the record would indicate 
that it was not necessarily true of every genus. In the matter of Juglans, 
it would seem that there may have been a separation as early as Miocene 
time, with the result that J. californica Wats. was derived perhaps through 
J. nevadensis Berry, J. rupestris Engelm., J. beaumontiu Axelrod emend., 
or other similar small-leaved, small-fruited species, which tolerated the 
more xeric environment of the southern parts of the United States, where- 
as the ancestors of J. hindsiui (Jeps.) Jeps., may well have advanced coast- 
ward before the cold, following down the streams of the Central Valley, 
never approaching the coast beyond the edge of the fog-drip belt, retain- 
ing its mesic characters of lofty height, lush foliage, and smooth, faintly- 
grooved flattened nut. 

It can be conciuded, therefore, that the climate and physiography were 
such as to make possible, or even probable, the independent evolution of 


1963 | THOMSEN: JUGLANS 9 


the southern California black walnut, J. californica Wats. and the north- 
ern species, J. Aindsii (Jeps.) Jeps., neither species being derived from 
the other. As a corollary, if the theory of separate and indigenous species 
is borne out by competent botanical investigation, the “anomaly” of dis- 
tribution raised so frequently in the past, does not exist; the presence of 
J. hindsii in north-central California assumes an aspect of rightness and 
logic, and requires no Indian trade to account for its narrow distribution 
in the Central Valley drainage basin and its focus in the San Francisco 
Bay area. 
Berkeley, California. 


LITERATURE CITED 


AXE LRoD, D. I. 1944. The Mulholland Flora, Carnegie Inst. Publ. 553:103-146. 
. 1950a. A Sonoma Florule from Napa, California. Carnegie Inst. Publ. 590: 
23-71. 
———-—. 1950b. Further studies of the Mount Eden flora, Southern California. 
Carnegie Inst. Publ. 590:75-117. 
Berry, E.W. 1927. The flora of the Esmeralda formation. Proc. U.S. Nat. Mus. 


72 23,.1-15, 
— . 1928. A petrified walnut from the Miocene of Nevada. Jour. Wash. Acad. 
18:6, 158-160. 


CHANEY, R.W. 1927. Geology and paleontology of the Crooked River Basin, with 
special reference to the Bridge Creek flora. Carnegie Inst. Publ. 346:45-138. 
——, 1938. The Deschutes flora of eastern Oregon. Carnegie Inst. Publ. 476: 

185-216. 
——-——. 1944a. The Dalles flora. Carnegie Inst. Publ. 553:285-321. 
1944b. The Troutdale flora. Carnegie Inst. Publ. 553:323-351. 
1944c. Pliocene floras of California and Oregon. Carnegie Inst. Publ. 
533:frontispiece. 

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

1944a. The Remington Hill flora. Carnegie Inst. Publ. 553:21-56. 

—. 1944b. The Table Mountain flora. Carnegie Inst. Publ. 553:57-90. 
DiterR, J. S. 1911. The auriferous gravels of the Trinity River basin, California. 
Contr. to Economic Geology, Dept. of Int. U.S. Geol. Sur. Bull. 470:11-29. 
Dorr, E. 1936. A late Tertiary flora from southwestern Idaho. Carnegie Inst. Publ. 

476:73-124. 

ExsAsser, A.B. 1960. The archaeology of the Sierra Nevada in California and 
Nevada. Univ. Calif. Arch. Surv. 51:1-93. 

Hrouicka, A. 1945. The Aleutian and Commander Islands and their inhabitants. 
The Smithsonian Inst., published by ‘ The Wistar Inst. of Anatomy and Biol- 
ogy, Phila. 43. 

Jepson, W. L. 1908. The distribution of Juglans californica Wats. Bull. S. Calif. 
Acad. Sci. 7:23-24. 

—————. 1909. Juglans, im Flora of California 1(2) :365. 

1910. The silva of California. Univ. Calif. Memoirs 2:192-195. 
1923. Juglans, 7x A manual of the flowering plants of California, p. 279. 
Assoc. Students Store, Univ. Calif. 

Know ton, F. H. (as quoted by J. S. Diller). 1911. U.S. Geol. Sur. Bull. 470:23-24. 

La Morte, R. S. 1936. The upper Cedarville flora of northwestern Nevada and 
adjacent California. Carnegie Inst. Publ. 455:57-142. 

Litiarp, J. B., R. F. Herzer and F. Fenenca. 1939. An introduction to the arche- 
ology of Central California. Sacramento Jun. Coll. Bull. 2:65. 


10 MADRONO [Vol. 17 


MacGrniTtrz, H. D. 1933. The Trout Creek flora of southeastern Oregon. Carnegie 
Inst. Publ. 416:21-68. 

1937. The flora of the Weaverville beds of Trinity County, California. 
Carnegie Inst. Publ. 465:85-151. 

. 1941. A Middle Eocene flora from the Central Sierra Nevada. Carnegie 
Inst. Publ. 534:1-178. 

Muwz, P. A., in collaboration with David D. Keck. 1959. A California flora. Univ. 
of Calif. Press, Berkeley and Los Angeles. 

Scott, R. A. 1954. Fossil fruits and seeds from the Eocene Clarno formation of 
Oregon. Palaeontographica, Band 96, Abt. B: 66-97. 

SMITH, J. P. 1919. Climatic relations of the Tertiary and Quaternary faunas of the 
California region. Proc. Calif. Acad. 4th Ser. IX:4, 123-173. 

SupworTH, G. B. 1908. Forest Trees of the Pacific Slope. U.S. Dept. Agr. 

Watson, S. 1875. Juglans californica Wats., in Revision of the genus Ceanothus, 
and description of new plants. Proc. Am. Acad. 10:333-350. 

Wo re, J.A. 1959. Tertiary Juglandaceae of western North America. Submitted 
in partial satisfaction of the requirements for degree of Master of Arts in Pale- 
ontology, Univ. Calif. 1-96. 

ZEINER, H. M. 1946. Botanical Survey of the Angel Mounds Site, Evansville, Indi- 
ana. Am. Jour. Bot. 33:83-90. 


A CONTROLLED HYBRID BETWEEN SITANION HYSTRIX 
AND AGROPYRON TRACHYCAULUM 


W.S. BoYLe 


Students of evolution have become increasingly aware of the extensive 
hybridization that exists between genera and species in the grass family, 
particularly the tribe Hordeae. Probably no other family in the plant 
kingdom is destined to undergo such a fundamental revision of concepts 
of genetic relationships among genera as is occurring gradually in the 
Gramineae. 

The present paper reports the meiotic chromosome behavior, fertility, 
and comparative morphology of a controlled hybrid between Sztanion 
hystrix (Nutt.) J. G. Smith and Agropyron trachycaulum (Link) Malte. 


MATERIALS AND METHODS 


Specimens of A. trachycaulum growing in fields near Logan, Utah, and 
those of Sitanion hystrix from Mantua, Utah, were transplanted to a 
field nursery in 1954. The crosses were made the following year. 

Forty florets involving several spikes of S. Aystrix were hand-emascu- 
lated in June. Mature culms of Agropyron trachycaulum were placed in 
bottles of water and the bottles taped to stakes driven in the ground 
beside the culms of Sztanion hystrix. Each culm of S. Aystrix, with its 
adjacent pollinators, was then covered with Kraft paper sacks. 

Two seeds were harvested in early August and planted later that same 
month in the greenhouse. The plants grew vigorously and a few spikes 
were produced the following year. One plant proved simply to be a selfed 


1963 | BOYLE: HYBRID GRASS al 


S. hystrix, but the other was indeed the hybrid. This was divided and 
repotted. 

From 1959-1960 the plants grew well in the greenhouse but the pro- 
duction of flowering culms was sporadic. It was concluded that the high 
summer temperatures in the greenhouse interfered with normal anthesis; 
therefore the five clones were transplanted to the field nursery. In the field, 
they grew very well and flowered normally (fig. 2). The importance of 
field observations was emphasized by the contrast between plants grown 
in the greenhouse and the same plants grown in the field nursery. In addi- 
tion to much greater height, the plants grown in the field possessed inflor- 
escences larger in all respects than those of plants grown in the greenhouse. 


B 


Fic. 1. Meiosis in the hybrid: A, metaphase I, pollen mother cell, 12 II, 1 IV 
(x 1100); B, telophase II, pollen mother cell, lagging chromosomes (x 1315). 


Numerous spikes were removed in the boot stage and fixed in New- 
comer’s solution (1953). Observations and photographs were taken from 
acetocarmine smears in temporary mounts. 


MEIOTIC CHROMOSOME BEHAVIOR 


METAPHASE I. Chromosome associations in the 204 pollen mother cells 
that were interpreted are summarized in Table 1. Although 24 different 
types of chromosome association were observed, approximately 75 per 
cent fell into the following categories: 14 II (23.5 per cent); 12 II, 11V 
(Zoro per cent): 13 1102 1 (l4./ percent); 11 11, 1 IV, 2 1 (iz7 per 
cent). Figure 1A is representative. Bivalents were present in all cells and 
averaged 12.2 per cell. Univalents averaged 1.13 per cell with a range 
of 0-6. Over half the cells possessed one or more quadrivalents, with an 


12 MADRONO [Viole17 


average of 0.6 per cell and a range of 0-3. Three trivalents and one hexa- 
valent were observed. 

TELOPHASE I, II. Approximately a third of the 847 cells examined at 
telophase I contained one or more lagging chromosomes (Table 2). They 
averaged 0.6 per cell and had a range of 0-6. Nearly half of the 1756 
cells examined at telophase II contained one or more lagging chromo- 
somes, with an average of 0.78 and a range of O-7 (fig. 1B). 


Fic. 2. The hybrid in the field (x 146). Fic. 3. Spikes: A, Sitanion hystrix; B, the 
hybrid; C, Agropyron trachycaulum (* %). 


Tetraps. Micronuclei averaged 0.37 per cell in the 2361 pollen grains 
examined at the tetrad stage. This is equivalent to 1.43 micronuclei per 
pollen mother cell. Approximately 70 per cent were without micronuclei. 
The maximum number of micronuclei in any one pollen grain was 3. 

PoLueNn. The pollen was almost entirely abortive. Of the 9064 mature 
pollen grains examined, only 3 appeared to be fertile. 


FERTILITY 


In a careful search of approximately 2500 florets, not a single seed was 
found. The hybrid is completely sterile. 


COMPARATIVE MORPHOLOGY 
The parental species are both highly variable, as the long synonomy 
lists suggest. However, obvious general differences exist between the two 
parents with respect to the morphology of the inflorescence. Except in 
size relationships, the hybrid is approximately intermediate between the 
two parents (figs. 3, 4). 


1963] BOYLE: HYBRID GRASS 13 


TABLE 1. CHROMOSOME ASSOCIATION AT METAPHASE I. 


I II II IV VI Number 
cells 

14 47 
12 1 47 

2 13 30 

2 11 1 26 

4 i did 
10 Z 11 

4 IK@) 1 ‘| 
i) 1 5 

Z 9 2 3 

3 11 1 Z 

1 13 1 1 

2 12 1 

6 11 1 

5 10 1 1 
Ww 2 1 

5 10 1 1 

4 9 1 1 

2 7 3 1 
11 1 ] 

4 11 1 
11 1 1 

5 10 1 1 
13 1 

1 12 1 1 

Average 1.13 ay 0.19 0.60 0.004 204 
per cell Total No. 


In Agropyron trachycaulum the spikelets occur singly at the nodes, the 
rachis does not disarticulate, the glumes are very broad, and both glumes 
and lemmas are awnless. In Sitanion hystrix the spikelets usually occur 
in pairs, the rachis readily disarticulates at maturity, the glumes are very 
narrow, and both glumes and lemmas terminate in long, frequently 
twisted awns. 

In the hybrid the spikelets occur singly at the nodes, the rachis tardily 
disarticulates at maturity, the glumes are moderately wide, and both 
glumes and lemmas have fairly short (1'4 cm.), diverging awns. The 
hybrid is a vigorous, tall bunchgrass apparently possessing considerable 
hybrid vigor. 


DISCUSSION 


The average number of univalents per pollen mother cell approximates 
the average number of micronuclei per pollen mother cell. The univalents 
would be expected to lag at first and second telophase and doubtless are 
the source of the micronuclei. The average number of laggards reported 
per cell at telophase I and II, however, was much lower than expected. 
A plausible explanation for this discrepancy could be our overly conserva- 


14 MADRONO [Vol. 17 


tive tendency in identifying laggards. Only chromosomes in the center 
of the plates were stipulated as laggards (fig. 1B). Very likely other 
chromosomes that approached the poles nevertheless failed to be included 
in the new nucleus and remained outside to form micronuclei. 

The absence of quadrivalents in cytological studies of the parents tends 
to suggest that both are allotetraploids, and that the pairing in the hybrid 
is allosyndetic. Therefore these two species may have more or less homol- 
ogous genomes. On the other hand, Wagenaar (1959) has convincingly 
demonstrated autosyndesis among Hordeum chromosomes in a hybrid 
between H. jubatum L. and Secale cereale L. No quadrivalents were ob- 
served in the Hordeum parent. Dewey (1961) has conclusively demon- 
strated autosyndesis among chromosomes of Agropyron repens (L.) 
Beauv. and A. desertorum (Fisch.) Schult. in a hybrid between them. 
Stebbins and Pun (1953) similarly demonstrated that chromosomes of 
A. intermedium (Host.) Beauv. paired autosyndetically in a hybrid be- 
tween that species and S. cereale. 


TABLE 2. FREQUENCY OF LAGGING CHROMOSOMES AT TELOPHASE I AND TELOPHASE II. 


Average number of Percent with one Number 


laggards per cell or mcre laggards of cells 
Telophase I 0.60 34.1 847 
Telophase II 0.78 48.5 1/56 


Since the writer cannot distinguish between the parental chromosomes 
in the hybrid, the precise nature of the pairing obviously cannot be in- 
ferred with confidence. The high frequency of quadrivalents, however, 
permits some speculation on homologies between these species. Over 50 
per cent of the metaphase I plates possessed one or more quadrivalents. 
It is most unlikely that reciprocal translocation is responsible for any 
substantial percentage of these quadrivalents since they are absent in 
approximately half the cells and occur in variable numbers of the cells 
that do contain them. Even if it is conceded that the bivalent pairing may 
be exclusively autosyndetic, which is by no means established, the high 
number of quadrivalents suggests that important homologies exist be- 
tween Agropyron trachycaulum and Sitanion hystrix. The degree of ho- 
mology cannot be accurately assessed at present, but these two species 
unquestionably are much more closely related than their present taxo- 
nomic status indicates. 

Stebbins et al. (1946) reported a controlled hybrid between Sztanion 
hystrix and a species of Agropyron (“San Benito”) believed to be most 
closely related to A. parishii Scribn. & Smith but originally identified as 
A, trachycaulum. The bivalent frequency (12.7 per cell average) closely 
approximates that of the hybrid reported in this paper. The frequency 
of univalents, laggards, and micronuclei in Stebbins’ hybrid, however, was 
much higher and the quadrivalent frequency was much lower than those 


1963 | BOYLE: HYBRID GRASS 15 


A B C 


Fic. 4. Spikelets: A, Sitanion hystrix; B, the hybrid; C, Agropyron trachycau- 
lum (X 2). 


found in the hybrid of this study. In the Stebbins’ contribution the 
authors suggested that A. saundersi (Vasey) Hitchc. is probably an F, 
hybrid between A. trachycaulum and Sitanion hystrix. The present study 
lends some, but not complete, support to this proposition. The type speci- 
men of Agropyron saundersi has considerably longer awns and shorter 
spikes than the controlled hybrid produced in the present study. Fur- 
thermore, in the type specimen of A. saundersti the spikelets are frequent- 
ly paired, in contrast to the single spikelets of the hybrid of the present 
study. Professor Arthur H. Holmgren has made the very plausible sug- 
gestion (oral commun.) that Sitanion longifolium J. G. Smith has served 
as one parent of Agropyron saundersiu. This species (sometimes referred 
to Sitanion hystrix) has large, coarse spikes with long, straight awns, and 
could well account for the differences observed between A. saundersti and 
the hybrid of the present study. Sitanion longifolium is known to be pres- 
ent in the general region where the type of Agropyron saundersii was 
collected. 


SUMMARY 

A controlled hybrid between Sitanion hystrix and Agropyron trachy- 
caulum is reported. Although 24 different types of chromosome associa- 
tions were observed at metaphase I in 204 interpreted pollen mother cells, 
75 per cent fell into the following four categories: 14 II; 12 II, 1 IV; 
13 II, 2 1; 11 I, 1 IV, 2 I. Bivalents were present in all cells and aver- 
aged 12.2 per cell. Over half of the pollen mother cells contained one or 
more quadrivalents. Univalents averaged 1.13 per cell. Lagging chromo- 
somes averaged 0.60 per cell at telophase I and 0.78 at telophase IT. 


16 MADRONO [Vol. 17 


Micronuclei averaged 1.43 per pollen mother cell. The hybrid is com- 
pletely sterile. 

Although it was not possible to distinguish between autosyndetic and 
allosyndetic pairing, the high frequency of quadrivalents in the hybrid 
suggests that important homologies exist between the parental species. 

The hybrid is morphologically intermediate between the two parents 
except for size relationships. Some support is given to the suggestion that 
Agropyvron saundersu had a similar origin. 


ACKNOWLEDGMENT 


The author wishes to express his appreciation to Professor Arthur H. 
Holmgren for many helpful discussions and to Robert Webster and Mrs. 
William Allderdice, student assistants. 


Utah State University 
Logan, Utah 


LITERATURE CITED 

Dewey, D.R. 1961. Hybrids between Agropyron repens and Agropyron desertorum. 
Jour. Hered. 52:13-21. 

Newcomer, E. H. 1953. A new cytological and histological fixing fluid. Science 
TS 2VG1: 

STEBBINS, G. L., and Func Tine Pun. 1953. Artificial and natural hybrids in the 
Gramineae, tribe Hordeae. VI. Chromosome pairing in Secale cereale « Agro- 
pyron intermedium and the problem of genome homologies in the Triticineae. 
Genetics 38:600-608. 

STEBBINS, G. L., J. I. VALENCIA, and R. M. Varencia. 1946. Artificial and natural 
hybrids in the Gramineae tribe Hordeae I. Elymus, Sitanion and Agropyron. 
Am. J. Bot. 33:338-350. 

WaceEnaar, E. B. 1959. Intergeneric hybrids between Hordeum jubatum and Secale 
cereale. Jour. Hered. 50:195-202. 


CYTOTAXONOMIC OBSERVATIONS ON MENTZELIA, 
SECT. BARTONIA (LOASACEAE) 


HENRY J. THompPson! 


A previous investigation (Thompson & Lewis, 1955) stated that infor- 
mation about chromosome numbers in Mentzelia would be of great value 
in the formation of evolutionary and taxonomic concepts in the genus. 
Section Trachyphytum, represented by twenty populations of ten species, 
was shown to be a polyploid complex on the base x9, with diploids 
(n=9), tetraploids (n=18), hexaploids (n—27), and octaploids (n=36). 
On the other hand, polyploids were not found in section Bartonza, al- 
though only two species were examined cytologically: Mentzelia multi- 

1 This work received support from a National Science Foundation research grant 


(G18720). The manuscript received the benefit of comments from professors Harlan 
Lewis and Peter H. Raven. 


1963 | THOMPSON: MENTZELIA Ly 


flora with n=9 and M. laevicaulis with n=11. The present paper reports 
chromosome numbers of n=9, 10, and 11, from forty-eight populations 
representing eleven of the approximately twenty species of section Bar- 
tonia, and establishes that aneuploidy, rather than polyploidy, is a major 
feature of evolution in this section. 

Section Bartonia is clearly distinct from all other sections of Mentzelza. 
In section Bartonia the seeds are flat and circular in outline, with the 
margin extended into a wing. The flowers are relatively large and the five 
petals grade into the stamens through petaloid staminodia or at least 
broad-filament stamens. The plants are rosette-perennials and flower in 
late spring or summer. The flowers open and shed pollen in the late after- 
noon or evening and are pollinated by bees or hawk moths. Most of the 
species occur in the Rocky Mountain area of the United States, but one, 
M. albescens, also occurs in Argentina and Chile. This delimitation of 
section Bartonia agrees with that of Urban and Gilg (1900) and also with 
the description of the section by Darlington (1934). Darlington, however, 
without discussion included M. torreyi and M.reflexa in section Bartonia, 
perhaps inadvertently, because she did not modify her description of the 
section to accommodate them. They are clearly more similar to species of 
other sections. 


CHROMOSOME NUMBERS IN BARTONIA 


The nomenclature under which the following chromosome numbers 
are reported largely follows the most recent monograph of the genus 
(Darlington, 1934). The characteristics of the species as understood in 
this study are given briefly so that the chromosome numbers are clearly 
associated with groups of natural populations. All observations of chrom- 
osomes were made in squashed microsporocytes, using procedures previ- 
ously described (Thompson, 1960). Voucher specimens are on file in the 
herbarium of the University of California, Los Angeles (LA). 


MENTZELIA LAEVICAULIS (Dougl. ex Hook.) Torr. & Gray n=11 
Thompson 3089, 3197, 5 miles northwest of Gardiner, Park County, Montana. 
Ne 1690, 35 miles northeast of Garrison, Millard County, Utah. 
3195, 42 miles southwest of Ely, Nye County, Nevada. 
3194, Montgomery Pass, Mineral County, Nevada. 

Mentzelia laevicaulis is widespread west of the Rocky Mountains, 
crossing that range into Montana and Wyoming. It is very distinct, with 
flowers that have five yellow petals 4-6 cm. long. There are no stami- 
nodia, but the five outer stamens have slightly broadened filaments about 
2 mm. wide. The bracts at the base of the capsule are linear to linear- 
lanceolate and entire or with up to four short, pinnate lobes. Over one 
hundred individuals from ten widely scattered populations (see also 
Thompson & Lewis, 1955) have been examined cytologically and all had 
eleven pairs of chromosomes with two of the pairs conspicuously larger 
than the other nine. 


18 MADRONO [Vol. 17 


MENTZELIA DECAPETALA (Pursh) Urban & Gilg n=11 
Thompson 3198, 5 miles northwest of Gardiner, Park County, Montana. 
sy 3206, 1 mile northwest of Florence, Fremont County, Colorado. 
Ernst 746, east of Hennessey, Kingfisher County, Oklahoma. 
Thomtson 3098, 5 miles south of Raton, Colfax County, New Mexico. 
ie 2084, 25 miles southeast of Lubbock, Crosby County, Texas. 
Mentzelia decapetala is widespread east of the Rocky Mountains from 
Idaho and Alberta south to Texas. In habit the plants are much like M. 
laevicaulis, but the flowers have ten white petals (five petals and five 
staminodia ) 5—8 cm. long. The bracts are pectinate and borne on the sides 
of the capsule. None of the chromosomes of M. decapetala is as large as 
the two largest chromosomes in the M. laevicaulis genome. The chromo- 
some number of MW. decapetala was reported as n=11 by Hamel (1938), 
but no voucher specimen was cited. Mentzelia decapetala and M. laevi- 
caulis hybridize readily where they grow together near Gardiner, Mon- 
tana (Thompson 3086, 3087, and 3199). The hybrids are common, easily 
recognized, produce very little good pollen, and set no viable seeds. 


MENZELIA NuDA (Pursh) Torr. & Gray n=10 
Thompson 1800, 2 miles north of Big Springs, Deuel County, Nebraska. 
os 3097, 15 miles south of Fountain, Pueblo County, Colorado. 
mt 1676, 6 miles east of Walsenberg, Huerfano County, Colorado. 
a 2085, 25 miles southeast of Lubbock, Crosby County, Texas. 

Mentzelia nuda occurs on the plains east of the Rocky Mountains from 
Montana to Texas. The plants are 5-10 dm. tall, with the older plants 
producing five or six strict stems from the caudex. The pubescence is 
dense and the leaves are the most scabrous in the section. The white petals 
and petaloid staminodia, about ten in number, grade into the stamens 
through numerous narrow staminodia. The bracts at the base of the cap- 
sule are laciniate. 

The identity of MM. nuda has been a matter of some confusion. Pursh 
(1814, p. 328) based his description on plants grown from seed gathered 
by Nuttall in 1811 along the Great Bend of the Missouri River. Accord- 
ing to Pennell (1936, p. 14) this locality is between the White River and 
the present city of Pierre, South Dakota. Pursh (loc. cit.) says, ““This 
species has smaller flowers, and the leaves are not so glaucous as the fore- 
going |M. decapetala|; in every other respect the above description is 
applicable to the present one... ”’. Considering these remarks and the 
locality, the name can apply only to the species considered here. I con- 
sider M. stricta (Osterh.) Stevens ex Jeffs and Little a synonym. The 
Nuttall specimen cited by Darlington (1934, p. 163) is not the type of 
M. nuda; Pursh apparently never saw it. Most plants cited by Darlington 
(1934, p. 162-3) as M. nuda are referable to M. multiflora. 


MENTZELIA STRICTISSIMA (Woot. & Standl.) Darl. n=10 
Thompson 2081, 4 miles west of Fort Sumner, De Baca County, New Mexico. 
2087, 26 miles southwest of Odessa, Ector County, Texas. 


This species occurs in eastern New Mexico and western Texas, partic- 


1963 | THOMPSON: MENTZELIA 19 


ularly in the drainage of the Pecos River. It is most similar to M. nuda, 
differing only in its smaller flowers (petals less than 25 mm. long) and 
more entire bracts. Mentzelia strictissima is not known to occur sympa- 
trically with M. nuda, and these two taxa may be only geographical races. 


MENTZELIA RUSBYI Woot. n—10 
Thompson 1670, 2 miles north of Eagle Nest, Colfax County, New Mexico. 
i 3103, 18 miles southwest of Las Vegas, San Miguel County, New 
Mexico. 
1665, 6 miles east of Flagstaff, Coconino County, Arizona. 

The plants of the collections cited here are erect, not branched below, 
and the inflorescence is compact. The rosette leaves are linear, about 
15 cm. long, 8 mm. wide, and dentate to nearly entire. The petals are 
15-20 mm. long, white and often with a tinge of apricot at the apex. The 
bracts at the base of the capsule are pinnately lobed. Harrington (1954) 
has considered M. rusbyi and M. nuda to be only varietally distinct |Z. 
nuda (Pursh) Torr. & Gray var. rusbyi (Woot.) Harr.]. 


<6 


MENTZELIA CHRYSANTHA Engelm. n=10 
Thompson 3096, 1 mile north of Florence, Fremont County, Colorado. 
Mentzelia chrysantha was described from plants collected near Canyon 
City, Colorado, from the same vicinity as the collection cited here. The 
plants of this species have numerous lower branches, which are decum- 
bent, and a compact inflorescence. The petals are golden-yellow and the 
leaves are lanceolate and shallowly lobed. The bracts at the base of the 
cylindrical capsules are entire and the seeds are narrowly winged. 


MENTZELIA SPECIOSA Osterh. n=10 
Thompson 3093, 5 miles south of Castle Rock, Douglas County, Colorado. 

The collection here referred to MW. speciosa fits the original description 
very well. The petals are golden-yellow, much like those of M. chrysan- 
tha, but the inflorescence is open, the stems reddish, not white, the leaves 
more deeply lobed, the seeds more broadly winged, and the lower branches 
are fewer and not decumbent. The name M. speciosa has not been used 
since Darlington (1934) placed it in synonomy under M. multiflora 
(Nutt.) Gray; however, the type of M. multiflora is from Santa Fe, New 
Mexico, and has straw-white petals. Furthermore, recent collections from 
the Santa Fe area that match the type of M. multiflora have nine pairs 
of chromosomes. Mentzelia speciosa is apparently restricted to the east 
slope of the Rocky Mountains of Colorado. 


MENTZELIA DENSA Greene n—10 
Thompson 1684, Cotopaxi, Arkansas River, Fremont County, Colorado. 

The plants of the population studied are 3—4 dm. tall with many 
branches from the caudex. The petals are golden-yellow, 16—18 mm. long, 
and grade through staminodia into the stamens. The capsules are cylin- 
dric, tapered at the base, 10-13 mm. long, and subtended by a linear, 
entire bract. The leaves are 4—8 cm. long, pinnate, usually with six lobes, 


20 MADRONO [Vot. 17 


these 4 mm. long and 2 mm. wide, with the midrib 2 mm. wide. These 
plants agree very well with Greene’s (1896) original description. Dar- 
lington (1934) cited only three specimens under MW. densa, all from the 
Grand Junction area of western Colorado, although Greene (1896) stated, 
‘Common in the Canon of the Arkansas in southern Colorado, and else- 
where among the foothills”’. 


MENTZELIA LACINIATA (Rydb.) Darl. n=10 
Thompson 1687, 11 miles west of Pagosa Springs, Archuleta County, Colorado. 
The plants of this population have golden-yellow petals and are gen- 
erally similar to M. densa but differ in having more deeply lobed, lacinate 
leaves and narrowly winged seeds. Mentzelia laciniata has been reported 
from southwestern Colorado and northwestern New Mexico. 


MENTZELIA MULTIFLORA (Nutt.) Gray n=—9 

Thompson 3095, 7 miles north of Penrose, Fremont County, Colorado. 
- 1675, 3 miles west of La Veta Pass Summit, Costilla County, Colo. 
i 3099, 8 miles south of Raton, Colfax County, New Mexico. 
" 3100, 18 miles southwest of Las Vegas, San Miguel County, New 

Mexico. 

ae 1672, 6 miles west of Red River, Taos County, New Mexico. 
s 1669, 7 miles east of Taos, Taos County, New Mexico. 
= 1668, 25 miles southwest of Taos, Taos County, New Mexico. 
i 1667, 38 miles west of Albuquerque, Bernalillo County, New Mexico. 
ce 2076, 15 miles northwest of Datil, Catron County, New Mexico. 
2074, 4 miles east of Quemado, Catron County, New Mexico. 
3107, 6 miles east of Pie Town, Catron County, New Mexico. 
be 2091, 5 miles west of Las Cruces, Dona Ana County, New Mexico. 
eB 2089, 6 miles east of Allamore, Hudspeth County, Texas. 
S 2058, 7 miles east of Pecos, Ward County, Texas. 
: 3212, 1 mile west of Sanders, Apache County, Arizona. 
1066, Highway 66 at Painted Desert, Apache County, Arizona. 
y 3222, 3 miles west of Winkleman, Pinal County, Arizona. 
2094, 21 miles northeast of Benson, Cochise County, Arizona. 
1664, 10 miles southwest of Prescott, Yavapai County, Arizona. 
- 1662, 2 miles northeast of Congress Jct., Yavapai County, Arizona. 
= 2067, Salt River at Highway 60, Gila County, Arizona. 
2066, 10 miles west of Globe, Gila County, Arizona. 

Raven 14818, 13 miles north of Puerto Penasco, Sonora, Mexico. 


Plants of the kind referred here occur in most of the western states 
and in northern Mexico and, as delimited in Darlington’s monograph 
(1934), they form the most variable complex in the section. All the plants 
in this group have petals 15-25 mm. long, yellow to nearly white, that 
grade through broad staminodia into the stamens. The capsules are cylin- 
dric to urceolate, 15-25 mm. long, usually with a linear, entire bract at 
the base and with seeds that are broadly winged. The leaves are pinnately 
lobed to dentate. Mentzelia multiflora (Nutt.) Gray obviously applies to 
this group of plants. The type was collected by Gambel near Santa Fe, 
New Mexico, and was described as having petals 34 inch long, straw- 
white in color. My collections from this region in New Mexico that agree 
with the original description have nine pairs of chromosomes. Mentzelia 


1963 | THOMPSON: MENTZELIA 21 


nuda (Pursh) Torr. & Gray is sometimes incorrectly applied to this group 
of plants. 

Four of the collections cited here, Thompson 1664, 1662, 2067, and 
2066, differ from the others in having the darkest yellow petals, the most 
urceolate capsules, and more entire leaves. Plants of this kind grow in the 
mountains of central Arizona between the southern desert and the Mogol- 
lon Rim. 


MENZELIA MULTICAULIS (Osterh.) Darl. n=—I11 
Thompson 3201, Dinosaur National Monument, Uinta County, Utah. 
a 3203, 8 miles west of Rio Blanco, Rio Blanco County, Colorado. 
x 3205, 1 mile north of Walcott, Eagle County, Colorado. 

Mentzelia multicaulis occurs in northwestern Colorado and northeast- 
ern Utah. The flowers have five yellow petals, but the next inner whorl 
may be either five staminodia or five stamens with filaments nearly as 
broad as the petals. A distinctive characteristic of M. multicaulis is that 
either the petals, the staminodia, or both are very broad, 9 mm. long and 
6 mm. wide. In this species the capsules are urceolate, 8 mm. long and 
subtended by a linear bract. The seeds have very narrow wings. The lower 
leaves are 4—6 cm. long, pinnatifid, the four to six lobes are nearly 1 cm. 
long and 2 mm. wide and the midrib is 3 mm. wide. The upper leaves are 
entire and linear. All of the plants observed were more than one year old 
and had several stems 1—3 dm. long from the old caudex. Mentzelia multi- 
caulis is most similar to M. humilis (Gray) Darl., but the latter differs in 
having the petals and staminodia short and narrow and seeds with broad 
wings. 

Chromosome numbers have been reported for two other species of sec- 
tion Bartonia. Mentzelia albescens (Gill.) Griseb., the only species of 
section Bartonia in South America, was reported as n=11 by Covas and 
Schnack (1946) from material collected at Lujan, Provincia de Mendoza, 
Argentina. Hamel (1938) reported n=9 from root-tip sections of plants 
identifid by him as M. humilis. Hamel did not cite a voucher specimen 
and the identity of his material is unknown. The seeds for his culture 
were obtained from the New York Botanical Garden. 


SUMMARY AND CONCLUSION 


Two sections of Mentzelia, sections Bartonia and Trachyphytum, are 
morphologically more similar to each other than to any of the other sec- 
tions. However, their validity as distinct phylads within the genus is indi- 
cated not only by consistent morphological differences but also by con- 
trasting patterns of chromosomal change in the course of their evolution. 
Section Trachyphytum has only one basic chromosome number (x=9) 
although it comprises diploid, tetraploid, hexaploid, and octaploid species 
that form one large polyploid complex. Section Bartonia, on the other 
hand, has no known polyploid species, but three basic chromosome num- 
bers are present in the group; one highly variable species has n=9, seven 


De MADRONO [Vol. 17 


species have n= 10, and four species have n=11. The prevalence of poly- 
ploidy in one section and aneuploidy in the other undoubtedly reflects a 
fundamental difference in genetic systems and a corresponding phylo- 
genetic separation of the two sections. 


Department of Botany, 
University of California, Los Angeles 


LITERATURE CITED 

Covas, G., and B. ScHNAcK. 1946. Numero de cromosomas en antofitas de la Region 
de Cuyo (Republica Argentina). Rev. Arg. Agron. 13:153-166. 

DARLINGTON, JOSEPHINE. 1934. A monograph of the genus Mentzelia. Ann. Missouri 
Bot. Gard. 21:103-226. 

GREENE, E. L. 1896. New or noteworthy species. Pittonia 3:99. 

Hamet, J. L. 1938. Etude de la mitose somatique et numération chromosomique 
chez quelques Loasacées. Rev. Cyt. Cytophys. Vég. 3:153. 

Harrincton, H. D. 1954. Manual of the plants of Colorado. Denver, Sage Books. 

PENNELL, F.W. 1936. Travels and scientific collections of Thomas Nuttall. Bar- 
tonia 18:1-51. 

PursH, F. 1814. Flora Americae Septentrionalis. Vol. I. 

Tuompson, H. J. 1960. A genetic approach to the taxonomy of Mentzelia lindleyi 
and M. crocea (Loasaceae). Brittonia 12:81-93. 

Tuompson, H.J., and H. Lewis. 1955. Chromosome numbers in Mentzelia (Loasa- 
ceae). Madrono 13:102-107. 

Urpan, I., and E. Gitc. 1900. Monographia Loasacearum. Nova Acta Abh. Kaiserl. 
Leop.-Carol. Deutsch. Akad. Naturf. 76:22-97. 


A REVISION OF THE GENUS THAXTEROGASTER SINGER 
RotF SINGER AND ALEXANDER H. SmitH! 


Since the original publication of the genus Thaxterogaster (Singer, 
1951) and the subsequent monograph (Singer & Smith, 1958), further 
studies have been made as time and available material permitted, and 
these have led to the discovery of additional species. Species are here 
grouped into sections, since certain distinct trends are now evident. Those 
species treated in detail in our monograph (doc. cit.) are not redescribed 
here, but detailed accounts of new or transferred taxa are treated critically. 

We wish to acknowledge assistance from the National Science Founda- 
tion, which made possible the studies at the Royal Botanic Gardens, Kew, 
England. We also express our thanks to Dr. G. Taylor, Director of the 
latter institution, for the privilege of studying the collections there, and 
to Dr. Clark Rogerson, Curator, New York Botanical Garden, New York, 
for the opportunity to study the collections of Hymenogaster at that 
institution. 


1 Papers from the Facultad de Ciencias Exactas y Naturales, Universidad de 
Buenos Aires, Buenos Aires, Argentina, and the University Herbarium and Depart- 
ment of Botany (Paper no. 1142), the University of Michigan, Ann Arbor, Michi- 
gan, U.S.A. 


1963 | SINGER & SMITH: THAXTEROGASTER 23 


KrEy TO SECTIONS OF THAXTEROGASTER 


A. Clamp connections present on hyphae of the gastrocarp. 
B. Spores small (rarely up to 13 uw long), either smooth or with an adhering ver- 


ruculose-rugulose exosporial ornamentation . . Sec. 4. Micros porogaster 

B. Spores larger and always distinctly ornamented. 
C. Spores angular when immature, subglobose . . . . Sect. 2. Blestogaster 
C. Immature spores not angular... . . . . . Sect. 3. Thaxterogaster 


A. Clamp connections absent. 
D. Spores with an apical beak and the wrinkled outer layer of the wall loosen- 


ing to a considerable extent . . . . . . Sect. 1. Scabrogaster 
D. Spores lacking a distinct apical beak anil the exosporial ornamentation adher- 
ing to the inner wall eee a ee ect. SA poy porasce, 


Sect. 1. Scabrogaster Singer & Smith, sect. nov. 


Sporis subrostrato-mucronatis, ornamentatione subrugulosa separabili. 
Typus. Thaxterogaster scabrosum (Cooke & Massee) Smith & Singer. 


1. Thaxterogaster scabrosum (Cooke & Massee) Smith & Singer, 
comb. nov. Secotium scabrosum Cooke & Massee, Grevillea 20:35. 1891. 

Peridium hemispherical, depressed, dingy-olive or grayish, minutely 
scabrid. Gleba lacunose, septa gill-like, waved and folded, dark reddish 
brown. Stipe very short, almost obsolete. 

Spores 15-18 (20) & 7.5—10 », dark rusty brown in KOH, near yellow- 
ish tawny in Melzer’s reagent; with a slightly wrinkled outer layer which 
tends to loosen variously, the plage area not differentiated; apex tapered 
to a blunt-pointed beak-like projection but this consisting entirely of wall 
material (no pore present). 

No other microscopical details obtainable from type. 

It is clear from the type, the original description, and Smith’s observa- 
tions on the spores that this species is a Thaxterogaster connecting that 
genus to the group of species in Hymenogaster sensu lato with spores hav- 
ing an outer wrinkled wall and an apical beak. This group in Hymeno- 
gaster makes up the largest element in that genus of diverse spore types. 

Type studied. “On the ground. Domain, Melbourne (Baron Muel- 

en > (Ke). 


Sect. 2. Blestogaster Singer & Smith, sect. nov. 


Sporis subglobosis, juventute plus minusve angulosis. 

Typus. Thaxterogaster brevisporum Singer. 

2. THAXTEROGASTER BREVISPORUM Singer, Persoonia 1:386. 1960. 

We emphasize the angularity of the spores because this feature may be 
of some importance in connecting the species to members of Hymeno- 
gaster sensu lato. 


Sect. 3. THAXTEROGASTER. 

Spores ellipsoid, over 13 » long, never angular, not with a distinct apical 
beak, the outer layer of spore wall adherent to inner layer; hyphae hav- 
ing clamp connections at the septa. 


24 MADRONO [Vol. 17 


Key TO SPECIES OF SECTION THAXTEROGASTER 


A. Peridium lacking a thick gelatinous epicutis. 
B. Peridium pallid, becoming brownish . . . . . . . 6. T. magellanicum 
B. Peridium violaceous. 
C. Spores mostly 15-18 » long; stipe poorly developed; growing under Notho- 


fagus pumilo .. 1 + © = 6 Gene ae ee ee (on Euiolacemn 
C. Spores mostly 13-15 » long; stipe well developed; growing under Nothofagus 
dombeyi and Saxegothaea .......... . .. 4.7. dombeyti 

A. Peridium with a thick gelatinous epicutis. 
D. Peridium white ftp i ad 3 SL GR 2A SN ie lencoce pial 
D. Peridium olive yellow to bister . . . ... .: =. =. =. «. 5. LT. pingue 


3. THAXTEROGASTER VIOLACEUM Singer, Mycologia 43:216. 1951, and 
BrittonvalOe 20/1958. 

4. THAXTEROGASTER DOMBEYI Singer, Persoonia 1:385. 1960. 

5. THAXTEROGASTER PINGUE (Zeller) Singer & Smith, Brittonia 10: 
211. 1958. Secotium pingue Zeller, Mycologia 33:211. 1941. 

6. THAXTEROGASTER MAGELLANICUM Singer, Mycologia 43:219. 1951, 
and Brittonia 10:208. 1958. 

7. THAXTEROGASTER LEUCOCEPHALUM (Massee) Singer & Smith, Brit- 
tonia 10:210. 1958. Secotium leucocephalum Massee, Grevillea 19:95. 
1891. 

The following data are taken from the holotype at Kew: Peridium 
duplex, the outer layer a thick gelatinous layer of appressed hyaline nar- 
row hyphae with clamp connections. Inner layer of floccose broader 
(5-12 ») hyphae. Tramal plates of interwoven hyphae yellowish in KOH 
(but not reviving well and apparently not gelatinous). Details of hyme- 
nium not obtained. Spores 12-16 & 7-8 mw, warty rugulose and rusty 
brown in KOH (as in Cortinarius). 

These data on the holotype verify the characters of the species as pub- 
lished previously by us (loc. cit.). 


Sect. 4. Microsporogaster Singer & Smith, sect. nov. 


A sectione Thaxterogastero sporis minoribus, interdum sublevibus 
differt. 
Typus. Thaxterogaster subalbidum Smith. 


Key TO SPECIES OF SECTION MICROSPOROGASTER 


A. Spores almost smooth; stipe-columella solid . . .. . . 8.7. subalbidum 
A. Spores verruculose-rugulose; sterile base chambered, columella hollow 
(at least when dried) fe Ss gee Shas DAMEN ee aba cap as Om wae TIE 


8. Thaxterogaster subalbidum Smith, sp. nov. 

Sporis sublevibus; stipite columellaque solidus. 

Typus. Thaxter, Fungi Hypogeus No. 4, March 5, 1906, Punta Arenas, 
Chile, South America (FH). 

Gastrocarp 1-2 cm. broad (estimated on basis of dried material), ir- 
regularly globose to convex, the surface silky and white, drying whitish; 
eleba dingy cinnamon brown, of minute chambers nearly filled with 


1963 | SINGER & SMITH: THAXTEROGASTER 23 


spores; stipe-columella well developed, solid, percurrent, the surface pal- 
lid, the interior slightly darker, extending well below the lower margin 
of the peridium, the estimated length 1—-1.5 cm., width 5-7 mm., the 
peridium either connected to the stipe-columella or free in places to expose 
slightly the gleba. 

Spores 10-13 6-8 uy, ellipsoid with a short-pointed, inconspicuous 
sterigmal appendage, the wall 1.3—2 » thick (as seen on fractured spores), 
the surface very minutely warty, wrinkled to punctate-roughened (almost 
smooth as seen under high dry), pale cinnamon in KOH; no apical dif- 
ferentiation. 

Basidia 28-33 8-10 p, clavate, 4-spored, hyaline in KOH. Cystidia 
none. Basidioles resembling immature basidia. Tramal plates with an 
indistinct subhymenium of narrow branched elements intergrading with 
the central area which is of interwoven filaments, some with somewhat 
enlarged cells, hyaline in KOH. Epicutis of peridium a thick layer of 
appressed, narrow (2-3 »), hyaline, smooth, thin-walled hyphae; this 
layer grading imperceptibly with the context which has broader (4—7 pz) 
hyphae, some oleiferous hyphae, and is slightly yellowish in KOH; clamp 
connections present. 

The nearly smooth spores separate Thaxterogaster subalbidum from 
the other known Thaxterogasters. In the material in the Zeller collec- 
tions (NY), a duplicate of the packet of specimens cited above is on the 
same sheet as the type of Hymenogaster fragilis. The description of the 
latter, however, obviously does not apply to ‘“‘Fungi Hypogeus No. 4,” 
which we here designate as the type of 7. subalbidum. Material of Hy- 
menogaster fragilis sent to Zeller, apparently by Thaxter, and in the 
Zeller collections, is marked with red crayon as an indication that it is 
the holotype of H. fragilis, but in the original description of that species 
the location of the “type” is given as the Farlow Herbarium. We do 
not know whether Zeller saw all the material or not. The holotype of 
T. subaibidum is that portion of “Fungi Hypogeus No. 4” deposited in 
the Farlow Herbarium. 


9. Thaxterogaster fragile (Zeller & Dodge apud Dodge & Zeller) 
Smith & Singer comb. nov. Hymenogaster fragilis Zeller & Dodge apud 
Dodge & Zeller, Ann. Missouri Bot. Gard. 21:646. 1934. 


Gastrocarp 1—2 cm. diam., subglobose to pear-shaped, the surface whit- 
ish or gray, drying pallid to pale cinnamon-buff; gleba chambered, fragile, 
the cavities fairly large for a fungus of that size and very irregular in 
shape, bright cinnamon as dried; stipe-columella consisting of a cham- 
bered sterile base, 6 mm. broad (reminding one of the marginate bulb in 
a Cortinarius), this extended into a hollow thin-walled percurrent colu- 
mella which when dried may be obliterated entirely except for the cavity; 
gleba attached to the length of the columella (if latter is still evident) ; 
peridium even, silky, extending to the stipe-columella (but possibly slight- 
ly free at times). 


26 MADRONO [Vol. 17 


Spores 10-12  7-8.5 », ovate-pointed and with a short, pointed sterig- 
mal appendage, dull cinnamon brown in KOH, thick-walled, warty- 
rugulose, the outer wrinkled layer not separating appreciably from the 
inner one; apical beak solid (lacking a pore). 

Peridium with a surface layer of indefinite thickness of appressed fila- 
ments 3-6 « diam., hyaline in KOH, smooth, thin-walled and with medal- 
lion clamps at cross walls. This layer intergrading with the context which 
is composed of broader hyphae and some laticifers (?). Details of hyme- 
nium and tramal plates not evident. 

Punta Arenas, Chile, March 1906. R. Thaxier. Word “type” under- 
lined in red on the packet in the Zeller collections (NY). 

The description of Zeller and Dodge obviously applies to this material, 
which is distinct from the March 5, 1906, collection (i.e., Thaxterogaster 
subalbidum) by the brighter cinnamon, more fragile gleba, ovate-pointed 
to almost broadly fusoid conspicuously roughened spores, chambered 
sterile base and hollow columella. 

Parks 2182, filed in the Zeller collections (NY) as Hymenogaster 
fragilis, is not the same as either of the Thaxter collections. The spores 
and the microscopic characters of the peridium are those of the type of 
Hymenogaster gilkeyae, but all the details of the hymenium and tramal 
plates had collapsed beyond the point of reviving. No clamps were pres- 
ent as far as could be observed. 

Sect. 5. Aporpogaster Singer & Smith, sect. nov. 

Hyphis defibulatis, sporis haud vel vix rostrato-apiculatus; strato 
externo sporarum haud separabili. 

Typus. Thaxterogaster conicum (Hesler) Singer & Smith. 


KEY TO SPECIES OF SECTION APORPOGASTER 
A. Gastrocarp globose, depressed; spores 7-llu broad . . . 10.7. porphyreum 
A. Gastrocarp typically conic and elongate; spores mostly 11-13 uw broad. 
11. T. conicum 

10. THAXTEROGASTER PORPHYREUM (Cunningham) Singer, Lilloa 26: 
105. 1953; see also Brittonia 10:212. 1958. Secotium porphyreum Cun- 
ningham, Proc. Linn. Soc. N.S. W. 49:114. 1924. 

11. THAXTEROGASTER CONICUM (Hesler) Singer & Smith, Brittonia 10: 
214. 1958. Secotium conicum Hesler, Jour. Elisha Mitchell Soc. 49:153. 
1933. 


Facultad de Ciencias Exactas y Naturales 
Buenos Aires, Argentina 


Department of Botany 
University of Michigan 
Ann Arbor, Michigan 


LITERATURE CITED 
SINGER, R. 1951. Thaxterogaster—a new link between gasteromycetes and Agaricales. 
Mycologia 43:215-288. 
SINGER, R., and A. H. SmitH. 1958. Studies on secotiaceous fungi I: a monograph 
of the genus Thaxterogaster. Brittonia 10:201-216. 


1963 | SPEESE & BALDWIN: HYMENOXYS ZY. 


CYTOPHYLETIC ANALYSIS OF HYMENOXYS ANTHEMOIDES 
BERNICE M. SpEESE and J. T. BALDWIN, JR. 


This paper reports the chromosomal number for Hymenoxys anthe- 
moides (Juss.) Cass., type species of the genus, and relates this datum to 
the chromosomal situation as known for Hymenoxys. The analysis is cyto- 
phyletic, in the original meaning of the word (Baldwin, 1939). 

Parker (1950) published certain new combinations in Hymenoxys 
Cass. that she later (1951, in manuscript) used in a monograph of this 
genus. Earlier, she had asked us to survey the chromosomes of Hymen- 
oxys and had supplied us with thirty-five collections of seeds representa- 
tive of both subgenera, of fourteen of the twenty-four species recognized 
by her, and of five varieties. 


Fic. 1. Specimens of Hymenoxys anthemoides grown from Argentine seed (Bald- 
win 15580). Six inch scale shown. 


28 MADRONO [Vol. 17 


Speese and Baldwin (1952) published the results of their studies of 
Hymenoxys: twelve species had 2n numbers of 30; H. acaulis (Pursh) 
Parker had 2n numbers of 60 in three varieties and of 30 in var. ivesiana 
Greene; H. odorata DC. had a 2n number of 22. Parker (1960) used 
chromosomal evidence and stem and leaf characters as bases for raising 
H. acaulis var. ivesiana to specific rank: H. ivestiana (Greene) K. F. 
Parker. 

Jackson (1957) reported an n number of 15 for H. argentea (A. Gray) 
Parker and thus substantiated our count of 2n of 30 for the species. Tur- 
ner, Beaman, and Rock (1961) published n numbers of 15 for two species 
of Hymenoxys from Nuevo Leon, Mexico: A. insignis (A. Gray) CKIL., 
a species of tall biennials not investigated by us and most closely related 
to H. grandiflora (Torr. & Gray) Parker for which we had found that 
2n=30; and H. odorata (Rock 264, TEX), annuals, for which we had 
reported a 2n number of 22 from New Mexico (Parker and McClintock 
7009, US) and Arizona (K. Parker 7459, US). For plants of H. odorata 
from almost the same Arizona station (P. Raven 11731, UC), Raven and 
Kyhos (1961) also determined an n number of 11: they thus corroborate 
our count for the species. In addition to the count of n—=15 for the Nuevo 
Leon material of H. odorata (Rock 264) reported in the 1961 paper, 
Turner obtained a similar count (fide Johnston) for a second Nuevo 
Leon collection (M. C. Johnston 5860). 


ee 0, MahOgeedpagagy 


Z 2, 


Fic. 2. Chromosomes of Hymenoxys anthemoides (the plants grown from Argen- 
tine seed): left, mitotic metaphase, 2n=30; right, metaphase I of meiosis, n=15. 
Chromosomes were drawn X 2200 and reduced by one-third in reproduction. 


Dr. Parker (letter of August 31, 1961) wrote us that all five specimens 
cited above are typical H. odorata. It is clear that the chromosomal sit- 
uation in this species needs further study. The chromosome counts re- 
ported in Turner, Beaman, and Rock (1961) are from pollen-mother- 
cell smears of buds fixed in the field during the summer of 1959. Our ex- 
perience has been that preparations from material so fixed are often dif- 
ficult to interpret, and especially so if the weather were hot at the time 
of fixation. 

Parker (1951) stated that H. odorata, a widely distributed annual in 
the Midwest and Southwest, is closely allied both to the Mexican H. 


1963 | SPEESE & BALDWIN: HYMENOXYS 29 


chrysanthemoides DC. and to the South American H. anthemoides ( Juss.) 
Cass., type species for the genus, but that these three annuals are quite 
distinct. Two other species are in South America: one is annual; the 
other, annual or biennial. Speese and Baldwin (1952) wrote: “If it 
should be discovered that the South American species—and especially 
the type species—fall into a chromosome series with H. odorata |2n=22: 
an annual] or into any series different from that evidenced by the ma- 
jority of Hymenoxys representatives as interpreted by Parker |2n=30 
or 60: mostly perennials], reason would then be at hand to suspect the 
validity of Parker’s treatment.” 

Parker, awaiting additional chromosomal data on Hymenoxys, has de- 
layed publication of her monograph, and both she and we have made a 
number of attempts to obtain viable seeds of South American species. 
Finally, on January 15, 1960, Dr. Arturo Burkart most kindly collected 
fruiting specimens of H. anthemoides and sent them to Dr. Parker, who 
placed a voucher specimen in the United States National Herbarium, 
Smithsonian Institution, and sent us plants with mature achenes. We 
grew seedlings (Baldwin 15580, US, fig. 1) and examined their chromo- 
somes: H. anthemoides has a 2n number of 30, an n-number of 15 (fig. 2). 

In summary, the basic chromosome number of sixteen of twenty-five 
species (twenty of thirty-one taxa) accepted by Parker (1951, 1960) for 
Hymenoxys and including the type species is 15; the plants are either di- 
ploid or tetraploid. The basic number for H. odorata is 11. (We assume 
the report of n—15 for this species to be incorrect.) These numbers indi- 
cate phyletic trends. From chromosomal evidence alone, one would con- 
clude that H. odorata is wrongly placed in this species. The chromosomal 
data for the other species, however, lend support from one more discipline 
to Dr. Parker’s interpretation of Hymenoxys. 


College of William and Mary, 
Williamsburg, Virginia. 


LITERATURE CITED 


BaLtpwin, J. T., JR. 1939. Certain cytophyletic relations of Crassulaceae. Chron. Bot. 
5:415-417. 


Jackson, R. C. 1957. Documented chromosome numbers of plants. Madrofo 14: 
111-112. 


PARKER, Kittie F. 1950. New combinations in Hymenoxys. Madrono 10:159. 
1951. Monograph of the genus Hymenoxys Cass. Unpublished manuscript, 
pp. 1-112. 
1960. Two species of Hymenoxys (Compositae) new for Arizona. Leafl. 
West. Bot. 9:92-93. 
Raven, P. H., and D. W. Kyuos. 1961. Chromosome numbers in Compositae: II. 
Helenieae. Am. Jour. Bot. 48:842-850. 
SPEESE, BERNICE M., and J. T. Batpwin, Jr. 1952. Chromosomes of Hymenoxys. 
Am. Jour. Bot. 39:685-688. 
Turner, B. L., J. H. BEAMan, and H. E. L. Rock. 1961. Chromosome numbers in 
the Compositae. V. Mexican and Guatemalan species. Rhodora 63:121-129. 


30 MADRONO [Vol. 17 


REVIEWS 


Native Orchids of Trinidad and Tobago. By RicHarp Evans SCHULTES. 275 pp., 
25 photographs, 74 drawings, 1 map. Pergamon Press, 1960. $15. 


Until recently the determination of orchids has often proved a difficult task. The 
literature pertaining to the 20,000 or so species is naturally voluminous, much of it is 
widely scattered and difficult of access, and in consequence most identifications have 
to be carried out by specialists at such large orchid herbaria and libraries as those at 
Harvard University. Within the last decade, however, a number of floristic treat- 
ments of the Orchidaceae have appeared, including those for the United States, 
Mexico, Guatemala, Venezuela, Brazil, Peru, and Malaya; now the general botanist 
and amateur student are much better equipped to work with this fascinating family. 

Dr. Schultes, Curator of the Botanical Museum of Harvard University, has pre- 
pared an excellent account of the orchids of Trinidad and its companion island, 
Tobago. The subject of this meticulously written and well-illustrated treatise is re- 
stricted to an area of less than 2,000 square miles, but the book will be of more 
than local interest because a large number of the species are found elsewhere in Latin 
America. Trinidad and Tobago are not true islands in the geologic sense, being non- 
volcanic in origin. Trinidad is only seven miles from the coast of Venezuela and 
endemism is low—7 per cent; as is well-known, on islands never connected with the 
mainland endemism is often very high (e.g., 75 per cent in Hawaii). The flora of 
Trinidad is also distinctly South American, rather than Caribbean, and this applies 
as well to the orchids. 

Schultes recognizes 181 species of Orchidaceae, two less than are found in the 
Gramineae, the largest family on the islands. Included in each generic description 
are characters found in the entire geographic range, whereas the species descriptions 
apply only to plants from the two islands. Descriptions and citations of specimens 
are very complete, and there are also presented such useful data as the flowering 
times, the etymological origin of the generic names, and the citation of notes pub- 
lished elsewhere. 

Most of the photographs are undistinguished, but are adequate for purposes of 
identification. Unfortunately, the one of Pleurothallis diffusa appears again on an- 
other page mislabeled Stelis muscifera. Quite outstanding are the drawings, many 
of which are reprinted from other recent floristic works on orchids. By such well- 
known artists as Gordon Dillon and G. Dunsterville, they add much to the value 
of the book. 

This work can be especially recommended as an introduction to a representative 
selection of orchid taxa, many of which are widespread in the Americas——Mvyron 
KimnacH, Botanical Garden, University of California, Berkeley. 


Aquatic Plants of the Pacific Northwest with Vegetative Keys. By (late) ALBERT 
N. Steward, La Rea Dennis, and HeLten M. Girxey. Illustrated. Oregon State 
College Studies in Botany, No. 11. Oregon State College Press, Corvallis. v + 184 pp. 
1960. $2.50. 


This manual was intended to cover “all true aquatics and also species whose life 
cycle includes some stage that requires either the saturation of the substrate with 
water or the presence of an ambient aqueous medium” occurring in Oregon, Wash- 
ington, British Columbia, and Alaska. The authors have not succeeded in doing 
this, but they have produced a manual with some merits which will be useful once 
its limitations are recognized. 

The text covers hepatics, mosses, vascular cryptogams, and angiosperms. Species 
found both in moist and dry areas were to be excluded. Main emphasis in the keys 
is on vegetative characters, although reproductive structures are used either secon- 
darily, or when vegetative traits fail. Because of the generous ecological qualifications 
given above, one cannot argue with the inclusion of Alnus oregona, Urtica lyallii, 
Physocarpus capitatus, Oenothera hookeri, or Gaultheria shallon in this manual, 


1963 | REVIEWS Spl 


although they would not be considered aquatic plants by many botanists. The 
incorporation of such doubtful species should not be troublesome to users of the 
manual. It is wiser for a work covering plants of a specific habitat to be over- 
generous than to be parsimonious. In contrast, however, a number of genera with a 
legitimate claim to inclusion have been omitted. Among these are Elatine, Lindernia, 
Camassia, Habenaria, Porterella, Limnanthes, Limosella, Navarettia, Tillaea, Berula, 
Sphenosciadium, and Lilaeopsis. Some of these genera are widespread and are likely 
to be picked up by persons collecting in the wet lands of the Pacific Northwest. 
Furthermore, in some genera such as Hydrocotyle, Cicuta, Eryngium, and Lobelia, 
not all species growing in moist areas are given, which increases the risk of mis- 
identification in these genera. I have not found any genera or species represented 
in Alaska, British Columbia, or Washington but not in Oregon in this manual, 
suggesting that it will have limited use outside the state. 

On the credit side, particularly noteworthy are the vegetative keys, especially 
to groups such as the Cyperaceae, which frequently do not flower for long periods 
of time and are difficult to identify when they do. Many of the vegetative distinc- 
tions between taxa will be of interest since these are often passed over in floras in 
faver of differences in the reproductive organs alone. 

This manual cannot be recommended for use in identification unless it is used 
in conjunction with the other manuals available for Oregon and Washington. It has 
some serious drawbacks which could easily be eliminated in a second edition by 
rewording the title, the preface, and by expanding the coverage to include those 
important groups which have been omitted—RoBERT ORNDUFF, Department of 
Biology, Reed College, Portland. 


A Flora of the Alaskan Arctic Slope. By Ira L. WicctIns and JOHN HUNTER 
THOMAS. vii + 425 pp. 1962. Univ. Toronto Press, Canada. $9.50. 


This book is a comprehensive manual of vascular plants, including complete keys, 
descriptions of all taxa, and distribution maps. It is based largely on recent exten- 
sive collections by the authors and others. The field, herbarium, and library work 
was supported largely by the Office of Naval Research and the Arctic Institute of 
North America. The taxonomic treatment generally follows Hulten’s and Anderson’s 
floras of Alacka, but it includes some original taxonomic interpretations, takes into 
account recent taxonomic papers, and cites many more collection localities, including 
range extensions to the American Arctic for numerous species. 


The authors have designed their book for use as a handbook for field identifica- 
tion as well as a technical reference work for professional taxonomists. The omission 
of synonymy and illustrations is unfortunate, as is the failure to cite distribution 
records from Hulten’s “Flora of Alaska” and from collections other than those 
listed by the authors. These deficiencies are more than balanced, however, by such 
features as complete keys and descriptions, helpful comments on taxonomic problems 
in many taxa, a glossary, a short bibliography, and detailed distribution maps ac- 
companied by a valuable gazetteer. 


The habitat notes and the sections on the environment contribute toward the 
integration of taxonomy and ecology. The essay on “minor habitats” (7.e., those dis- 
turbed by man) emphasizes dramatically “the serious nature of comparatively minor 
disruptions of the normal balanced environmental and ecological conditions” in the 
Arctic. The vastness and remoteness of the Arctic tend to make people in general 
overlook this fact, but it is all too evident to the visitor to the extensively disturbed 
Point Barrow region. On the other hand, the American Arctic presents unsurpassed 
and largely unexplored opportunities for detailed study of the effects of ecological 
disturbance in a region where man has as yet effected only very local changes. The 
new flora of the Arctic Slope will be a valuable tool in the hands of biologists fol- 
lowing this as well as many other lines of investigation—S. GaLEN SmitTH, Depart- 
ment of Botany and Plant Pathology, Iowa State University, Ames. 


32 MADRONO [Vol. 17 


The Savory Wild Mushroom. A Pacific Northwest Guide. By Marcaret Mc- 
Kenny (with the collaboration of D. E. Stuntz). xiv + 133 pp., 1 text-figure, 33 
black-and-white and 48 color photographs. University of Washington Press, Seattle. 
1962. $3.95. Paperback. 


One of the most welcome aspects of this attractive handbook of mushrooms is 
the inclusion of a brief, up-to-date chapter on mushroom poisons, written by Dr. 
Varro E. Tyler, Jr., Professor of Pharmacognosy, University of Washington. For this 
reason, this book should be of particular interest to physicians; it provides a con- 
venient source of information on types of poisons, symptoms, and treatment, as well 
as descriptions and illustrations of the most dangerous of the poisonous mushrooms. 
Although subtitled “A Pacific Northwest Guide,” this handbook should be of interest 
to any collector, because it includes a large number of more or less cosmopolitan 
species. 

Following a brief introduction to the nature of mushrooms and instructions for 
their collection, the species are organized into three groups—edible mushrooms, 
poisonous mushrooms, and nonpoisonous mushrooms to be avoided. Most of the 
common edible and poisonous species are included. Enough precautions about col- 
lecting, determining, and eating are mentioned to make it possible to recommend this 
book, with no serious reservations, to the amateur collector as a reliable handbook. 
For the gourmet, there is a chapter of tantalizing recipes for cooking mushrooms.— 
ISABELLE TAVARES, Department of Botany, University of California, Berkeley. 


NOTES AND NEWS 


The year 1962 marked the passing of two noted botanists, John James Thornber, 
who died in Tucson,. Arizona, on November 22 at the age of ninety, and Joseph F. C. 
Rock, who died in Honolulu, Hawaii, on December 5 at the age of seventy-nine. 
Dr. Thornber, who joined the staff of the University of Arizona in 1901, was a bota- 
nist of international repute and was Dean of the College of Agriculture at the Uni- 
versity of Arizona from 1922 to 1928. His intimate knowledge of the Arizona flora 
has not been surpassed. Dr. Rock was Professor of Botany at the University of 
Hawaii from 1911 to 1919. He spent many years exploring in remote parts of south- 
western China. In addition to his contributions as a plant explorer, he was a noted 
linguist, cartographer, anthropologist, and ornithologist. Many of his plant discoveries 
were introduced into cultivation through the University of California Botanical 
Garden. His beautifully prepared specimens are deposited in herbaria throughout the 
world. 


The Segundo Congreso Mexicano de Botanica, organized by the Sociedad Botdnica 
de México, will be held in San Luis Potosi from September 17 to 20, 1963. Queries 
concerning the Congress may be addressed to Biol. Fernando Medellin, Apartado 
Postal 458, San Luis Potosi, S.L.P., México. The highly successful first Congress was 
held in Mexico City in October, 1959. Among the 172 participants were 26 from the 
United States. 


NOTICE TO CONTRIBUTORS. With my retirement from the University 
of California in June, 1963, I am also retiring from the Editorial Board of 
Madrono, a post that I have held for twenty-nine years. In anticipation of 


this time, I am asking all contributors to send new manuscripts to Dr. John 
H. Thomas, Division of Systematic Biology, Stanford University, California. 
Dr. Thomas will take over the duties of the editorship beginning with the 
July, 1963, issue of Madroho.—HErBeErtT L. Mason, Chairman, Editorial Board. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Second Edition, 1954). 

Articles may be submitted to any member of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume I, 1916-1929 . . . . . $5.00 
Volume IT, 1930-1934... . .  . . » 5:00 
Volume IIT, 1935-1936 Mae ee Bee ag 0.0) 
Volume IV, 1937-1938. . . . . 5.00 
Volume V, 1939-1940 . . . . . 5,00 
Volume VI, 1941-1942 ge genet hy ESTO 
Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 oo Me al a 1GKOO 
Volume X, 1949-1950 . . . . . = 7.00 
Volume XI, 1951-1952 rhe rae 2 3)4 0,0, 
Volume XII, 1953-1954 . . . . 8.00 
Volume XIII, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 et a OOO 
Volume XVI, 1961-1962 . . .  . 12.00 


Single numbers: 


Volumes: andll 2.0% Soe 50 
Volumes II through XIV... .. 1.00 
Volumes V" 5 3 “2s | heap ae 1.25 
Volume XVI. (2 «vale eee 1.50 


The subscription price of MaproNo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


ADRONO 


VOLUME 17, NUMBER 2 APRIL, 1963 


Contents 


PAGE 


QUATERNARY CLOSED-CONE PINE FLORA FROM TRAVER- 
TINE NEAR LITTLE Sur, CALIFORNIA, Jean H. Langen- 
heim and J. Wyatt Durham 33 


CHROMOSOME NUMBERS OF SOME PHYTOGEOGRAPHICAL- 
LY INTERESTING CHILEAN Piants, D. M. Moore 52 


CHROMOSOME COUNTS IN SECTION ERYTHRANTHE OF 
THE GENUS MIMULUS (SCROPHULARIACEAE). II., 
Robert K. Vickery, Jr., Barid B. Mukherjee and 
Delbert Wiens 53 


AN ANALYSIS OF VARIATION IN VIOLA NEPHROPHYLLA, 
Norman H. Russell and Frank S. Crosswhite 56 


Review: H. H. Allan, Flora of New Zealand 
(Robert Ornduff) 66 


NOTES AND NEws: CHROMOSOME NUMBERS IN CROSSO- 
soMA, Peter H. Raven and Marion S. Cave 68 


A WEST AMERICAN JOURNAL OF BOTANY 


BLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


HERBERT L. Mason, University of California, Berkeley, Chairman 
EpcAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 

HERBERT F. COPELAND, Sacramento College, Sacramento, California 

Joun F. Davinson, University of Nebraska, Lincoln 
MitpreD E. MArtuias, University of California, Los Angeles 24 
MARION OWNBEY, State College of Washington, Pullman 
REED C. Rotiins, Gray Herbarium, Harvard University 
Ira L. Wiccrns, Stanford University, Stanford, California 


Secretary, Editorial Board—ANNETTA CARTER 
Department of Botany, University of California, Berkeley 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Herbert L. Mason, Department of Botany, University of California, 
Berkeley. First Vice-President: Paul C. Silva, Department of Botany, University of 
California, Berkeley. Second Vice-President: Robert F. Hoover, California State 
Polytechnic College, San Luis Obispo. Recording Secretary: Mary L. Bowerman, 
Department of Botany, University of California, Berkeley. Corresponding Secretary, 
Margaret Bergseng, Department of Botany, University of California, Berkeley. 
Treasurer: Douglas M. Post, Biology Department, San Francisco State College, San 


Francisco, California. 


Sn 


1963 | LANGENHEIM: LITTLE SUR FLORA 33 


QUATERNARY CLOSED-CONE PINE FLORA FROM 
TRAVERTINE NEAR LITTLE SUR, CALIFORNIA 


JEAN H. LANGENHEIM AND J. Wyatt DURHAM 


Interest in the relation of plants to deposition of calcareous charged 
water as travertine, tufa or sinter has existed at least since the time of 
Vergil and Pliny. Most of the studies have been concerned with currently 
active springs, and discussion has tended to center around the role of 
bacteria and algae, primarily, in producing deposition or at least in in- 
creasing precipitation of calcium carbonate from calcium bicarbonate 
(Agardh, 1827; Cohn, 1862; Weed, 1888; Meunier, 1899; Jones, 1914; 
Kellerman and Smith, 1914; Emig, 1917; et al.). In some cases other 
plant structures such as leaves, twigs, wood, etc., have been noted. The 
Little Sur travertine terraces in Monterey County, California, are inter- 
esting not only in terms of their formation and related geologic history, 
but also because the plant fossils give additional evidence regarding the 
distribution of the closed-cone pine forest along the central coast of Cali- 
fornia during the Pleistocene. 

The travertine outcrop was discovered by Durham during a class field 
trip in October, 1958, and collections were made at that time as well as 
during a similar trip the following year. Additional collections and recon- 
naissance studies were made by both authors later in the second year. 
Durham is responsible for the geologic observations and discussion herein. 
Thanks are due to Robert Simmonds for running the amino acid tests on 
the gastropod shells and to Allyn B. Smith for identifying the shells. 
Likewise appreciation is expressed to H. L. Mason for criticizing the 
manuscript and to Jack Wolfe for checking the determination of fossils 
as well as for criticism of the manuscript. Most of the plant fossil speci- 
mens are located in the Paleobotanical Collections of the Department of 
Botany, University of Illinois. Duplicate specimens are deposited in The 
Museum of Paleontology, University of California at Berkeley. Voucher 
specimens for description of the modern vegetation are in the Herbarium 
of the Department of Botany, University of Illinois. 


FORMATION OF TRAVERTINE 


The term “travertine” has been used for the Little Sur deposits because 
it seems to be the most inclusive term for deposits of this type. Emig 
(1917) has pointed out that the use of terms associated with this type of 
deposit has been varied and confused. He uses the term “travertine” for 
deposits of white, gray or brown concretionary calcium carbonate with 
cavernous and irregularly banded structure, soft and chalk-like to hard 
and crystalline, often containing leaves, twigs and mosses. Emig likewise 
indicates that ‘“tufa” is a “more ancient term” for travertine which was 
used by Vergil and Pliny in the same sense that travertine is now used in 


MaproNo, Vol. 17, No. 2, pp. 33-68. April 29, 1963. 


34 MADRONO [Vol. 17 


A ag 

VEL perp | 

\ ENS AOE! src : me : ; bass <a 2 i/ 

NWSW, @ Aas oe Ree ON: / 
\ \Wi7 lV Bod : eae 5 7m y \ ~ \ 1) 
2 a rz ey ‘ avy : side 


Fic. 1. Sketch of travertine terraces as seen from south. California State High- 
way 1 in foreground. 


Italy. He considers tufa as a cellular variety of calcite that has been de- 
posited from calcareous springs around nuclei of algae, mosses, leaves, 
twigs and other plant structures. Another term which has indiscriminate- 
ly been interchanged with travertine by some is “calcareous sinter” 
(Weed, 1888). In its original German sense sinter means dross from iron 
and implies an initial process of heating before deposition takes place. 
Thus Emig thinks that deposits formed in the immediate vicinity of hot 
springs may be correctly called “‘sinter.”” Twenhofel (1950) indicates that 
calcareous deposits around springs are known as travertine, but distin- 
guishes tufa and sinter as porous and travertine as compact and banded. 

The material from the Little Sur occurrence varies from a very soft 
chalky, cavernous rock to hard, finely crystalline concretions and bands. 
It varies in color from white to creamy to ochreous, and at times is 
stained with ferrous iron. Because of the proximity of the Serra Hill fault 
(Trask, 1926), it seems probable that the terraces were produced as a 
result of hot springs activity along this fault. Hot springs are also known 
to occur at the present time about twenty-one miles south of the area 


1963] LANGENHEIM: LITTLE SUR FLORA 315) 


along the Sur Thrust. Preservation of the leaves is often relatively poor, 
probably because of the effect of their having fallen into a hot spring. as 
well as the manner in which the calcium carbonate was precipitated. Fur- 
thermore, it is difficult to uncover the fossils in their entirety because 
they do not occur along a bedding plane and because frequently the 
margins of the leaves are curled under. 


LITTLE SUR TRAVERTINE TERRACES 


The terraces are located in the Santa Lucia Mountains, Monterey 
County, California. They are exposed along State Highway 1, nearly one 
mile north of the mouth of Little Sur River (lat. 36° 20.7’ N., long. 121° 
53.45’ W.). 

There are four prominent terraces on the east side of the road and on 
the south side of a prominent ravine (fig. 1), and there is seemingly at 
least one below the road on the cliffs extending to the ocean. There is con- 
siderable marble (Sur Series) cropping out of the hill slopes to the east 
of the presumed location of the Serra Hill Thrust fault (Trask, 1926) 
and also on the north side of the ravine. According to Trask’s map, the 
bedrock underlying the terrace deposits should be sediments of the Creta- 
ceous ‘‘Chico Group.” The south side of the present ravine on the north 
edge of the terraces has been cut into travertine which is 40—50 feet thick 
in places. It seems probable that the hot springs forming these deposits 
were in an old swale associated with the Serra Hill fault, and were located 
on an old surface prior to the development of the details of the present 
topography; the cascading pools were responsible for the sequence of 
terraces now observed. For convenience, the terraces have been numbered 
in sequence from the highway up the slope to the highest recognized, with 
the forefront of Terrace 1 being exposed in part in the road cuts. The 
apparent terrace below the road was observed from a distance only and 
is not included in the following descriptions. 

Terrace 1 extends approximately 120 feet from north to south; Ter- 
race 3, which is smaller, extends 200 feet from north to south; Terrace 2 
is a little over 150 feet in the same direction as is Terrace I. The frontal 
deposits of Terraces 1 and 2 are each at least 40—50 feet thick. 

The thickest accumulation of travertine is near the south margin of 
the adjacent ravine, seemingly indicating that the center of accumulation 
was either at that point or near the axis of the present ravine. The de- 
posits thin to the south, but Terraces 3 and 4 extend from the ravine on 
the north to the gully to the south where their southern extent is trun- 
cated by the present topography. Terraces 1 and 2 do not extend that far 
south. 

The lower part of the deposit consists of occasional pure layers and 
tongues of travertine intercalated with alluvial gravels. In the roadcut 
exposures and outcrops at the north end of Terrace 1, the upper 15—20 
feet of sediments are much purer travertine than the lower part. The 
lower part of this interval has beds with concentrated masses of Cupressus 


36 MADRONO [Vol. 17 


branchlets as well as a few pine cones. Just below the Cupressus band 
there is a zone with abundant leaves of plants such as Ceanothus, Garrya, 
Rubus, Ribes, etc. Occasional masses of abundant Equisetum or leaves 
of the above plants are scattered in the exposures along the roadcut. 

Much of the surface of the undisturbed exposures above the highway 
is covered with a dense caliche that probably conceals the fossil content. 
The best fossil collecting was in the roadcut and in the exposures at the 
north end of Terrace 1 on the south side of the ravine. On the upper ter- 
races the most abundant fossils observed were sedge-like leaves with 
scattered Garrya leaves. On the forefront of Terrace 2 clumps of sedges 
were noted in growth position, apparently indicating water flowing down- 
hill through a series of small pools. 

The time of formation of the travertine terraces can not be determined 
directly at the present time. Trask (1926) considered the faulting in the 
area to have occurred at the end of the Pliocene, but this might indicate 
mid-Pleistocene in present-day terms. Topographically, the terraces ap- 
pear to be situated on the upper part of the “steeply inclined slopes 
below” of Trask (1926, p. 176). However, the present sea cliffs in this 
area are obviously being eroded into this older topography and both the 
north and south edges of the terraces have been truncated by the erosion 
producing the present topography. Thus the terraces must be younger 
than Trask’s “steeply inclined slopes below,” and older than the pres- 
ent cycle of erosion. The ravine on the north is more than 50 feet deep 
where it cuts through the terraces and probably required considerable 
time for its formation. 

Two gastropods which occur commonly in the travertine were identi- 
fied as Plespericola pinicola Berry and Helminthoglypta dupetithouarst 
(Deshayes) by Allyn G. Smith of the California Academy of Sciences who 
stated that he can not distinguish the fossil specimens from those living 
in the area today; consequently he would think that the terraces were 
Sub-Recent in age. However, Mr. Robert T. Simmonds (unpublished 
Ph.D. thesis, Univ. of Illinois) has been studying gastropods of the gen- 
eral Polygyra group from the Recent and the Pleistocene to see if it is 
possible to establish ages by means of amino acid deterioration in the 
shells. Mr. Simmonds ran chromatogram tests for amino acids on the 
shells of fossil specimens of Helminthoglypta dupetithouarsi from the 
travertine and on present-day specimens from the terraces. On the basis 
of his amino acid dating technique, Simmonds states that the fossil 
shells in the travertine definitely predate historic times and in all prob- 
ability are more than 10,000 years old. A possible uncertainty in accept- 
ing the amino acid date might be the effect of heat (from the warm water 
of the hot springs) on deterioration of the amino acids. It is thought, 
however, that by the time the water had reached the surface and the 
travertine had been precipitated, the temperature must have been low- 
ered to a point where it would not seriously affect the amino acids. The 
age of 10,000 or more years would appear to be substantiated by the in- 


1963] LANGENHEIM: LITTLE SUR FLORA OW 


Ss 


Trinidad 
Head 


DISTRIBUTION MAP 
OF QUATERNARY AND RECENT 


CLOSED-CONE PINE FORESTS 


Drakes Bay & 


Mussel Rock--- 


“San Bruno 
Point Ano Nuevo 


panncug NN oe ES 


“San Simeon 
47 Cambria 


Morro Bay CY pacho Hills 


// Purisima Ridge 
Carpinteria 


Fic. 2. Map showing distribution of Pleistocene and recent closed-cone pine forests 
in California. Slanted lines indicate present-day distribution and black circles indi- 
cate fossil records of closed-cone pine forests. 


ferences from the topographic relationships of the terrace deposits. 

The plant fossils do not furnish conclusive evidence as to the age of 
the deposits as during the Pleistocene the coastal region of California did 
not have changes in flora comparable to those elsewhere apparent during 
the change from glacial to interglacial epochs. Mason (1934) indicates 
that northern plants range farther south but the general aspect of the 
flora was the same. A lag in plant response to climatic fluctuations has 
made the presence of northern species useless as a means of detailed cor- 
relation at the present time. It is only at the periphery of the distribution 
of the flora that differences can be noted which may be used for this 
purpose. 


38 MADRONO [Vol. 17 


Fossit PLANTS FROM THE TRAVERTINE 


Most of the fossils preserved in the travertine are impressions of leaves, 
with a few casts of stems and cones. In most of the other Pleistocene 
deposits recording the history of the closed-cone pines in California, ie. 
Carpinteria, Santa Cruz Island and Tomales Bay, there have been many 
more fruits or seeds preserved than leaves. Several samples of the traver- 
tine were tested for pollen and spores, but no recognizable remains were 
observed. 

Of the species represented by the travertine fossils, the following are 
considered to be forest plants: Pinus radiata, P. muricata (?), Pseudot- 
suga menziesu (?), Cupressus cf. goveniana, Quercus agrifolia, Myrica 
californica, Ribes cf. sanguineum var. glutinosum, Rubus cf. parvti- 
florus (?), R. cf. vitifolius (2), Ceanothus cf. griseus, Garrya elliptica; 
and the following hydric or streamside plants: Equisetum cf. hiemale 
var. californicum, Carex spp., Juncus spp., algae. 

Descriptions of the fossils and discussions of the modern occurrence of 
the species represented follow. 


PINUS RADIATA Don and P. MuricaATA Don (fig. 3, g and /). Casts of 
three partially preserved cones were found in the lower beds of Terrace 1. 
Preservation is poor, or at least the cones appear to have been weathered 
previous to being incorporated into the travertine. One specimen has only 
the lowermost scales preserved; however, the umbos are rounded to a 
knob which is a distinctive character of P. radiata. Another specimen has 
what appears to be one side of the cone preserved. It is 8 cm. long and 
ca. 5 cm. wide. The umbos on the lower scales are rounded, but they are 
also of a form which might be interpreted as that of P. muricata. The 
third specimen is so poorly preserved that it can only be stated that there 
is indication of a pine cone. However, below the piece of cone there ap- 
pears to be a fascicle of three needles. Pinus muricata has two needles in 
a fascicle whereas P. radiata may have either three or two needles. The 
needles appear to be narrower than the average for extant P. radiata. 
As a result of some operation that caused the margins of the needles to 
become inrolled toward the midvein, the needles are circular in cross sec- 
tion and their width has been reduced. This inrolling may have occurred 
if they dropped into hot spring water. There also appear to be other scat- 
tered fragments of needles which may be turned on edge. 

Fossil cones and wood of P. radiata have been found in the Pliocene 
Merced Sandstone (Glen, 1959), the Pleistocene deposits at Mussel Rock, 
at Carpinteria, and at Tomales Bay. Pinus radiata is the most abundant 
species in the Tomales deposit where it is represented by wood, cones, 
and leaves. The only previously known fossil occurrence for P. muricata 
is the Tomales Formation where it was far less abundant than P. radiata. 

At present P. radiata is a coastal endemic of central western California, 
occurring from Pescadero, San Mateo County south to San Simeon, Mon- 
terey County. A variety also occurs on the coastal islands off of southern 


1963 | LANGENHEIM: LITTLE SUR FLORA 39 


California. Pinus muricata ranges discontinuously through about ten de- 
grees of latitude, from Trinidad Head in Humboldt County to La Purisi- 
ma Ridge in Santa Barbara County, thence southward through insular 
California to Guadalupe Island. On Huckleberry Hill near Monterey 
both P. radiata and P. muricata occur together but P. muricata tends to 
occur on shallow soils. 


PSEUDOTSUGA MENZIESII (Mirb.) Franco(?). Another cast which is 
apparently a cone has a maximum length of 2.5 cm. and diameter of ca. 
2 cm. It suggests either Picea or Pseudotsuga. There is no indication of 
the bract subtending the cone scale which is the usual means of identi- 
fying living Pseudotsuga menziesii. This has been true of all of the other 
Pleistocene records of the species (Mason, 1927, 1940; Potbury, 1932; 
Chaney and Mason, 1934); however, in Carpinteria and Tomales de- 
posits, the portion of the bract lying underneath the cone scale and pro- 
tected thereby, is preserved. This serves as a means of separating fossil 
cones of Pseudotsuga from those of Picea. In the Tomales flora certain 
fossils were identified as Picea because the cones bore a greater number 
of scales in proportion to their size than do those of Pseudotsuga and also 
because needles and twigs were found. Preservation of the Little Sur 
material is too poor to make positive determination possible. Mason 
(1934) states, however, that Tomales is the southern limit of the known 
range of Picea in the Pleistocene. 

Pseudotsuga perhaps shared dominance with Pinus radiata and P. muri- 
cata of the forests about Tomales Bay in the Pleistocene as indicated by 
the hundreds of parts of cones. Pseudotsuga has also been reported from 
the Willow Creek flora on Santa Cruz Island. It has been recorded in the 
Pliocene of California (Dorf, 1930) where it is most often associated with 
Sequoia. 

The modern range of Pseudotsuga menziesii along the coast is from 
Salmon Creek in the Santa Lucia Mountains northward to the northern 
end of Vancouver Island, British Columbia. 


CUPRESSUS cf. GOVENIANA Gord. (fig. 3, b). Concentrated masses of 
branchlets of Cupressus occur in the lower part of Terrace 1, in the same 
interval where the pine cones were found. The branchlets are ca. 1—1.5 
mm. wide; the leaves likew.se are approximately 1—-1.5 mm. long, oppo- 
site, appressed, acute but tending to be more or less blunt. Two cupres- 
soid fructifications were found. 

Species of Cupressus are common in the California Pleistocene, being 
reported from La Brea, Carpinteria, Santa Cruz Island and Tomales de- 
posits. Mason (1927, 1934), however, points out the difficulty in specific 
identification of fossil material because of the close relationships among 
the various modern species. Determinations must be based largely upon 
ecological considerations suggested by the association of species and in 
many cases such a determination can not be accepted as final. 


40 MADRONO [Vol. 17 


Cupressus macrocarpa is restricted to granodiorites on the headlands 
of Monterey Bay where it is an associate of Pinus radiata. Cupressus go- 
veniana occurs around Monterey on the Monterey Shales, and is asso- 
ciated in the highlands with both Pinus radiata and P. muricata. In Men- 
docino County Cupressus pygmaea occurs in the vicinity of Fort Bragg, 
where it is an associate with Pinus muricata, Pseudotsuga menziesi, Picea 
sitchensis, Tsuga heterophylla and Sequoia sempervirens. The only other 
possibility for these fossils is Cupressus sargentit which has the widest 
range of the species of Cupressus in California. It is not strictly coastal 
although it is reported from some localities near the sea. It ranges from 
the southern Santa Lucia Mountains north to Mendocino County. De- 
spite its wider geographic distribution, it tends to be limited to serpentine 
outcrops. Thus from the location, general association and the absence of 
granodiorites in the vicinity of the terraces, it appears that the branch- 
lets in the travertine are more likely to be referable to C. goveniana than 
the other species. 


QUERCUS AGRIFOLIA Nee (fig. 3, e; fig. 4, e). Upon initial study, the 
abundant specimens of Garrya elliptica were thought to be variable forms 
of Quercus agrifolia. However, close investigation of the venation pat- 
terns showed that a few specimens only were probably Quercus. These 
specimens occurred in the same beds with those assigned to Garrya. No 
fructifications were found. 

Quercus agrifolia has been reported in the Pleistocene both from the 
asphalt beds of Rancho La Brea (Frost, 1927) and Carpinteria as well 
as the Tomales Formation (Mason, 1934). The Carpinteria material is 
more nearly like the shrub form of the species, i.e. those occurring on 
sterile shoots or crown sprouts that come up after fire. Neither acorns nor 
cups were found in the Carpinteria deposits, although they are common 
at Tomales Bay and elsewhere in the Pleistocene. 

The modern Q. agrifolia typically is a coastal oak ranging through the 
coastal mountains of central and southern California. In the isolated rem- 
nants of the original closed cone-pine forests scattered from La Purisima 
Ridge northward to Sonoma County coast, Q. agrifolia is the dominant 


EXPLANATION OF FIGURE 3. 


Fic. 3. Fossils from Little Sur travertine deposits. a, Rubus cf. parviflorus Nutt. 
Portion of folded leaf. & 1. Univ. I'linois Pa‘eobot. Coll., Hypotype L-1. 6, Cupressus 
cf. goveniana Cord. Branchlet and possible frutification. * 1. Univ. Illinois Paleobot. 
Coll., Hypotype L-3.c. Rubus cf. parviflorus Nutt. Portion of leaf. x 1. Univ. Illinois 
Paleobot. Coll., Hypotype L-2. d, Ribes cf. sanguineum var. glutinosum (Benth.) 
Loud. Portion of leaf. * 1. Univ. Illinois Paleobot. Coll., Hypotype L-4. e, Quercus 
agrifolia Nee. Portion of leaf. * 1. Univ. Illinois Paleobot. Coll., Hypotype L-6. 
f, Ribes cf. sanguineum var. glutinosum (Benth.) Loud. Portion of leaf. & 1. Univ. 
Illinois Paleobot. Coll., Hypotype L-5. g. Pinus muricata Don? Portion of cone. X 1. 
Univ. Illinois Paleobot. Coll., Hypotype L-7. h, Pinus radiata Don. Portion of cone. 
x 1. Univ. Illinois Paleobot. Coll., Hypotype L-8. 


1963] LANGENHEIM: LITTLE SUR FLORA 41 


g 


Fic. 3. Fossils from Little Sur travertine deposits. 


42 MADRONO [Vol. 17 


understory tree in the forest. From the basis of general association, it 
seems probable that these fossil leaves are referable to QO. agrifolia. 


MyrIcA CALIFORNICA C. & S. A single leaf impression is suggestive of 
Myrica californica. It is ca. 4 cm. long and 1 cm. wide, and the dentition 
is not preserved because of strong inrolling of the margin of the leaf. 
Although preservation is not sufficient for unquestioned identification, 
the venation seems to indicate this species. 

M yrica californica is represented in the Pleistocene deposits of Tomales 
Bay by an abundance of fruits and a few leaf impressions from indurated 
nodules. It is also known from the Santa Cruz Island and Carpinteria 
floras. No records are available from the more arid Rancho La Brea 
deposits. 

The extant Myrica californica is strictly a coastal plant, attaining its 
best development only on the coastal side of the mountains in valleys im- 
mediately exposed to the sea. It ranges from the Puget Sound region 
southward through the coastal area of Oregon and northern California, 
thence becoming rarer southward where it extends to the Santa Monica 
Mountains of southern California. Mason (1934) suggests that the south- 
ernmost records of present distribution are probably relicts of Pleisto- 
cene distribution. The distribution of Myrica californica ends abruptly 
in the region of what probably was the southernmost extension of the 
cool humid Pleistocene climate. 


RIBES cf. SANGINEUM var. GLUTINOSUM (Benth.) Loud. (fig. 3, d and 
f). Ribes leaves occur frequently with Rubus in beds of Terrace 1. As in 
the case of the other leaves, none are preserved in entirety, with masses of 
fragments occurring on top of each other. Almost all of the fragments are 
bent and frequently the edges turned under. The leaves are rounded in 
general outline with palmate divergence of three major veins indicating 
three lobes. They average 2—5 cm. wide. These characters tend to indicate 
the Ribes sangineum group, although the venation pattern fits the variety 
glutinosum better than the typical form. The variety now occurs in open 
places or brushy sites or in the closed-cone pine forest from Santa Bar- 
bara to Humboldt County. There is no previous Pleistocene record. 


Rusus cf. PARVIFLORUS Nutt. and Rubus cf. viTIFoLIus C. & S. (fig. 3, 
a and c). In the lower beds of Terrace 1 Rubus leaves occur commonly. 
They tend to be found with masses of Ribes leaves and usually apart from 
the large concentrations of Ceanothus and Garrya which occur in the 
same sequence of beds. As usual only fragments of the leaflets are pre- 
served, with the specimens being bent and folded so that it is difficult to 
determine the nature of the whole leaflet in most specimens. Some of the 
specimens are generally ovate in shape with others being palmately lobed. 
The size likewise varies greatly with the amount of the fragment pre- 
served. Venation characters help indicate the size, however, especially by 
indicating the possibility of lobing. In many fragments the venation is 


1963 | LANGENHEIM: LITTLE SUR FLORA 43 


pinnate toward the terminal portion of the leaflet, but where the lower 
portion of the leaflet is preserved there is palmate divergence even though 
the lobe may not be intact. 

The characters of some of the fragments suggest Rubus vitifolius as 
leaflets of this form may be either ovate or palmately lobed. However, 
other specimens, in terms of larger size and nature of the lobing appear 
to fit Rubus parviflorus better. Thus there appears to be little question 
of the presence of Rubus, but the amount of a leaflet preserved raises 
a question as to specific determination. 

The only known Pleistocene record of Rubus vitifolius is from Tomales 
Bay (Mason, 1934) although Mason assumed that it was present in most 
of the coastal region of California. At present Rubus vitifolius is one of 
the most abundant shrubs in the valley and low hill country of California, 
occurring chiefly along streams and springy flats. 

Rubus parviflorus fruitlets occur in the Pleistocene beds of Tomales 
Bay and from the San Bruno deposits. At the present time it is common 
along canyon streams and in open woods from southern California to 
Alaska. A clearly marked variety, called velutinus, occurs at the present 
time in the closed-cone pine and redwood forests near the coast from 
Santa Barbara to Mendocino County. It frequents streambanks and moist 
swales not too exposed to the sun. 


CEANOTHUS cf. GRISEUS (Trel.) McMinn (fig. 4, b and c). Ceanothus 
leaves are abundantly represented in Terrace 1 in the beds which likewise 
frequently contain Garrya and sedge-like leaves. The leaves appear to 
have fallen in great quantities into the pool at one period, as the speci- 
mens are frequently laid one on top of the other. The specimens are frag- 
mentary with the edges of the leaves universally curled under. This is due 
not only to the possible curling effect of hot water, but also to the natural 
tendency of leaves of some species of Ceanothus to be revolute. Because 
the edges are curled, it is impossible to determine the margin. The shape 
generally is oblong-ovate to broadly elliptical. The portion preserved 
varies in length up to 4 cm., indicating that some were even longer. The 
greatest width is 2.5 cm. with 1.5 cm. being common. Some show three 
prominent veins, although a few have only one. The lateral “‘reticulate” 
venation pattern is also prominent. These characters fit within the range 
described for C. thyrsiflorus or C. griseus. At one time C. griseus was con- 
sidered a local variant of C. thyrsiflorus (Jepson, 1925), although recent- 
ly it has been raised to specific status (McMinn, 1942; Munz and Keck, 
1958). 

Ceanothus thyrsiflorus is native to the cool coastal region from Santa 
Barbara County, California, north to Douglas County, Oregon, being 
especially abundant in the redwood belt of central and northern Califor- 
nia. On the Monterey Peninsula it is replaced by C. griseus which occurs 
with the closed-cone pines and on the open slopes in the chaparral. Both 
of these species hybridize with other species of Ceanothus, providing ad- 


44 MADRONO [Vol. 17 


ditional sources of variation. The general characers preserved in these 
fossils place them within either the group of C. thyrsiflorus or that of 
C. griseus. The habitat and the associated community however suggest 
C. griseus. 

Ceanothus thyrsiflorus has been reported from the California Pleisto- 
cene beds of Santa Cruz Island, Carpinteria, San Bruno and Tomales 
Bay. Mason (1934) states that the one leaf impression from the Tomales 
flora looks much like the gviseus variant, but restricts his identification 
to C. thyrsiflorus. 


GARRYA ELLIPTICA Dougl. (fig. 4, d and f). Fragments of leaves that 
appear to be Garrya occur most abundantly in beds of Terrace 1, just 
below the zone containing the masses of Cupressus branchlets. However, 
they occur occasionally in deposits of the upper terraces where sedge-like 
leaves are the predominant observed fossils. The fact that only Garrya 
was found with the sedges, however, may be more a function of the expo- 
sures being covered with caliche and thus either masking the other fossils 
or rendering access to fossils very difficult in the upper terraces. On the 
other hand, the leaves of Garrya are more coriaceous than most of the 
others found and thus may have been preserved better under the possibly 
more unfavorable conditions in the upper hot pools. 

The portions of the leaves preserved vary from 3 to 6 cm. in length 
and they indicate that the entire leaves must have been oblong or elliptic. 
Few leaf margins are intact, the edges usually being curved under. The 
general character of the leaf is within the range of variability occurring 
in Garrya elliptica, although one could easily confuse leaves of this group 
with those of the live oaks (particularly Quercus agrifolia). 

Garrvya elliptica is a coastal species ranging through west-central Cali- 
fornia from Monterey north to Humboldt County. It is frequent in the 
understory of the closed-cone pine forests at Monterey and may serve as 
an indication of considerable fog and a rainfall of at least twenty to forty 
inches. 

The genus Gavrya has been described from Pliocene beds of Bennett 
Valley and Coalinga, California (Dorf, 1930). In the Pleistocene the only 
known records are based on specimens closely referable to the modern 
G. elliptica. The Pleistocene specimens from Carpinteria, and Santa Cruz 
Island are both south of the present range. 


EQUISETUM Cf. HIEMALE var. CALIFORNICUM Milde (fig. 4,a). Portions 
of stem occur thoughout the beds from Terrace 1 and are particularly 
abundant in the exposure along the roadcut. They are preserved as in- 
ternal casts, showing only the nodes and no sheaths; the stems average 
8-10 mm. in diameter. The internodal ridges are well preserved; the aver- 
age number is approximately 30. 

The specimens could be referable either to Equisetum laevigatum or 
E. hiemale var. californicum as far as diameter (2—8 mm. or 5-15 mm. 
respectively) is concerned. However, the fossils are larger and more 


1963 | LANGENHEIM: LITTLE SUR FLORA 45 


Fic. 4. Fossils from Little Sur travertine deposits. a, Equisetum cf. hiemale var. 
californicum Milde. Portion of stem. X 1. Univ. Illinois Paleobot. Coll., Hypotype 
L-9. b. Ceanothus cf. griseus (Trel.) McMinn. Portion of leaf. * 1. Univ. Illinois 
Paleobot. Coll., Hypotype L-10. c, Ceanothus cf. griseus (Trel.) McMinn. Portion 
of leaf. x 1. Univ. Illinois Paleobot. Coll., Hypotype L-11. d, Garrya elliptica Doug]. 
Portion of leaf. & 1. Univ. Illinois Paleobot. Coll., Hypotype L-12. e, Quercus agri- 
folia Nee. Portion of leaf. & 1. Univ. Illinois Paleobot. Coll., Hypotype L-14. f, Gar- 
rya elliptica Dougl. Portion of leaf. 1. Univ. Illinois Paleobot. Coll., Hypotype L-13. 


robust in general than E. laevigatum. The two distinct rows of silica 
tubercles also tend to indicate E. hiemale var. californicum. 

Potbury (1932) reported small portions of Equisetum stems with nodes 
and portions of internodes from the San Bruno deposits. There was no 
previous report of Hquisetum in the California Pleistocene but Potbury 
predicted that it would be recorded. 

Equisetum hiemale var. californicum is found living alongside streams 
and in marshy places from southern California to Humboldt County and 
thence north to Alaska. 


CAREX spp. and JuNcus spp. Abundant sedge-like leaves and stems 
occur throughout the beds of the various terraces. There are the usual 
signs of curling or inrolling of the leaves. They vary in width from 2 mm. 
to 18 mm., the most common range being 2.5—5 mm. They give the gen- 
eral appearance of leaves of sedges with very prominent vertical stria- 


46 MADRONO [Vol. 17 


tions. A few flat leaves approximately 15-18 mm. wide might indicate 
Typha latifolia; however, lack of horizontal striations makes this iden- 
tification seem unlikely. The stems are circular and hence would not 
be sedges which have a triangular cross section. Along the streams and 
springs in the area at present both sedges and rushes are common. 
Grasses, however, can not be excluded as a possibility, but sedges seem 
more probable in a hot springs environment. Along the edge of Ter- 
race 2 masses of sedge appeared to be in growth position, bent in such 
a manner as to indicate water flowing downhill. On this terrace almost 
any portion of travertine sampled for fossils produced masses of these 
Carex and/or Juncus leaves. 

Carex is known from both the San Bruno and Tomales Pleistocene 
deposits in California. In both cases, perigynia, which are a much better 
means of identification than leaves, were preserved. There is no record of 
Juncus from the California Pleistocene, but there is no reason why it 
should not be expected. 


AtGaAE. In several beds tangled masses of fibrous filaments occur 
which give the appearance of algae. A sample was tested for the presence 
of spores but no recognizable forms were recovered. Thus determination 
of these obvious algal remains can not be made. 

Certainly the presence of algae is to be expected in travertine deposits. 
Davis (1897) states that algae are responsible for the peculiarities in 
structure of travertine, i.e. the formation of filaments, ropes, beads, tufts, 
etc. Otherwise he believes that the calcium carbonate would be laid down 
“like coats of whitewash.” The algal inhabitants of hot springs are all 
members of one or two families, closely related to one another, as condi- 
tions are generally not suitable for most algae. Emig (1917) states that 
in the Arbuckle Mountains of Oklahoma there is a natural cycle of 
plants that accompanies the appearance and development of travertine. 
During the earliest states of development the unicellular green algae 
(i.e. Protococcus) and blue-green algae (i.e. Oscillatoria and Lyngbya) 
are present, followed by filamentous green algae (i.e. Vaucheria, Oedo- 
gonium, Rivularia, Cladophora, et al.) that grow in felt-like masses. Also 
mosses that aggregate in dense tufts are present. 


VEGETATIONAL RELATIONSHIPS 


An assemblage of the recognized plants in the fossil assemblage indi- 
cates a closed-cone pine forest with streams or springs along or through 
the forest. The forest apparently included Pinus radiata, possibly P. muri- 
cata, and Cupressus goveniana. There is likewise a possibility of Pseudo- 
tsuga menziesi. The understory trees include an abundance of Garrya 
elliptica, Ceanothus griseus, lesser amounts of Quercus agrifolia and pos- 
sibly Myrica californica. Understory shrubs include Rubus parviflorus, 
Rubus vitifolius and Ribes sanguineum. Springs are necessary to the for- 
mation of travertine, and thus the presence of Equisetum as well as 
sedges, rushes, and algae was to be anticipated. 


1963 | LANGENHEIM: LITTLE SUR FLORA 47 


It appears probable that in this area there was a closed-cone pine forest 
along a canyon or swale with travertine being deposited along a stream— 
hot spring area. Perhaps as the hot springs became more active, imme- 
diately adjacent trees and shrubs were killed, possibly accounting for the 
abundance of needles of Cupressus and leaves of Ceanothus and Garrya. 
Likewise as activity became even more extensive, only sedges and rushes 
could persist, hence their prevalence in certain portions of the terraces. 

The present vegetation surrounding the area of the terraces gives no 
evidence of closed-cone pine forest. It is predominantly a coastal sage 
pattern, which generally is characteristic of the drier and rockier slopes 
of the outer California Coast Ranges. The major dominant in the Little 
Sur area is Artemisia californica with Baccharis pilularis very abundant. 
Salvia mellifera, Rhus diversiloba, and Pteridium aquilinum also are 
common. Eriogonum spp., Eriodictyon crassifolium, Diplacus aurantia- 
cus, Dendromecon rigida, Agastache spp. are scattered although locally 
common. 

Temporary streams occupy most of the nearby ravines and gullies, al- 
though a few apparently have a year-round source of water. An occasional 
wind-trained redwood and Photinia arbutifolia occur in such sites. About 
a quarter of a mile south of the terraces there is a semi-permanent 
stream with Salix scouleriana, Amelanchier alnifolia, Cornus californica 
and Rkamnus spp. forming a dense shrubby overstory. The understory 
consists of Equisetum telmateia and Stachys chamissonis, with Rubus 
vitifolius climbing over the other plants. Species of Carex and Juncus 
are also present. 

Coastal sage is the predominant vegetation pattern northward from 
the terraces to approximately Bixby Creek (about 3 miles from Little Sur 
River) where species of Ceanothus and Arctostaphylos form a dense 
chaparral cover. From thence north to Carmel, coastal sage and chaparral 
alternate, with the latter occupying the more mesic sites such as north- 
facing slopes. Pinus radiata and Cupressus goveniana do not appear until 
near Carmel. The forest occupying the ridge between Carmel Bay and 
Monterey Bay is probably the best developed closed-cone pine forest in 
central and southern California. This forest occupies the entire Monterey 
Peninsula to the exclusion of other forest types, but extends only a few 
miles into the interior, and only a short distance to the north as well as 
to the south. Whether this forest was more extensive in the early history 
of white man is not known, but there is record of lumbering operations. 
Also pine roots have been found in the soil to the north and Mason (1934) 
states that it is probable on the basis of typically associated species that 
the forest once extended northeastward to the Salinas River. 

To the immediate south of the area, coastal sage is replaced by grass- 
land with scattered oaks (primarily Quercus agrifolia), thence into a local 
redwood forest in the Big Sur area. Southward the only known groves 
of Pinus radiata occur near San Simeon and Cambria. The forests are 
similar in aspect to those of the Monterey region but not as rich in species. 


48 MADRONO [Vol. 17 


Occasional individuals of P. muricata and the kinds of shrubs suggest 
more favorable soil conditions than at Monterey. In the Pecho Hills, 
pine is largely replaced by a dense live oak assemblage. Neither pines 
nor Ceanothus occur in an assemblage at Purisima Ridge that is other- 
wise like the Monterey forest. Pinus radiata var. binata likewise occurs 
on Guadalupe Island. 

Despite the fact that obviously only a small portion of the forest flora 
was preserved in the travertine, the pattern of the flora fits that of the 
modern Monterey closed-cone pine forest. The entire forest there is dom- 
inated by Pinus radiata; the most conspicuous understory tree is Quercus 
agrifolia with scattered trees of Myrica californica and Umbellularia cali- 
fornica. The most characteristic feature of the forest is the extensive 
chaparral-like ground cover made up of species of Ceanothus and Arcto- 
staphylos associated with a number of other shrubs such as Adenostoma, 
Lonicera, Ribes, Rosa, Rubus, Symphoricar pos, et al. At slightly higher 
elevations Cupressus goveniana occurs, whereas Pinus muricata is found 
in areas of greatest summer fog and heaviest winter rainfall. Mason 
(1934) points out that local variations in ecological aspect of the forest 
flora seem to be related to depth and character of the soil as well as atmos- 
pheric moisture. Thin, rock soil layers contain dense brush and in some 
cases take on an aspect of typical pine barrens. The largest trees and 
most open forest occur when soil is deep and ground water abundant. 
From the possible presence of P. muricata as well as Cupressus, it appears 
that the forest at the time of travertine deposition might have been at a 
site with considerable fog. Myrica likewise indicates this. The preponder- 
ance of Garrva elliptica and relatively few preserved specimens of Quer- 
cus agrifolia are noteworthy. It is likewise interesting that there is an 
abundance of Ceanothus but no record of Arctostaphylos, which one 
would expect to be preserved more eas:ly than the Ceanothus. The abun- 
dance of Ribes and Rubus are to be expected in a more mesic aspect of 
the forest such as one finds along streams or shaded canyons, and prob- 
ably on deeper soils. 


VEGETATIONAL HISTORY 


The present distribution of the closed-cone pine forest is highly dis- 
continuous (fig. 2). The northernmost outpost is Trinidad Head in north- 
ern California, where it is almost obscured by redwood forest. There then 
is a gap 100 miles south to Inglenook where it occurs as typical coastal 
forest for about 100 miles to Fort Ross. After a 50-mile break it recurs 
on Inverness Ridge across from the fossil localities at Tomales Bay. Some 
remnants also occur on San Geronimo Ridge in Marin County. There is 
another 75 mile gap to Point Ano Nuevo where another small grove 
occurs, then another 40 miles to the south end of Monterey and Carmel 
bays where there is an extensive forest. It is another 60 m.lies south to 
San Simeon and Cambria before the forest type is encountered again, and 
thence southward to the isolated patches occurring at Pecho Hil's and 


1963 ] LANGENHEIM: LITTLE SUR FLORA 49 


La Purisima Ridge. There is another break on the mainland of 500 miles 
to Point San Quintin, Baja California, Mexico. Remnants of closed-cone 
pine forests also occur on Cedros Island 150 miles south of Point San 
Quintin, and on Guadalupe Island 200 miles off the coast of Baja Cali- 
fornia. They also occur on Santa Rosa and Santa Cruz islands of the 
Channel group. 

These modern forests are not homogeneous floristically or ecological- 
ly. Evidence from fossil deposits from such widely separated areas as 
Tomales Bay, Carpinteria and Santa Cruz Island indicates that this for- 
est assemblage is a relict of a past flora and that it was more homogeneous 
throughout its range in the past than today (Mason, 1934). Mason also 
points out that the present discontinuity is due to significant geological 
events, and that for a relatively long interval of time the forest type 
occupied an extensive area, of which its modern occurrences are but 
fragments. In all localities in the Pliocene and Pleistocene where fossil 
records of closed-cone pines are known, the site of deposition has been 
along one of the major fault lines in the California Coast Ranges. Like- 
wise in all cases the areas occur along fault zones near a block that has 
been positive throughout the Pliocene at least. Most conspicuous are the 
blocks severed from the mainland by the San Andreas fault. In the Pleis- 
tocene deposits some of the plant materials have come from lands up- 
lifted since the Pliocene but presence of sediments from older landmasses 
has always been demonstrable. 

Mason likewise states that the closed-cone forests today are confined 
to lands that were islands throughout much of the Tertiary and particu- 
larly during the Pliocene. Homogeneity of species content and absence of 
other contemporary assemblages in the Pleistocene support this hypoth- 
esis. This insular relationship is further supported by the high degree of 
endemism among the forest species: 58 per cent of the woody plants are 
endemic to the California floristic province with 29 per cent of the above 
restricted to closed-cone pine forests. This is remarkable considering the 
very small area of the California province occupied by closed-cone pines. 
Because most of the endemic species have closely related species else- 
where in the Coast Ranges, it appears probable that there was insular 
isolation with water barriers. Also the relation of the modern flora to that 
of the islands off of southern California is important. Several species such 
as Vaccinium ovatum and Arbutus menziesiu have their southward exten- 
sion on islands or occur on the southern mainland only in localities that 
appear to have been Tertiary islands. Pinus radiata grows in the general 
region of Monterey Bay, Morro Bay and on Guadalupe Island (200 miles 
off the coast of Baja California), but does not grow on the mainland in 
between. The sporadic occurrence of Cupressus from Guadalupe Island 
to coastal Mendocino County also is another similar distribution. Cupres- 
sus is known to have occurred in every closed cone-pine forest of which 
there is fossil record. In many areas it now is extinct, e.g. Santa Barbara 
region and Santa Cruz Island. Likewise Pinus radiata has disappeared 


50 MADRONO [Vol. 17 


from both of these localities, and both P. radiata and Cupressus are now 
absent from Tomales Bay although they were common there during the 
Pleistocene. 

Mason further indicates that the major differences in aspect that pre- 
vail in the forest throughout its range today are due primarily to the 
mingling of the original species with those of migrating populations and 
consequent selection of floras due to changing climatic conditions in Pleis- 
tocene and Recent time. The California insular floras are considered by 
Mason to represent the original vegetation of coastal California prior to 
their invasion by continental floras. The discontinuous closed-cone pine 
forests along the California coast are the last remnants of these insular 
floras. The fossil occurrences described in this investigation give addi- 
tional documentation of the former more extensive distribution of this 
vegetation type. 


SUMMARY 


An assemblage of plant fossils from travertine terraces in Monterey 
County, California, provides additional documentation of a former more 
extensive distribution of closed-cone pine forests during the Quaternary. 
The travertine terraces are considered to be 10,000 or more years old as 
evidenced by amino-acid dating of gastropod shells in the travertine and 
substantiated by the topographic relationships of the deposits. The plant 
fossils are relatively poorly preserved impressions of leaves, with a few 
casts of stems and cones. Despite the fragmentary record, the forest ap- 
parently included Pinus radiata, Cupressus goveniana and possibly Pinus 
muricata and Pseudotsuga menziesu. An abundance of Garrya elliptica, 
Ceanothus griseus, lesser amounts of Quercus agrifolia and possibly My- 
rica californica occurred as understory trees. Rubus parviflorus, Rubus 
vitifolius and Ribes sanguineum var. glutinosum comprise the recorded 
understory trees. Equisetum hiemale var. californicum as well as sedges, 
rushes and algae also occurred commonly. Thus it appears probable that 
there was a closed-cone pine forest along a canyon or swale with traver- 
tine being deposited along a stream—hot spring area. The present vege- 
tation surrounding the area of the terraces, being predominantly a coastal 
sage pattern, gives little evidence of closed-cone pine forest. The pattern 
of the fossil flora, however, fits closely that of the modern closed-cone 
pine forest on the Monterey peninsula. The location of the travertine 
terraces also generally supports Mason’s (1934) conclusion that fossil 
records of closed-cone pines have occurred near fault blocks that were 
positive during the Pliocene. This insular relationship helps to account 
for the highly discontinuous and endemic nature of the floras. 


Biological Laboratories, Harvard University, 
Cambridge, Massachusetts 


Department of Paleontology, 
University of California, Berkeley 


1963] LANGENHEIM: LITTLE SUR FLORA on 


LITERATURE CITED 


AGARDH, C. 1827. Aufzahlung einiger in den 6streichischen Landern gefundenen neuen 
Gattungen und Arten von Algen, nebst ihrer Diagnostik und beigefugten Be- 
merkungen. Flora 10:625-640. 


CHANEY, R.W. and H. L. Mason. 1930. A Pleistocene flora from Santa Cruz Island, 
California. Carnegie Inst. Publ. 415:1-24. 


. 1933. A Pleistocene flora from the asphalt deposits of Carpinteria, Cali- 
fornia. Carnegie Inst. Publ. 415:45-79. 


Coun, F. 1862. Einen Vortrag uber die Algen des Karlsbader Sprudels und deren 
Antheil an der Bildung des Sprudelsinters. Flora 45:538—-540. 


Davis, B. M. 1897. The vegetation of the hot springs of Yellowstone Park. Science, 
NS 6:145-157. 


Dorr, E. 1930. Pliocene floras of California. Carnegie Inst. Publ. 412:1-108. 
Emic, W.H. 1917. Travertine deposits of Oklahoma. Okla. Geol. Surv. Bull. 29:1-76. 
Frost, F. H. 1927. The Pleistocene flora of Rancho La Brea. Univ. Calif. Publ. Bot. 
14:73-98. 
GLEN, WILLIAM. 1959. Pliocene and lower Pleistocene of the western part of the 
San Francisco peninsula. Univ. Calif. Publ. Geol. Sci. 36(2) :147-198. 
Jepson, W.L. 1925. A manual of flowering plants of California. Univ. Calif. Press. 
Berkeley. 
Jones, J. C. 1914. The tufa deposits of the Salton Sink. Carnegie Inst. Publ. 193: 
79-84. 
KELLERMAN, K. F. and N. R. SmitH. 1914. Bacterial precipitation of calcium car- 
bonate. Jour. Wash. Acad. 4:400-402. 
Mason, H. L. 1927. Fossil records of some west American conifers. Carnegie Inst. 
Publ. 346:139-158. 
. 1934. Pleistocene flora of the Tomales formation. Carnegie Inst. Publ. 
415:81-179. 
. 1940. A Pleistocene record of Pseudotsuga macrocarpa [Carpinteria, Cali- 
fornia]. Madrono 5:233-235. 
. 1942. Distributional history and fossil record of Ceanothus, in Ceanothus 
by M. van Rensselaer and H. E. McMinn. Santa Barbara Bot. Gard. 
. 1957. A Flora of the Marshes of California. Univ. Calif. Press. Berkeley 
and Los Angeles. 
McMinn, H. E. 1942. A systematic study of the genus Ceanothus, in Caenothus by 
M. van Rensselaer and H. E. McMinn. Santa Barbara Bot. Gard. 


Meunier, S. 1899. Observations relativ du depot de certain travertins calcares. 
Compt. Rend. Acad. Paris 129:659-666. 


Muwz, P.A. and D. D. Keck. 1959. A California flora. Univ. Calif. Press. Berkeley 
and Los Angeles. 


PotBury, SUSAN S. 1932. A Pleistocene flora from San Bruno, San Mateo County, 
California. Carnegie Inst. Publ. 415:25-44. 

SIMMONDS, R. T. 1962. Wisconsin Geochronological determinations based upon amine 
decay rates. Ph.D. dissertation, Dept. Geol., Univ. Illinois, Urbana. 

Trask, P. 1926. Geology of Point Sur Quadrangle, California. Univ. Calif. Publ., 
Bull. Dept. Geol. Sci. 16:119-186. Pl. 16. 

TWENHOFEL, W. H. 1950. Principles of sedimentation. McGraw-Hill. New York. 


WEED, W. H. 1888. Formation of travertine and siliceous sinter by the vegetation 
of hot springs. U.S. Geol. Surv. Ann. Rept. 9:613-676. 


5Z MADRONO [Vol. 17 


CHROMOSOME NUMBERS OF SOME 
PHYTOGEOGRAPHICALLY INTERESTING CHILEAN PLANTS 


D. M. Moore 


Within the flora of Chile there are a number of floristic elements of 
which perhaps three may be singled out as having particular phytogeo- 
graphical interest. Two, in North Central and Southern Chile, show ties 
across the equator with, respectively, Southwestern North America and 
parts of the North Temperate region and a third, in South Chile, is linked 
over Antarctica with Australia and New Zealand. Taxonomists are cur- 
rently investigating in detail several groups of species within these ele- 
ments because the inter-continental affinities are, in several cases, at the 
specific level and they are especially relevant to problems of plant evo- 
lution and distribution. 


TABLE 1. CHROMOSOME NUMBERS OF SOME CHILEAN PLANTS. 


SPECIES NUMBER LocaLiTy COLLECTION 
GOODENIACEAE 
Selliera radicans Cav. n=—8*  W. side of Tumbes Pen- Moore 286 
insula, Prov. Concepcion, UCB, US, RSA 
Chile 
COMPOSITAE 
Franseria chamissonis n=18* ca.1.5km.N.of Lirquen, Moore 291 
subsp. bipinnatisecta Prov. Concepcion, Chile UCB, US, RSA 
(Less.) Wiggins & 
Stockwell 


Adenocaulon chilense Less. 2n=46** ca. 200 m., Perez Rosales Moore 327 
Pass, Prov. Llanquihue, UCB, US 
Chile 

Adenocaulon bicolor Hook. 2n=46** ca. 400m., near Corvallis, Chambers 1535 
Benton County, Oregon, LA 


U.S.A. 
Adenocaulon bicolor Hook. 2n=46** Shinano-Oiwake, Mt. I. Fukuda 
var. adhaerescens Asama, Nagano Prefec- in October, 1960 
Makino ture, Honshu, 


Japan, ca. 1000m. 


* Meiosis in pollen mother cells. 
** Mitosis in root tips. 


The three Chilean species for which data are recorded in this note are 
all comparable in having the same chromosome numbers as have been 
determined for them or their close relatives in other continents. Thus, 
as in the counts for Chilean specimens presented in Table 1, Selliera 
vadicans Cav. has eight bivalents at meiosis in plants from New Zealand 
(Hair and Beuzenberg, 1960) and Franseria chamissonis Less. subsp. 
bipinnatisecta (Less.) Wiggins and Stockwell forms eighteen bivalents 


1963 | VICKERY, MUKHERJEE & WIENS: MIMULUS D0 


in Californian material (Wiggins and Stockwell, 1937). Representatives 
of three disjunct taxa of the genus Adenocaulon have all been counted for 
the first time (Table 1) and all share a somatic complement of forty-six 
chromosomes. Further studies of relationships within this genus are in 
progress. 

I wish to thank Mr. Donald Kyhos for the preparation showing meiosis 
in pollen mother cells of Franseria. Dr. Kenton L. Chambers, Oregon 
State University, Corvallis, and Mr. I. Fukuda, Tokyo Women’s Chris- 
tian College, Japan, very kindly collected the seed of Adenocaulon. My 
collecting trip to Chile was financed from National Science Foundation 
Grant No. G-13518 made to Dr. Harlan Lewis. 


Department of Botany, 
The University, 
Leicester, England. 


LITERATURE CITED 


Hair, J. B. and E. J. BEUZENBERG. 1960. Contributions to a chromosome atlas of the 
New Zealand Flora. 4. Miscellaneous families. N. Z. Jour. Sci. 3:432-440. 
Wiccins, I. L. and P. STOCKWELL. 1937. The maritime Franseria of the Pacific Coast. 
Madrono 4:119-120. 


CHROMOSOME COUNTS IN SECTION ERYTHRANTHE 
OF THE GENUS MIMULUS (SCROPHULARIACEAE). II’ 


RoBERT K. VICKERY, JR., BARID B. MUKHERJEE 
AND DELBERT WIENS 


This investigation formed part of our long range biosystematic study 
of section Erythranthe (Vickery, 1956; Vickery, Mukherjee, and Wiens, 
1958). It had two purposes. The first was to determine the chromosome 
numbers of two rare species, Mimulus eastwoodiae Rydberg and M. nel- 
son Grant. The second was to analyze the genome homologies of the 
more common species of the section. 

For the cytological portion of the investigation, buds expected to con- 
tain the desired stages of meiosis were placed for 24 hours in a freshly 
prepared fixative consisting of 3 parts absolute ethanol to 1 part glacial 
acetic acid saturated with ferric acetate. The anthers were then dissected 
from the buds, squashed, and lightly stained in aceto-carmine. For each 
determination, the chromosomes of ten or more cells were carefully 
studied and counted under a phase contrast microscope. Many of the 
configurations were recorded with the aid of a camera lucida. 

Both M. eastwoodiae and M. nelsoni were found to have n=8 chrom- 
osomes (fig. 1 and table 1) as do the three more common species of the 
section, M. cardinalis Douglas, M. lewisti Pursh, and M. verbenaceous 


1 This work was supported by grants from the National Science Foundation and 
the University of Utah Research Fund. 


54 MADRONO LVol. 17 


(Greene (Brozek, 1932; Vickery, Mukherjee, and Wiens, 1958). Mimulus 
eastwoodiae (fig. 2) is known from only six or seven localities in southern 
Utah and northern Arizona. The populations occur in shaded seeps in the 
desert at elevations below 4,500 feet. Generally, suitable seeps are on 
perpendicular or overhanging sandstone walls. In contrast, M. nelsoni 
(fig. 2) grows by small mountain streams in the cloud forest, 8,000 feet 
or above, in the central portion of the Sierra Madre Occidentale of Mex- 


5848 6211 3423 


— 
20 10 O 10 micra 


Fic. 1. Camera lucida drawings of meiotic chromosomes of M. eastwoodiae (5848), 
M. nelsonii (6211), and the Fy hybrid (3423) of M. cardinalis (5311) « M. verbe- 
naceous (5924). All pollen mother cells are at or near metaphase I. 


TABLE 1. CHROMOSOME CouUNTS IN MIMULUS, SECTION ERYTHRANTHE. 


A. CULTURES OF NATIVE SPECIES. 


n=8 M. eastwoodiae 
Arches National Monument, Grand County, Utah, altitude 4,200 feet, 
Vickery 339 (5848). 
n=8 M. nelsonii 
Crest of Sierra Madre on Durango-Mazatlan Road, Durango, Mexico, 
altitude 9,000 feet, Wiens 2642 (6211). 


B. INTERSPECIFIC AND INTERPOPULATION HyprIDs.” 


n—=8 Fy (3424) of M. verbenaceous (5924) * M. cardinalis (5316). 

n=8 Fy, (6031) of M. lewisii (5875), Wasatch Mountains form) x M. lewisii 
(5032), Sierra Nevada form). 

—8 Fy, (3423) of M. cardinalis (5311) * M. verbenaceous (5924). 

—8 Fy, (3425) of M. cardinalis (5031) X M. lewisii (5875, Wasatch Moun- 
tains form). 

n=8 Fo (3502) of M. cardinalis (5077) & M. lewisii (5051, Sierra Nevada 
form). 


2 The chromosomes showed regular pairing and/or regular segregations in all cases. 


1963 | VICKERY, MUKHERJEE & WIENS: MIMULUS ap) 


6211 fe 


| 
C 


Fic. 2. Photographs of the two rare species of Mimulus studied in this investiga- 
tion, M. eastwoodiae Rydberg (5848, UT 28,932) and M. nelsoniz Grant (6211, 
UT 54,858). 


< 


mM. 


ico. The full range of this unusual and strikingly beautiful species is not 
known. We collected it near the crest of the Durango-Mazatlan road, 
which is only the second locality reported (Grant, 1924). 

For the cytogenetic part of the investigation, each interspecific com- 
bination was made reciprocally and was repeated using different cultures 
of the parental species. From one to ten flowers were carefully hand- 
pollinated for each combination. The putative hybrid seeds were sown 
and the resulting seedlings carefully checked to verify their hybrid nature. 
The hybrids that were investigated cytologically included combinations 
involving the three major species M. cardinalis, M. verbenaceous, and 
M. lewis (table 1). They included the two distinctive forms of the latter 
species. In all cases the hybrids showed regular bivalent chromosome asso- 
ciations at metaphase I and normal segregations of the chromosomes in 
the later stages of meiosis. 

The results of the present investigation taken in conjunction with the 
previous study (Vickery, Mukherjee, and Wiens, 1958) suggest that the 
common chromosome number of section Erythranthe is n=8. The regular 
bivalent association and normal segregation of the chromosomes of the 
hybrids involving four of the main taxa suggest the presence of essen- 
tially homologous genomes in these forms and possibly throughout the 


56 MADRONO [Vol. 17 


section. Apparently, evolution in section Erythranthe is proceeding prin- 
cipally by the accumulation of diverse genes in the various populations 
and species rather than by the accumulation of chromosomal differences. 


Department of Genetics and Cytology 
University of Utah, Salt Lake City, Utah 


Department of Obstetrics and Gynecology 
Columbia University, New York City, New York 


Department of Biology 
University of Colorado, Boulder, Colorado 


LITERATURE CITED 


Brozek, A. 1932. Mendelian analysis of the “red-orange-yellow” groups of flower 
colours in Mimulus cardinalis Hort. Preslia 11:1-10. 

GranT, A. L. 1924. A monograph of the genus Mimulus. Ann. Missouri Bot. Gard. 
11:99-388. 

PENNELL, F.W. 1951. Scrophulariaceae zm Illustrated Flora of the Pacific States by 
Leroy Abrams. Stanford Univ. Press. Vol. III, pp. 688-731. 


VickErY, R.K., Jr. 1956. Data on intersectional hybridizations in the genus Mim- 
ulus (Scrophulariaceae). Proc. Utah Acad. 33:65-71. 


VicKERY, R.K., Jr., B. B. MUKHERJEE, AND D. WIENS. 1958. Chromosome counts 
in section Erythranthe of the genus Mimulus (Scrophulariaceae). Madronfo 14: 
150-153. 


AN ANALYSIS OF VARIATION IN VIOLA NEPHROPHYLLA 
NorMAN H. RUSSELL AND FRANK S. CROSSWHITE! 


The genus Viola is well known to be one of the more taxonomically 
“difficult” of the temperate angiosperms. Though in North America the 
“species” were sorted out in what appeared at the time to be a satisfac- 
tory manner (Brainerd, 1920), subsequent studies have shown that their 
limits are anything but clear. In particular regions it is possible to distin- 
guish separate forms easily; in others there is so much morphological 
and ecological variability that distinct forms or even morphological types 
are very difficult to describe. Polyploidy, introgression, genetic drift in 
isolated populations, and other hypotheses have been used to explain this 
situation. 

More important than the explanation of this morphological and physio- 
logical variation is the accurate and objective description of it. A method 
for the more objective comparison of units (individuals and aggregations 
of individuals) has been suggested by the senior author elsewhere (Rus- 
sell, 1961, 1962) and is used in the present analysis and description. It 
consists of the preparation and correlation of multiple pair comparisons, 


1 This research was supported by National Science Foundation grants G—4994 
and G-12113. 


1963 | RUSSELL AND CROSSWHITE: VIOLA oi, 


ei 
SGP 3 aga) snl \ wo Ree oJ 
50 Se é ty ee ae) Ree Vaan Sen ea | 
Se ile ; nf de Sia if ee a p oe Nie vl Nr 
: - i : We ee 
co ~— <A ws \ Hi oa ss 
; . soo ae a poh! if V “= | 
c, 2 - @ 0 e \, “\ a - he 
Sooo a eS i 
Pd ay. SS a ane so eee _ Hoare ee 
ee e ‘ : L 
ee i ; e A ae a ° Ce; = od % OF 
~ 40 a, teas , oe | “fee oo! ° oe % 
5.08 ee - - i) 3 e fa e Ve e mae 
bd 2 oo i ' e e es zi i) 
r ; O ee ' e N eo ogee oF 
' oo? ,e¢ ; of rae 5 we re? bl 
er e HH \ - e, 
\ é be 
\ ek aia ee ; 
oe e .° ‘“ “ 
30 Dd wee one pee 
. \e J %e = O. 
. | 
Bais \ 
Shae Dawe ON 


Fic. 1. Geographical distribution of Viola nephrophylia Greene. 


using an index similar to that described by Anderson (1936), and is 
described in greater detail below. 

Becker (1925), in the generally accepted classification of the genus 
Viola, recognized eleven sections, the most complex of which is the sec- 
tion Plagiostigma. This section was further divided into several subsec- 
tions, the best known of which is the subsection Boreali-americanae, a 
group of perhaps twenty-six so-called “species,” found exclusively in 
North America. Of these, only a single one, Viola nephrophylla Greene 
(Pittonia 3:144—-45. 1896), occurs to any extent in the Rocky Mountains, 
the remainder being found almost exclusively in eastern and central North 
America. Viola nephrophylia has the largest range of any of the stemless 
blue violets. A map of its distribution (fig. 1) is based upon the examina- 
tion of specimens from about sixty herbaria and represents the subjec- 
tive decisions of the senior author on the basis of ten years of field and 
herbarium study of the genus Viola. Each dot, therefore, represents the 
place of collection of a herbarium specimen that, in his opinion, would key 
to V. nephrophylia. As is true of all such maps, it suffers from these sub- 
jective decisions and, therefore, must represent, perhaps, only an approxi- 
mation to reality. The species, very likely, also occurs in Mexico, but we 
have not seen specimens from there. In many parts of its range, particu- 
larly where it does not grow with other kinds of stemless blue violets, 
it is morphologically distinct. However, in many situations in eastern 
North America, it is difficult to distinguish from related violets. 

The habitat of V. nephropAylla varies widely in the Rocky Mountains. 
Usually we found it growing in moist, grassy, grazed fields, frequently 
in the shade of willows. Other populations were found along the shaded, 
sandy edges of canyon streams. In the eastern United States it grows 
both in open, grazed, poorly drained meadows and along the rocky shores 
of lakes in glaciated country. Figure 2 indicates the general appearance 
of this violet during the spring flowering period. 


MADRONO [Vol. 17 


On 
oo 


Fic. 2. Viola nephrophylla, spring appearance, < %rds. 


COLLECTION METHODS 


Fifteen samples were taken by the junior author during the growing 
season of 1960, wherever they could be found, in rather extensive travels 
through Arizona and Colorado. Their locations are shown in figure 3. 
The number of specimens taken at each location varied from sixteen to 
fifty, depending upon the size of the local population. Ordinarily fifty 
specimens were obtained, but in some instances this was not possible. 
Plants were collected no closer together than six feet, to lessen the pos- 
sibility of sampling two members of the same clone. They were usually 
measured or scored while fresh, but in some instances the plants were 
washed, pressed, and dried before examination. All the plants measured, 
and the measurement data, are deposited in the herbarium of Arizona 
State University. 


1963 | RUSSELL AND CROSSWHITE: VIOLA 59 


GUNNISON 


(ees 


(COL 


GUNNISON 


SAGUA 


Fic. 3. Locations of population samples in Arizona and Colorado. 


After a preliminary inspection of herbarium material, those character- 
istics showing the most conspicuous differences between individuals were 
chosen for analysis. The following characters were measured or scored 
as indicated: 

1. Length of the lamina of the largest mature leaf. 

2. Breadth of this lamina. 

3. Distance from the apex of this lamina to one of the basal lobes. 

4. The angle made by one-half of the apical margin of this leaf with 

the horizontal. 


60 MADRONO [Vol. 17 


TABLE |. DISTRIBUTION (EXPRESSED AS PERCENTAGES) OF LAMINA LENGTH/BREADTH RATIOS, 
AND INDEX VALUES ASSIGNED FOR THE FIFTEEN POPULATIONS ANALYZED. 


Hybrid Index Scores 


O I 2 
Crosswhite 
Coll. No. 50259 60-69 70579 80-89 90-99 100-109 -LI0-L1I9.—s 120-129 130-159 
14504 2.00 18.00 30.00 
970A f°) fo) fe) 
965 12.50 56.25 31.25 
971 ) Co) 35.00 
1239 2.00 16.00 50.00 
62 f°) 26.67 40.00 23.33 10.00 0) (e) fo) fc) 
843 fe) f°) 16.00 48.00 24.00 8.00 O (o) 4.00 
1064 ) 8.00 20.00 36.00 32.00 4.00 fo) ) fo) 
1065 Oo f°) 12.00 60.00 18.00 8.00 2.00 (o) 0) 
1397 0) 17.50 65.00 17.50 ) ) 0) ) fo) 
1304 ) 4.00 36.00 44.00 16.00 0) ) fo) fo) 
1062 ) Co) 26.67 53.33 13.33 f°) 0) 6.67 ) 
N29 0) 4.00 14.00 30.00 48.00 4.00 (o) f°) fe) 
1087 ) 2.22 2.22 2.0.00 40.00 26.67 6.67 2.22 ) 
/054 ) ) 4.00 24.00 28.00 24.00 16.00 4.00 fe) 


5. Number of teeth on one-half the margin. 

6. Pubescence of the upper lamina surface (scored as 0 = glabrous, 
1 = slightly pubescent, 2 = moderately pubescent, and 3 = heavily 
pubescent). 

7. Pubescence of the lower lamina surface (scored as indicated for 
the upper surface). 

8. Pubescence of the margin of the lamina (scored as 0 = glabrous, 
1 = hairy over half or more of its extent). 

9. Pubescence of the petiole (scored as 0 = glabrous, 1 = 10 or more 
hairs present). 


These leaf characteristics are those used by the senior author in studies 
on other stemless blue violets (Russell, 1955, 1956a, 1956b). It might 
have been desirable to measure other plant structures also, but either 
no conspicuous differences were noted in these or they were not present in 
all the samples. During the summer, when the majority of the samples 
were taken, no open flowers are produced, and, at times, no cleistogamous 
flowers or fruit. Rhizome differences, though present, were small and dif- 
ficult to measure accurately. 


PREPARATION OF THE INDEX 


After compilation of the data (nine measurements or scores for each 
specimen), three ratios were calculated: lamina length/lamina breadth; 
lamina length/length from lamina apex to lobe; and lamina length/lami- 


1963 | RUSSELL AND CROSSWHITE: VIOLA 61 


TABLE 2. INDEX VALUES ASSIGNED FOR EACH OF THE 
FIVE CHARACTERS USED IN THIS STUDY. 


Ranges in Characters 


Character Value O Value | Value 2 


Lamina length/ 
breadth ratio Omos NGOmed S10) |S) 


Lamina length/length 
to lobe ratio SOT ISS) Ome SBOriIg 


Lamina length/twice 
no. lamina teeth pOORmo2 Aa\O= S/S) 


Apical angle of 
lamina ZO So- 40°-49° One coe 


Total lamina* 
pubescence ‘Sr .s) SyrS) O22 


*Pubescence was scored on an orbitrary scale for both leaf surfaces, 
the margin, and the petiole. The value of 8 represented the greatest 
hairiness found, and that of O complete glabrousness. 


na teeth. The distributions for each of the characteristics and ratios were 
then plotted for each sample. An example of one such distribution is 
shown in Table 1, for the lamina length/breadth ratio (an approximate 
measurement of overall shape). In the event real intra- or intersample 
differences were noted, index values (0, 1, and 2) were assigned to the 
extremes and median conditions found. This is illustrated in Table 1, and 
in Table 2 the index value assignments for all the characteristics used in 
the subsequent analyses are given. 

In Table 3 the collections are arranged in order of the value of the 
mean index, in an attempt to reveal the existence of altitudinal or lati- 
tudinal clines. Total sample variation did not fall into any such pattern, 
although the Arizona samples generally had low values. Only a morpho- 
logical “cline” can be shown, and we were unable to discover any geo- 
graphical or ecological factor to which this could be definitely related. 

The samples at opposite ends of Table 3 show considerable difference, 
enough so that if only these two extremes were known they might be 
called different “species” under present nomenclatural practices in plant 
taxonomy. As an example, the index distributions of two extreme collec- 
tions have been plotted in figure 4, on a percentage basis. The differences 


62 MADRONO [Vol. 17 


TABLE 3. SUMMARY OF DIFFERENCE INDEX SCORES 
FOR FIFTEEN POPULATION SAMPLES. 


Crosswhile No. 

Coll.No. Elev. Latitude Spec. O | 2 3 4 5 6 7 8 9 10 Mean Location 
/450A 5800' 35° 50 hon (ey a2 9S of | 3.12 Oak Creek, Ariz. 
970A 6000' 34°5' 16 4 5 2 4 | 3.63 Lakeside, Ariz. 
965 5800' 31°50! 16 A Ae ee i 3.81 Portal, Ariz. 

971 6000' 34°5' 40 1 7 1 6 9 | 4.48 Bog Creek, Ariz. 
1239 9400' = 38°50! 50 OB Ba 7? ey ey! 4.76 Cement Creek,Colo. 
62 8400' 38°30' 30 ome OVE OF cso { 4.80 Almont, Colo. 

843 6000! 34°20! 25 2 eae 7 ile al \ 5.32 East Verde, Ariz. 
/064 9200' 39° 25 2a 5 6 4 2 5.44 Gothic, Colo. 
/065 9300' 39° 50 10) 10) 1/2 13s 2 5.90 Gothic, Colo. 

1397 7900' 38°20! 40 6 7 12 7 8 6.10 Gunnison, Colo. 
/304 9200' 40° 25 2 8 6 4 4 | 6.12 Nederland, Colo. 
1062 8400' 38°30' 15 2 A 4e ee lee: 6.13 Almont, Colo. 
M29 9000' 38°50! 50 fee Ae He Te eeg 6.54 Crested Butte,Colo. 
/087 9500' 39° 4! WS eR ee A 7.29 Gothic, Colo. 
1054 9400' 39° 25 3 5 10 7 8.84 Gothic, Colo. 


between these two curves were analyzed in the following manner (Russell, 
1961, 1962): 

1. Percentage of the distance from the mode of one aggregation to the 
extreme value of the scale assigned to the other aggregation. 

2. Percentage of the distance on the total index scale that the range 
of values for the aggregation does not cover. 

3. Percentage of the total index for the range discontinuity (plus val- 
ues ) between the pair of samples or the total overlap (minus values) 
between them. 

4. Percentage of the total index for the distance between the modes 
of the two curves. 

These four descriptive features of the curves are analyzed in such a way 

as to show the greatest morphological separation of the two populations. 


TABLE 4. MATRIX OF TOTAL DIFFERENCE INDICES FOR ALL SAMPLE COMPARISONS, 
TO BE READ HORIZONTALLY. 


Crosswhite 

Coll No” I4SQA 970A (905 97) 1239 “M162 2845" 004 1065, 51807 904 Oe Ve Oe Coa Meany 
/450A x -29 -29 «14 fe) 44 39 44 89 63 67 73 72 (WO 150 48.50 
970A - 43 x -33 «13 O 50 42 50 100 7I 75 8l 8l 122 167 55.43 
965 -43  - 83 x 13 ) 50 42 50 100 7 ie 8I 8 122 167 51-86 
97/ - 1 =43 43 X -15 fh 14 -8 5! 15 26 S20 S278 eles 16.00 
1239 14 = fan eaulie e229) Xx OF F=I5 fe) 50 14 25 31 3 78 122 20.50 
62 10 -l2 =“|20-37 4-30 aXe 25 Elo (o) Oo -25 -19 -I9 33. 78 = -6.00 
843 39 14 14-14-1525 X  -50 =| -42 | -6 18 33 «O«77 3.07 
1064 49 29 2913 O =i2 0 15 x 14-33 Oy 57 -8 50 100 14.93 
1065 100 88 88 63 63 38 38 2 KS) 34 25) 50-14 43 27.57 
1397 87 72 72 «44 43 25 15 -8 -33 x | -8 =i, 43 100 31.86 
/304 55 37 38 O13 13-12 13-2 -50 -49 Xeuse5 -8 43 99 10.43 
1062 67 50 50 25 25 0) 2s 2| —6(7 00150 -6r XP es2 28 = 85 8.93 
M29 66 50 50 25 25 fe) O -22 -29 -14 -1I5 -12 Xx Oo 668 13.71 
1087 110 = 100 100 «78 78 55 56 44-14 14 15 7 18 x 8 47.79 


1054 190 190 190 168 167 «145 145 144 100 128 128 l21 118 66 X 142.86 


1963 | RUSSELL AND CROSSWHITE: VIOLA 63 


Two separate analyses must be made, one for each population as com- 
pared to the other. The results of the analyses of the curves shown in 


figure 4 are: 
Crosswhite 965 ........ 64/27/-9/55  Crosswhite 1054............ Total Index—137 
Crosswhite 1054 ........ 82/64/-9/55  Crosswhite 965..........-- Total Index—192 


Average Difference Index—164.5 


Pictorialized scatter diagrams (figs. 5 and 6) illustrate the population 
differences further. 


40 


30 


% 90 


O | 205 4 5 6 7 8 9 #10 
INDEX VALUE 


Fic. 4. Index distributions for two extreme populations; further explanation 
im text. 


Total difference indices were computed for the other possible compari- 
sons and were smaller than the above, grading down to minus values. 
They are shown in Table 4. When all the index distributions were con- 
verted to percentages and summed, a curve was obtained (fig. 7) which 
does not seem to indicate the presence of two or more types or “‘species”’ 
in this complex, but instead indicates a series of variable populations cen- 
tering about an average overall morphological condition. A more pene- 
trating analysis, using techniques such as those suggested by Rogers and 
Tanimoto (1960) might indicate whether or not more than one morpho- 
logical type is present, and such an analysis is being planned. Other taxo- 
nomists have given specific rank to certain aberrant or differing types in 
the range of Viola nephrophylla; namely V. arizonica Greene, V. cognata 
Greene, V. prionosepala Greene, V. McCabeiana Baker, and V. Clauseni- 
ana Baker. We do not interpret the present descriptive data to support 
the recognition of different morphological types or “species.” 


64 MADRONO [Vol. 17 


65 35 


c Spe 


5 
er 
; 4 x if 
55 
© 25 44 
4 
q 
z 
50 Z 20 ity 
= FSC 1054 
45 ' + 
i? LAMINA LENGTH 
40 
aL ' Fig. 6 
z 9) 
2 35 
Ww 
a 
o 
z 30|[ | 30 
= 
aq 
* dtd 
25 aD FSC 965 20 
% 
10 
20 . 
15 ro) 
5 20 25 30. 35 40 45 o 12 3 4 5 6 7 8 9 0 
LAMINA LENGTH INDEX VALUE 
Fig. 5 Fig. 7 


Fics. 5-6. Pictorialized scatter diagrams of Viola nephrophylla populations: 
5, Crosswhite 965; 6, Crosswhite 1054. Fic. 7. Distribution (converted to percentage 
values) of indices for all populations. 


A sample collected in Oak Creek Canyon, Arizona, during September, 
1960, deserves special mention (Crosswhite 1450A).The specimens differ 
(fig. 8) only in the relationship of lamina breadth to lamina length, the 
ratios exposing approximately equal numbers of plants with leaves wider 
than long and with leaves longer than wide. We have considered that the 
two differing regression lines may be explained either as the result of 
measurement errors, ‘‘juvenile’” and mature leaves having been measured 
in equal numbers, or as the result of a nearly equal distribution of the 
members of a pair of alleles differently affecting lamina growth in the 
area collected. We have no data to test the second hypothesis, but have 
re-examined and remeasured the leaves and have, we believe, eliminated 
the possibility of error in choice of leaves. 

Th taxonomic disposition of such aggregations of plants as that which 
we now call Viola nephrophylia still, of course, remains subject to the 
caprices of taxonomists who, under the present international rules, may 
justify their nomenclatural decisions by reference to their intuitive judg- 
ments. In this exploratory study, considering only a few samples in a part 
of the range of V. nephrophylla, we have submitted what we feel is good 
evidence for the rejection of the synonyms listed earlier for the Rocky 


1963 | RUSSELL AND CROSSWHITE: VIOLA 65 


95 
zi 
-9 
85 
75 - -0 -9-0 
—O 
me 
| $0 T 
65 -4 
as - ~ 9 
55 7 -9 4 to-0 i 
: 4% BB 
S © 
f a) 4 > fo! 
< a6 
z 35 =) 
é é 
25 e 
FSC 1l1450A 


15 25 35 45 55 65 75 85 95 


LAMINA LENGTH 
Fic. 8. Pictorialized scatter diagram for population sample, Crosswhite 1450A. 


Mountain area. More elaborate studies are in progress on populations 
from the eastern and north-central part of the United States. 


Arizona State University 
Department of Botany, Tempe, Arizona 


LITERATURE CITED 


ANDERSON, E. 1936. Hybridization in American tradescantias. Ann. Mo. Bot. Gard. 
23:511-525. 
BECKER, W. 1925. Viola in Engler, A. and K. Prantl, Die Naturliche Pflanzenfamilien, 
ed. 2, vol. 21:363-376. 
BRAINERD, E. 1921. Violets of North America. Vt. Agr. Exp. Sta. Bull. 224. 
RoceErs, R. J., AND T. T. TAnrmorTo. 1960. A computer program for classifying plants. 
Science 132:1115-1118. 
RusseEL1, N. H. 1955. Local introgression between Viola cucullata Ait. and V. septen- 
triontalis Greene. Evolution 9:436—440. 
. 1956a. Techniques for species comparison in violets. Proc. Iowa Acad. 
Sci. 63:157-160. 
. 1956b. Regional variation patterns in the stemless white violets. Am. Midl. 
Nat. 56:491-503. 
. 1961. The development of an operational approach in plant taxonomy. 
Syst. Zool. 10:159-167. 
. 1962. A different approach to taxonomy. Assoc. Intern. Syst., Caen, 
France. 


66 MADRONO [Vol]. 17 


REVIEW 


Flora of New Zealand. By H. H. Arran. Vol. I. Indigenous Tracheophyta: 
Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. Government 
Printer, Wellington. liv + 1085. (April) 1961. $14.70. 

Although it was never published, the first flora of New Zealand was compiled 
nearly two centuries ago by Daniel Solander, a student of Linnaeus. Following it 
were works by notable botanists such as Allan Cunningham, Joseph Hooker, Thomas 
Kirk, and T. F. Cheeseman. This first volume of the newest “Flora of New Zealand” 
by H.H. Allan follows in the best tradition of New Zealand botany and is a distin- 
guished contribution to the world’s botanical literature. 

The part issued includes all vascular plants indigenous to New Zealand except 
monocotyledons, presumably to be the subject of a later volume. Solander’s manu- 
script described 360 species of vascular plants; Allan deals with 1457 species and 
280 varieties of native vascular cryptogams, gymnosperms, and dictoyledons. If a 
rough estimate of the number of monocotyledons were added, the total number of 
native taxa would considerably exceed 2000. Allan originally conceived a revision 
of the second edition of Cheeseman’s “Manual of the New Zealand Flora” (1925) 
which was out-of-date and had long been unobtainable, but it became clear that a 
complete reworking of the flora was necessary. Unfortunately, Allan did not live to 
see the publication of the work to which he had devoted so much of his time and 
energy. It was completed and guided through the press under the able and sympa- 
thetic direction of Lucy B. Moore. Most of the Flora was prepared by Allan, but sev- 
eral groups were treated by Miss Moore, including difficult genera such as Myosotis 
and all of Hebe except the whipcord hebes, and others were handled by M. B. Ashwin, 
who contributed sections on Euphrasia, the whipcord hebes, Parahebe, and Pygmea. 

The preface to the work is followed by several pages of useful annals devoted 
to bibliographic citations and short notes on botanical literature relevant to New 
Zealand covering the period from Solander’s manuscript of 1769 to papers published 
in 1958. A short section discusses the New Zealand Botanical Region, taken to in- 
clude the Kermadec Islands, Three Kings Islands, Chatham Islands, the subantarctic 
islands (Antipodes, Aucklands, Campbell, Macquarie, and the Snares), and the three 
main islands of New Zealand. Next are pages explaining abbreviations used in the 
text, a list of authors of New Zealand taxa, and a synopsis of the classes and orders 
of the plant groups covered by the Flora. The arrangement of ferns follows a system 
by Holttum and that of the dicotyledons follows the first edition of Hutchinson’s 
“The Families of Flowering Plants.” The artificial keys to the families of dicotyle- 
dons and to the genera are praiseworthy for their simplicity; they seldom use more 
than single pairs of characters for making a choice. If satisfaction fails here, there 
are additional generic keys following each family listing in the main portion of the 
book. 

New taxa are circumscribed in English in the text with their Latin diagnoses 
appended in a special section. Also included are a glossary covering technical termi- 
nology used in the descriptions and a series of drawings illustrating terms describing 
leaf morphology. The alphabetical list of Maori plant names will be of special use 
to visitors, since even professional botanists in New Zealand often refer to plants by 
names such as kowhai, ti, rimu, manuka, puriri, and pohutukawa. Some of these 
names have inspired genera such as Hoheria from houhere, Tupeia from tapia, 
Corokia from korokio, and additional specific names such as totara, maire, and 
taraire. A lengthy but useful group of supplementary notes precedes the index, the 
final section of the book. These notes make some typographical or factual correc- 
tions, additional comments, and amendments of parts of the text based on recent 
collections or newly-published monographs. They bring the work up to date through 
1960 and correct virtually all errors in the text. 

The original and critical nature of the Flora is reflected by the fact that three 
new genera, 29 new species, and 61 new varieties are described in it. The new genera 


1963 | REVIEW 67 


are: Kirkophytum, which includes two species formerly referred to Stilbocarpa 
(Araliaceae) ; Neopanax, to accommodate species formerly found in Nothopanax 
(Araliaceae) ; and Kirkianella as a monotypic genus for what has unhappily been 
called Crepis novae-zelandiae, a composite of uncertain affinities. Other genera have 
been dropped from the New Zealand flora, including Leucopogon, which Allan rele- 
gated to Cyathodes. 

In this volume, 290 genera are recognized. Northern hemisphere botanists may be 
surprised to find familiar genera such as Myosotis (34 spp.), Ranunculus (43 spp.) 
and Epilobium (50 spp.) so amply represented. The largest genus is Hebe, with 79 
species recognized—this even after the removal from it of peripheral entities such as 
Pygmea and Parahebe. Next in size are Celmisia (58 spp.), Epilobium (50 spp.), 
and Coprosma (45 spp.). Nearly two-thirds of the three dozen endemic genera are 
monotypic. The vascular flora as a whole is about 80 per cent endemic according 
to an estimate made some years ago by Cockayne. Each island in the botanical 
province has its own endemic flora, and a number of these species have extremely 
limited ranges, being known only from a few or single colonies. Of particular phyto- 
geographical interest are the Three Kings Islands, only 8 square kilometers in area, 
which support about a dozen endemic species and two endemic genera. One of these 
genera, Plectomirtha (the only representative of the Anacardiaceae in New Zealand), 
is known only as a single tree! At least two species, Tecomanthe speciosa (the only 
member of the Bignoniaceae in New Zealand) and Alectryon grandis (Sapindaceae) 
occur only as single individuals on these islands! 

Introduced plants were not recorded in this volume, but in view of their large 
number (576 species reported by Cheeseman in 1925) their omission is understand- 
able. Since many of these introduced species belong to genera not otherwise found in 
New Zealand, there is little chance that they will be confused with indigenous taxa. 
Species such as Oxalis corniculata and Picris hieracioides were included because they 
were collected early in the 19th century only shortly after settlement of New Zealand 
began, but they are probably introductions. Others, such as Sonchus asper, presum- 
ably arrived with the Maori, since they were collected by the first Europeans to land 
in New Zealand. 

In the text, the species are given ample descriptions with a full author citation, 
place of publication, type locality, and notes on range and habitat. Synonymy is not 
complete, but covers most names relevant to New Zealand material. Many regional 
floras serve only as guides to identification and little else, giving no hint as to the 
botanical problems present in the area covered. But, as pointed out in the preface to 
the Flora, “it is recognized that many species are inadequately known and a second, 
but not necessarily secondary, objective has been to indicate directions in which 
more investigations are needed,” which objective has been admirably fulfilled. Par- 
ticularly outstanding are the copious notes discussing various problematical specimens 
in herbaria, and field observations which will be of great help to botanists dealing 
with these groups in the future. One of the most valuable and novel features of the 
Flora is the inclusion of sections augmenting the generic treatments with comments 
on heteroblasty, sexual expression, polymorphy, hybridism, horticultural forms, taxa 
of uncertain position, taxonomic synopses, notes on special problems, synopses of 
growth forms, and general field observations and remarks reflecting Allan’s wide 
knowledge of plants in the field and the literature concerning them. 

A recurrent theme in the Flora is the suggestion that hybridization is responsible 
for the variation within many plant groups. Although both men had published on 
the subject separately, in 1934 Leonard Cockayne and Allan published a list of nearly 
500 wild species-hybrids in the New Zealand flora. This paper must have met con- 
siderable skepticism and opposition from botanists in other areas of the world, where 
interspecific sterility barriers are the sine qua non of orthodox biosystematy. Never- 
theless, Allan received support from subsequent workers in the dominion and else- 
where, and continued to maintain that hybridization is a potent force in the evolu- 
tion of the New Zealand flora. It is evident from numerous well-documented exam- 


68 MADRONO [Vol. 17 


ples that actual, rather than apparent, hybridization does occur and is largely respon- 
sible for the taxonomic complexity of these groups. 

The Flora is illustrated with several excellent line drawings by Nancy Adams, 
who also designed the attractive dust jacket. To achieve a volume of handbook size, 
very thin paper was used; the nearly 1100 pages make a book only 2 cm. thick. 
The copious notes are printed in 6-point type, which seems too small to be read 
comfortably for very long. The printing and binding are very well done. The small 
size of the volume should not belie the riches it contains. RoBERT ORNDUFF, Depart- 
ment of Botany, Duke University, Durham, North Carolina. 


NOTES AND NEWS 


CHROMOSOME NUMBERS IN CROSSOSOMA. Since the relationships of the small fam- 
ily Crossosomataceae have been a subject of discussion, it is of interest to record the 
chromosome numbers of two species of the only genus. Crossosoma californicum Nutt. 
is confined to Santa Catalina, San Clemente, and Guadalupe islands off the coast of 
southern California and Baja California, whereas C. bigelovii Wats. is found about 
the margins of the Sonoran Desert in California, Arizona, Baja California, and So- 
nora. Crossosoma parviflorum Rob. & Fern. and C. glaucum Small, both described 
from Arizona, are probably not distinct from C. bigelovii at a specific level, and so 
the family probably consists of only two species. The chromosome number of C. cali- 
fornicum was determined from buds collected from Pebbly Beach Canyon near the 


Fic. 1. Chromosomes of Crossosoma at meiotic metaphase I, a, C. californicum ; 
b, C. bigelovii. Both figures * 2600. 


water purification plant, Santa Catalina Island, Los Angeles County, California 
(Taylor & Ornduff 4383, UC); from material propagated at Rancho Santa Ana 
Botanic Garden, taken from a collection (Wolf 1487, RSA; fig. 1a) made at the junc- 
tion of Pebbly Beach and Renton Mine roads, Santa Catalina Island; and from 
material of undetermined origin cultivated in the East Bay Regional Parks. All of 
these collections had a gametic chromosome number of n=12, with no meiotic irreg- 
ularities observed, as did a single collection of C. bigelovit from Morongo Valley, 
Riverside County, California (Davis 105, RSA; fig. 1b). The twelve pairs of rela- 
tively small chromosomes found in these plants are markedly different from the five 
very large pairs found in Paeonia (Ranunculaceae), with which Crossosoma has been 
allied. They are, however, more or less similar to the chromosomes found in a num- 
ber of other families of angiosperms. PETER H. RAvEN, Division of Systematic Biology, 
Stanford University, California, and Marion S. Cave, Department of Botany, Uni- 
versity of California, Berkeley. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Second Edition, 1954). 

Articles may be submitted to any member of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: ~ 


Volume I, 1916-1929 . . . . = . $5.00 
Volume IT, 1930-1934 . . . . = .  §.00 
Volume IIT, 1935-1936 oi arn Mh ert eo) 
Volume IV, 1937-1938 2. % "ia 35208 
Volume V, 1939-1940. 2 ... 4%. 5.00 
Volume VI, 1941-1942 ote cate akee MORO 
Volume VII, 1943-1944 . . . . 5.00 
Volume VITI, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 Re a Onl) 
Volume X, 1949-1950... 0 .0) « 2 9 ag00 
Volume XI, 1951-1952 Re ee ee te 0, 
Volume XII, 1953-1954 . . . . 8.00 
Volume XIII, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . .  . £8.00 
Volume XV, 1959-1960 oe, Ri em EO 
Volume XVI, 1961-1962 2 ican a. e200 


Single numbers: 


Volumes and il 2)" 209 ae eee 50 
Volumes II through XIV. . ..... 1.00 
Volume Sev toc: eee ee 25 
Volume’ XVI. 3. es eet eee #250 


The subscription price of MapRoNo is $6.00 per year. 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 membership in the California Botanical Societv on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


x ee ee 
6 Sea SS SS 


Aes 
MADRONO 


VOLUME 17, NUMBER 3 JULY, 1963 


Contents 
PAGE 


A NEw SPECIES OF ISOPYRUM ENDEMIC TO THE QUEEN 
CHARLOTTE ISLANDS OF BRITISH COLUMBIA AND ITS 
RELATION TO OTHER SPECIES IN THE GENUS, J. A. 
Calder and R. L. Taylor 69 


CYTOPHYLETIC ANALYSIS OF HYMENOXYS ODORATA: A 
RECAPITULATION, B. L. Turner (th 


A NOoTE ON TAXONOMIC CHARACTERS IN LOLIUM, 
Frank C. Vasek and J. Kirk Ferguson 79 


CLEISTOGAMY IN THE MALVACEAE, Paul A. Fryxell 83 


SWALLENIA, A NEW NAME FOR THE CALIFORNIA GENUS 
ECTOSPERMA (GRAMINEAE), Thomas R. Soderstrom 
and Henry F. Decker 88 


Reviews: Irving W. Knobloch and Donovan S. Correll, 
Ferns and Fern Allies of Chihuahua, Mexico (Rolla 
M. Tryon); K. R. Sporne, The Morphology of Pteri- 
dophytes (Job Kuijt); C. Leo Hitchcock, Arthur 
Cronquist, Marion Ownbey, and J. W. Thompson, 
Vascular Plants of the Pacific Northwest (Robert 
Ornduff) 89 


NoTES AND News: NEW DISTRIBUTIONS FOR FOUR 
GRASSES IN OREGON, Kenton L. Chambers and La Rea 
J. Dennis; CALypso BULBOSA IN THE SANTA CRUZ 
Mountains, Thomas A. Crandall; NoTES ON TETRA- 
COCCUS AND CHILOPSIS IN BAJA CALIFORNIA, MEXICO, 
Edmund C. Jaeger 91 


A WEST AMERICAN JOURNAL OF BOTANY 


‘PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofo, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


EpcAR ANDERSON, Missouri Botanical Garden, St. Louis 
LymMAN BENSON, Pomona College, Claremont, California 
HERBERT F. COPELAND, Sacramento College, Sacramento, California 
Joun F. Davipson, University of Nebraska, Lincoln 
MivprepD E. MATHIAS, University of California, Los Angeles 24 
ROBERT ORNDUFF, University of California, Berkeley 
MaArRIon OwNnBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
Ira L. Wiccins, Stanford University, Stanford, California 


Editor—JoHN H. THoMAS 
Dudley Herbarium, Stanford University, Stanford, California 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Herbert L. Mason, Department of Botany, University of California, 
Berkeley. First Vice-President: Paul C. Silva, Department of Botany, University of 
California, Berkeley. Second Vice-President: Robert F. Hoover, California State 
Polytechnic College, San Luis Obispo. Recording Secretary: Mary L. Bowerman, 
Department of Botany, University of California, Berkeley. Corresponding Secretary, 
Margaret Bergseng, Department of Botany, University of California, Berkeley. 
Treasurer: Douglas M. Post, Biology Department, San Francisco State College, San 
Francisco, California. 


1963 | CALDER & TAYLOR: ISOPYRUM 69 


A NEW SPECIES OF ISOPYRUM ENDEMIC TO THE QUEEN 
CHARLOTTE ISLANDS OF BRITISH COLUMBIA AND ITS 
RELATION TO OTHER SPECIES IN THE GENUS! 


J. A. CALDER AND R. L. TAYLOR 


In 1957, a party composed of D. B. O. Savile and the authors carried 
out a botanical survey of the Queen Charlotte Islands from late May until 
the latter part of August. During the course of the survey, several collec- 
tions of a new taxon belonging to the genus /sopyrum were made in the 
Queen Charlotte Ranges. This paper is devoted to the description of this 
new species and its relation to other members of the genus. 

Four species of /sopyrum are presently recognized in North America. 
Three are found in western United States and their distributions include 
southern Washington, Oregon, and California. The fourth species, /. biter- 
natum, is restricted to eastern United States and Canada. The new 
species, J. savilez, is completely disjunct from these four taxa and is found 
only in a small area of the Queen Charlotte Islands. 

Isopyrum savilet is the fourth endemic to be described from this region 
as a result of the 1957 survey, the others being: Saxifraga taylori (Calder 
and Savile, 1959), Saxifraga punctata ssp. carlottae (Calder and Savile, 
1960), and Ligusticum calderit (Mathias and Constance, 1959). Another 
distinct endemic, Senecio newcombet, was previously described by E. L. 
Greene in 1897 from a collection made by C. F. Newcombe during a 
survey of the Queen Charlotte Islands in the same year. In addition to 
these endemic taxa, there still remain a few undescribed entities in our 
collection from the alpine and subalpine areas in the Queen Charlotte 
Ranges. Although the endemics are few in number, the degree of endem- 
ism is high for such a small flora. The phytogeographic significance of 
this endemism will be fully discussed in a forthcoming treatment of the 
flora of the Queen Charlotte Islands. 

We would like to express our appreciation to the curators of the follow- 
ing herbaria for the loan of specimens or the opportunity to examine 
material in their respective institutions: University of California, Berke- 
ley; University of Oregon; Peck Herbarium, Willamette University; 
University of Washington; Washington State University; University of 
Wyoming; New York Botanical Garden; and Gray Herbarium, Harvard 
University. We would like to express our appreciation to the artist, Miss 
C. Mentges for the excellent illustrations, to B. Boivin for the Latin 
diagnosis, and to C. Crompton for technical assistance. 

It isa pleasure to name this species after our close friend and colleague, 
D. B. O. Savile, who has collected widely in the Pacific Northwest and 


1 Contribution No. 259 from the Plant Research Institute, Research Branch, 
Canada Department of Agriculture, Ottawa, Ontario. 


Maprono, Vol. 17, No. 3, pp. 69-92. July 15, 1963. 


70 MADRONO [Vol. 17 


whose suggestions and stimulating discussions have provided a greater 
insight into the botanical problems of this region. 


KEY TO THE NORTH AMERICAN SPECIES OF ISOPYRUM 


Tepals 3.5-6.0 mm long, veins few and prominent; filaments flat, narrowly tri- 
angular; follicles stipitate. Oregon and northern California............ I. stipitatum 
Tepals 7.0-17.0 mm long, veins many and inconspicuous; filaments filiform, clavel- 
late to clavate; follicles sessile. 
Leaflets puberulent beneath; flowers in a cyme. Southern Washington and north- 
ern: Oresoni 2. eee ee ee I. hallii 
Leaflets glabrous beneath; flowers solitary. 
Roots never tuber-like; lobes of leaflets with a shallow glandular notch at apices; 
tepals usually 12.0-16.0 mm long. Queen Charlotte Islands................ I. savilei 
Roots often tuber-like; lobes of leaflets glandular apiculate at apices; tepals 
usually 7.0-10.5 mm long. 
Tuber-like roots, fasciculate; follicles 10.0-12.0 mm long; styles recurved, 


1.0 mm long or less. Central and southern California............ I. occidentale 
Tuber-like roots never fasciculate; follicles 5.0-6.5 mm long; styles straight, 
ca. 1.5 mm: long. Hastern North Americas: I. biternatum 


Isopyrum savilei Calder and Taylor, sp. nov. Perenne, erectum, gla- 
brum, valde rhixomatiforme (10.5)—15.0-31.0-(36.0) cm; folia inferne 
glaucina, bi-ternatisecta; flores solitarii, terminales vel axillare; tepala 5, 
alba, decidua (9.8)—12.6-15.0-(16.8) mm long; (6.9)—8.2—10.2—(11.2) 
mm lat.; stamina 40—60, filamentis filiformis, clavatis, 5.0-8.0 mm long; 
carpellis sessilibus, 2—8; folliculi dense aggregati, arcuati, 11.0-15.0 mm 
long; semina 2-8, laevigata, ovoidea, apiculata cum raphid, 2.0—-2.3 mm 
long. 

A delicate upright perennial, glabrous throughout, strongly rhizoma- 
ous (10.5)—15.0-31.0-(36.0) cm high; leaves glaucous beneath, twice 
ternately compound, leaflets strongly 2—3-lobed, lobules entire to 3-lobed, 
with shallow glandular notches at apices, basal leaves usually one, cauline 
1—-several; flowers solitary, terminal or axillary; tepals 5, white, occa- 
sionally tinged pink at apex, readily deciduous, (9.8)—12.6—15.0—-(16.8) 
mm long, (6.9)—8.2-10.2-(11.2) mm wide; stamens usually 40-60, 
filaments filiform, clavate, 5.0-8.0 mm long; carpels sessile, 2—8; fruit 
a head of upright to strongly arcuate follicles, follicles 11.0-15.0 mm. 
long with recurved beaks; seeds 2-8, essentially smooth, ovoid, promi- 
nently apiculate with distinct raphe, 2.0—2.3 mm long. 

Type: 20 miles south of Moresby Logging Camp near an alpine lake, Moresby 
Island, Queen Charlotte Islands, British Columbia, Calder et al. 23055 (DAO). 

GRAHAM IsLAND: Empire Anchorage, Athlow Bay, Calder & Savile 21464; head 
of McClinton Bay, Masset Inlet, Calder et al. 21578; east side of Shields Bay, Rennell 
Sound, Calder & Taylor 23294; mountain north of Mt. Stapleton, Shields Bay, 
Rennell Sound. Calder & Taylor 23375. MorEesBy IsLtanp: Mt. de la Touche, Fairfax 
Inlet, Tasu Sound, Calder & Taylor 23566; mountain at west end of Mosquito Lake, 
Caider & Taylor 23721; 20 miles south of Moresby Logging Camp near an alpine 
lake, Foster & Joslin 56 (UBC); Tasu Inlet, June 26, 1961, Foster & Bigg (UBC). 

Isopyrum savilei is restricted to the Queen Charlotte Ranges at high 
elevations except on the west coast where subalpine conditions extend 


1963 | CALDER & TAYLOR: ISOPYRUM (p 


Fic. 1. Isopyrum savilei: A, habit, « ™%; B, stamen, x 5; C, fruit, x 2; D, seed, 
se 10: 


down to sea level. It is a species usually found in moist, shady, rock run- 
nels or in cliff crevices, but it occasionally extends onto talus slopes where 
suitable habitats exist. It is associated with many species, but is fre- 
quently found with Lloydia serotina, Saxifraga mertensiana, Anemone 
narcissifiora (s.l.), Romanzo fia sitchensis, Heuchera glabra, and Pingui- 


WD MADRONO [Vol. 17 


Fic. 2. Leaflets of North American species of Jsopyrum: A, I. savilei; B, I. biter- 
natum; C, I. occidentale; D, I. hallii; E, I. stipitatum. (All ca. X 2.) 


cula vulgaris. Although noted in all alpine areas surveyed, it was never 
a conspicuous element of the vegetation. 

A detailed comparison has been made between the five North American 
species, emphasizing those morphological characters which we feel are 
most diagnostic (table 1). We fully realize that additional characters 
such as the shape of the leaflets, stamen number, and follicle shape, which 


1963 | CALDER & TAYLOR: ISOPYRUM fle 


have been used by other authors, could have been included. However, the 
number of stamens and shape of the follicle is variable and cannot be 
used readily to separate the species. On the other hand, leaflet characters 
are distinct, but difficult to describe adequately. For this reason we have 
included detailed morphological comparisons of actual leaves utilizing a 
chloral hydrate/sodium hydroxide clearing technique (fig. 2). This 
method of comparison of leaf types clearly shows the venation patterns, 
lobule apiculation (or lack of same), and the degree and types of lobing. 
The distribution of the four western species is shown in Figure 3. 

Isopyrum savilei is strikingly distinct from the other specie of /so- 
pyrum that occur in North America in several morphological characters, 
e.g., the strongly rhizomatous nature of the root system, the shallowly 
notched tips of the ultimate leaf segments, the large showy flowers, the 
arcuate follicles, and the essentially smooth apiculate seeds with promi- 
nent raphes (fig. 1). 

The discovery of this endemic is significant as it provides further evi- 
dence of the close relationships between the North American and Japa- 
nese species. Close scrutiny of the distinctive western North American 
species, Jsopyrum hall, reveals that it possesses many similar charac- 
ters to the Japanese species, /. raddeanum, such as: pubescence of leaves, 
apiculate tipped leaflets, and seed coat characters. These observations 
confirm those of Drummond and Hutchinson (1920. p. 154) who stated, 
“The remarkably close affinity of two species of this genus, /. [nemuion | 
Raddeanum from Manchuria, and FE. Hallw from Oregon, is worthy of 
note.” Another group of Japanese species have glandular, notched tips 
on the ultimate leaf segments, smooth seeds, rhizomatous root systems, 
and usually two carpels; characters which are also found in Jsopyrum 
savilet. On the other hand, /. savilei also shows close relationships with 
the American taxa with respect to lack of staminodia, clavate filaments, 
follicle size, and deeper lobation of leaf segments. 

The North American species have been segregtated under Enemion 
(Drummond and Hutchinson, 1920) and this segregation was based on 
the tenuous character of carpel number and the presence or absence 
of petals; two morphological units which are extremely difficult to eval- 
uate in the family Ranunculaceae. They recognized seven genera; how- 
ever, only /sopyrum and Enemion are pertinent to the present discus- 
sion. Hnemion was proposed by Rafinesque (1821) to include a group of 
species differing from /sopyrum by the absence of petals and this sepa- 
ration was supported by Drummond and Hutchinson. It should be em- 
phasized that the use of the terms petals and sepals with respect to the 
genera in question was not supported by anatomical studies by either 
Rafinesque, or Drummond and Hutchinson. Indeed, the latter authors 
have based their segregation primarily on phyletic grounds rather than 
on critical evaluation of morphological characters. We believe the outer 
showy organs arranged in a spiral fashion are not sepals, but are best 
classified as tepals in accordance with modern terminology. In addition 


74 


MADRONO 


[Vol. 17 


e. bY Ks) 
WZ 


‘> 


Lsovilee 


L halts 


L.stip/tatum 
I. occidentale FZ 


Fic. 3. The geographical distributions of western North American Jsopyrum. 


WwW 


CALDER & TAYLOR: ISOPYRUM 


1963 | 


9}e[noIde 

ATyvomM pue ‘punol 
‘gsosni A][3UO0I}s 
‘SUO] WW /*[-p'T 
a}ye}yIds ! passoid 
-de pue yystidn 

: SUO[ UW Q'8-0'S 
Jepnsuel} A[MOIeU 
-Snouriquisw ‘yey 
SuO|T WU 8°7-6'T 
[el1a}eW polip 

UI JUIUTWIOId SUTOA 
:OpIM WU 7° 7-S'T 
‘SUO] WW (Q'9)- 
s's-2'p—-(S'e) ‘ory 
IIM0IqO ‘paley 
A[suo01}s xode 


AIBPI[OS SIIMOLY 


snoiqeys !ayepnoide 
Ie[npurls saqo] 
$JOOI (UWIIOJISN}] ) 
a¥T-19qn} ‘pauaysry} 
‘JAOYS JO VpOIOse] 
:SNo}eWLOZIYI OU 


Ws (S°2T)-O'IT-S's 
WN4D41G 14S * J 


dye[noide ATyeom 
pue ‘proao ‘aso[n3sni 
:SU0[ WU 7°7-0'Z 
a[ISSas !yUISIBAIP 
AJPPIM 0} yYUsIIdN 
:SUO] WW O'/-O'p 
I} RAPID 

APYPIM ‘UAOF YY 


SUO] WW 7Z°8-]'S 


[el1a}e8Ul potip 

UI snondIdsuOdUT SUIOA 
‘ OPIM WUT Co—E"¢ 

: SUO] WW ¢°6—-S*'Y ‘OTM 


poiey 
A[BUOI}S JaAoU xode 


ISOWAD SIIMOT] 


y}yeaueq JUs[N 
-daqnd ‘oazernoide 
Ie[Npur[s soqo] 


4904S}001 APOOM 
‘ynoys ‘y10Ys 
-SnOJeUIOZIYI JOU 


wd (0°L9)-O'S9-0'S¢S 
myDY *] 


yuadsaqnd 

AJaynutu ‘}UsuTUTOId 
oydei ‘proao ‘asopn3ni 
APysys 10 y}oouS 
‘SUO, WW 9Q°7-1'Z 
J[ISses !JUIBIIAIP 
ATapIM 0} YYysIUIdN 

- sUO,T WU S°9-0'S 


IJET[PAP]O ‘WO FT, 


suo, Wu 7°¢~-1°7 


[PI19}eW Pelip UI sno 
-noidsuosutl suraA Sapir 
Wu 9'9-7'p ‘suo, WU 
(ZIT)-T6-Z'2 “oy 
poiey 

A[SuO1}s JaAoU xode 


AIeP]OS SIOMOY 


SnNOIqe[S 5 9}e| 

-noide re[npurys saqo] 
SIITIISEJ 

UI JaAdU Jnq ‘sno 


-19qN} SaWIT}IWIOS s}OOI1 
:Sno}ewoziyt ApyVoM 


Ws O'Te-O'TT 
WUNIDULIILG "J 


aye[noido AyT}UoU 
-Tm01d pue ‘ploAo 
‘yyoows A][eUassa 
‘BuO, WU ¢'7-0'Z 


a[ISsos a} eNII1e 
A[Su01}s 0} JYysIAdN 
‘ 3UO] WU O'S T—-O'TT 


9} PAPO 
APYeaM ‘ULIO FY 


BUOT WW Q'8-0'S 


[el1oyeW palip ul 
Snonotdsuoour suloA 


-OpIm WW (7° TT)— 201-78 


—(6°9) ‘Buoy WU (g°9T)— 
O'ST-9°ZI-(8'6) ‘9}TYM 


poley 
A[BUO01}S JaAouU xode 


AIE}OS SIIMOY 


snoiqeys 
- Yoyou Ae[NyF ueps 
MOT[VYS B YIM soqo] 


WIOflUdl ‘asopnsni 

: SUO[ WU Q'7—-/'T 
J[ISSIS | }JUIBIIAIP 
AJapIM 0} 7YBIUdN 
:SUO] WU Q'ZI-O'0I 
I}BALTO 

AP{RIM ‘UWIOFTTY 


suo] WU ¢'S—-7'7 


[el10}eW ~pelip 

UI SNONSTdSUOSUT SUTIOA 
-Opim wut /°o-8'¢-(Z'¢) 
“SUOT UWL (E"1T)—£°6-0'L 
yuid Ay[euotsess0 ‘aqyTyM 


poley 
A[BUOI}S IaAoU xode 


AIVPIOS SIOMOLT 


SNOIqR]S $9}°] 
-noide aepnpurys saqo] 


SJOOI (UIIOJISN]) dy] 


pousayoty} JOU s}OOI -Jaqny ‘pauayory} ‘y10Ys Jo 
-SnO}eULOZIYA A[SUOI}S a[IIOSeJ {Ssno}eWOZIYyI jou 


Wd (O'9¢)— 
(OP ES Ok E(USu oy) 
1a]LAUDS *] 


wd (0'Or)- 
0'0£-0'ST-(0'L) 
91D]UIP1IIO “J 


WOUAdOST JO SAIOUdS NVOIWAWY HLYON AHL JO NOSIYVdWOD YW ‘1 ATav 


Spaas 


SofH[O} 
dIN}eIA] 


S}UIUILTI 


SUITES 


sjeday, 


apunpedg 


JUDISIIOYUT 


Sjoyea’] 


UI9}SAS JOO 


1Y4SIOH 


76 MADRONO [Vol. 17 


to the tepals, Drummond and Hutchinson have noted small petaloid 
structures in the European /. thalictroides, and stalked and bilobate 
structures in the Japanese species, /. stolonifera and I. trachyspermum, 
respectively. We feel they have misinterpreted these petaloid structures 
as petals and have subsequently placed too much stress on their value 
as characters for segregation of the genera. After careful examination of 
these structures, we have concluded they are staminodia and represent 
a transitional series from a sessile petaloid organ to one which is stalked 
bilobate. These structures are readily identifiable as staminodia and close- 
ly approximate the stamens with clavellate filaments found in several 
Isopyrum species. It should be pointed out that we have not completed 
a detailed ontogenetic study of these structures. 

Drummond and Hutchinson considered that both /sopyrum and Ene- 
mion were derived from Paraquilegia, a “primitive ancestoral”’ genus 
comprised of four species from the mountainous regions of central and 
southern Asia, and that they represent separate and distinct lines of 
divergence. We have no evidence that Paraquilegia does not represent the 
ancestoral progenitor of this complex and we do not disagree with their 
concept that a ‘“‘natural” group exists in Japan and North America. How- 
ever, we do think that the two groups, i.e., the Japanese and the North 
American, do not represent divergent lines of evolution, but rather that 
they represent overlapping stages within a single line of development. 
These two groups are closely related and there is no significant morpho- 
logical evidence for separating them into two separate and distinct genera, 
hence we consider that the North American taxa belong in the genus 
Isopyrum. 

Plant Research Institute 


Canada Department of Agriculture 
Ottawa, Ontario 


LITERATURE CITED 


Caper, J. A. and D.B.O. Savite. 1959. Studies in Saxifragaceae—II. Saxifraga sect. 

Trachyphyllum in North America. Brittonia 11:228-249. 
. 1960. Studies in Saxifragaceae. III. Saxifraga odontoloma and lyallii, and 

North American subspecies of S. punctata. Canad. Jour. Bot. 38:409-435. 

DrumMmMonpD, J. R. and J. Hutcurnson. 1920. XXIII. A revision of Isopyrum (Ra- 
nunculaceae) and its nearer allies. Kew Bull. No. 5:145-169. 

GREENE, E. L. 1897. New or noteworthy species XX. Pittonia 3:249. 

Maruzas, M. and L. Constance. 1959. New North American Umbelliferae—III. Bull. 
Torrey Club 86:374-382. 

RAFINESQUE, C.S. 1820. Enemion biternatum. Jour. Phys. Chim. Hist. Nat. 91:70. 


1963 | TURNER: HYMENOXYS del 


CYTOPHYLETIC ANALYSIS OF HYMENOXYS ODORATA: 
A RECAPITULATION! 


B. L. TURNER 


In a recent article in Madrono, Speese and Baldwin (1963)? stated, 
“The basic number for H. odorata is 11. (We assume the report [sic] 
of n = 15 for this species to be incorrect.)’’ In spite of this statement, 
the authors succinctly summarized the reported chromosome counts for 
H. odorata by referring to 3 populational counts from the United States 
(reported independently by 2 different groups of workers) as being di- 
ploid with n = 11; they also referred to chromosome counts by myself 
from 2 Mexican populations which had n = 15. To convince the reader 
that the published count of n = 15 for H. odorata might be in error, they 
pointed out that this count (Turner, Beaman & Rock, 1961) was made 
from “‘pollen-mother-cell smears of buds fixed in the field during the sum- 
mer of 1959. Our experience has been that preparations from material 
so fixed are often difficult to interpret, and especially so if the weather 
were hot at the time of fixation.” This, in spite of the fact that a camera 
lucida drawing included in the published account showed a meiotic figure 
with n = 15. 

Upon reading Speese and Baldwin’s comments, I felt compelled to take 
a second look. The following account, though phrased in an admittedly 
personal way, is my version of the story: 

When I first received the pickled buds of H. odorata from Mexico in 
1960, I was reluctant to examine these since I recognized the species as 


TABLE 1. CHROMOSOME COUNTS OF HYMENOXYS ODORATA FROM MEXICO 


COLLECTION CHROMOSOME COUNT SOURCE 

Coahuila: 12 mi S of Saltillo al (buds) 

Powell & Edmondson 528. TEX 211 = 30 (root tips) 
Nuevo Leon: 39 mi S of Saltillo 

Powell & Edmondson 543. TEX one = 30 (root tips) 
Nuevo Leon: 24 mi S of Galeana 

Crutchfield & Johnston 5860. TEX if ema Et (buds) 
Nuevo Leon: 41.2 mi S of Saltillo 

Rock M264. TEX n= 15 (buds) 
Nuevo Leon: 1 mi S of San Roberto. 

Thompson & Doolin 2163. TEX N15 (buds) 


1 The rhetorical definition of recapitulation is preferred here being “A form of 
peroration in which the respective processes, as of explanation, conviction, excita- 
tion, and persuasion, pursued in a discourse, are concisely repeated for the purpose 
of more complete effect.” (Funk and Wagnalls, New Standard Dictionary, 1945 
edition.) 

2 The authors were apparently unaware of an earlier report of n= 15 for H. 
anthemoides from Argentina (Solbrig, 1962). 


78 MADRONO [Vol. 17 


being one already counted by Speese and Baldwin (1952). However, I 
thought the single previous count needed checking, so I examined the 
material and to my surprise it showed n — 15. I double checked this by 
counting the meiotic material from several different florets and heads; 
all counts were n — 15. As indicated by Speese and Baldwin in their ref- 
erence to a Johnston collection (5860), also from Mexico, I again counted 
n = 15 for H. odorata; this time I was not surprised, but I noted in my 
lab book and indicated on the collector’s label, ““‘n = 15, clearly, det. 
B. L. Turner from PMC’s.”’ 

My next encounter with the chromosomes of H. odorata came in the 
fall of 1961 when a graduate research assistant, A. M. Powell, counted 
n = 15 for a collection of his from Nuevo Leon, Mexico, from the same 
general area of the previous Mexican collections with n = 15. By this 
time, / was beginning to question the counts of n = 11 reported for H. 
odorata by Speese and Baldwin (1952) as well as those of Raven and 
Kyhos (1961). My curiosity now being whetted, I collected in the spring 
of 1962, buds from 3 populations of H. odorata (2 in Texas and 1 in 
California, the latter from the same area from which Raven and Kyhos’ 
counts were obtained). I was delighted to find n = 11 in all these collec- 
tions (Powell and Turner, 1963); my confidence in my scientific col- 
leagues now re-established I forgot the issue until the recent publication 
of Speese and Baldwin (1962), statements from which are quoted above. 

My first reaction to Baldwin and Speese’s comments was mild irrita- 
tion; this soon gave way to the haunting fear that my observational 
senses were being affected by the too frequent exposure to acetocarmine 
fumes, to say nothing of the PDB to which we are all accustomed; 
finally I couldn’t bear the onus of my conscience and decided to germi- 
nate seeds from the original Mexican collection or collections to deter- 
mine if indeed the mitotic counts might not tell a different story, one told 
from the cool confines of a petri dish instead of the hot atmospheric con- 
finement accorded the original material. In spite of the trivial nature of 
the “experiment,” my excitement was high. I personally transferred the 
root tips through their various solutions, and by observation time, I was 
in a fit of fear and hopefulness difficult to imagine by anyone not caught 
in similar observational disputes. 

The mitotic counts proved to be 2n = 30 (Table 1), much as you must 
have guessed by this time or else I would not have gone to the trouble 
to write this paper. Hymenoxys odorata is obviously dibasic with x = 11 
and x = 15 (so far as known). Whether this constitutes cytophylesis in 
the sense of Baldwin (1939)? I leave to the judgment of the reader. 


The Plant Research Institute and Department of Botany 
University of Texas, Austin 


3 Baldwin does not define the term cytophyletic in this paper. One can only infer 
its meaning from a single pertinent sentence as follows: “That the Crassulaceae are 
so chromosomally variable and yet closely related makes a cytophyletic approach to 


1963 | VASEK & FERGUSON: LOLIUM 79 


LITERATURE CITED 


BALDWIN, J. T., JR. 1939. Certain cytophyletic relations of Crassulaceae. Chron. Bot. 
5:415-417. 

PowELL, A.M. and B.L.TurRNeER. 1963. Chromosome numbers in the Compositae. 
VII. Additional species from the southwestern United States and Mexico. 
Madroho (in press). 

RAVEN, P. H. and D.W. Kyuos. 1961. Chromosome numbers in Compositae. II. 
Helenieae. Am. Jour. Bot. 48:842-850. 

Sotsric, O. T. 1962. Numero cromosomico de una Compuesta entrerriana (Hyme- 
noxys anthemoides). Darwiniana 12:521. 

SPEESE, BERNICE M. and J.T. BALpwin, JR. 1963. Cytophyletic analysis of Hyme- 
noxys anthemoides. Madrono 17:27-29. 

—. 1952. Chromesomes of Hymenoxys. Am. Jour. Bot. 39:685—-688. 

Turner, B.L., J.H. BEAmaAn, and H.F.L. Rock. 1961. Chromosome numbers in 
the Compositae. V. Mexican and Guatemalan species. Rhodora 63:121-129. 


A NOTE ON TAXONOMIC CHARACTERS IN LOLIUM 
FRANK C. VASEK AND J. KIRK FERGUSON 


The introduced grasses, Lolium perenne L. and L. multiflorum Lam., 
are listed in many manuals (e.g. Abrams, 1940; Munz, 1959; Hitchcock, 
1950) as two distinct species, separated from each other primarily on 
the basis of whether the lemma is awned or not. However, in southern 
California, awned and awnless plants frequently grow in mixed stands. 
A study was begun to determine whether the presence or absence of an 
awn is sufficient grounds for distinguishing the two species and to inves- 
tigate the possibility that other criteria might be more valid. A sample 
of 50 plants was collected from a mixed population, growing alongside 
U. S. Highway 60 near the campus of the University of California at 
Riverside and studied to determine whether characters that distinguish 
the two species were correlated. In addition, seeds were collected from 
an awnless plant growing in the Riverside locality and from an awned 
plant and an awnles plant growing at Zumwalt Meadows, Kings Canyon 
National Park, California. The latter locality was selected because it is 
climatically and ecologically greatly different from the former locality. 
The seeds were planted in the greenhouse and the resulting progenies were 
studied for character correlation. 

According to descriptions found in manuals (Fernald, 1950; Hitch- 
cock, 1950; Munz, 1959) L. multiflorum is characterized by awned (at 
least the upper) lemmas 7 to 8 mm. long, 10 to 20 florets per spikelet, and 


an understanding of the family sound: differences in chromosome number and be- 
havior and morphology allow a recognition of trends, and the trends are funda- 
mental, their divergences and convergences constituting a basic revolutionary pat- 
tern; taxonomic categories with names and ranks are conveniences and may or may 
not have real significance.” 


80 MADRONO [Vol. 17 


spikelets twice as long as the glumes, while L. perenne is characterized by 
awnless, or nearly awnless, lemmas 5 to 6 or 7 mm. long, 6 to 10 florets 
per spikelet, and spikelets little exceeding the glumes. Based on these de- 
scriptions, the 50 plants in the Riverside sample would be identified as 
follows: 33 L. multiflorum and 17 L. perenne on the basis of awns vs. 
nearly awnless, if nearly awnless is interpreted to include awns up to 
2mm.; 6 L. multiflorum, 6 either species, and 38 L. perenne on the basis 
of floret number; and no L. multiflorum, 3 either species, and 47 L. 
perenne on the basis of lemma length. 

Actually, the sample from Riverside (fig. 1) indicates continuous vari- 
ation in both floret number and spikelet glume ratio. These traits are not 
correlated with awn length and three-fourths of the plants are not con- 
sistently one species or the other. The Riverside sample thus appears to 
represent one segregating population. 


Meo 
2 | 
Of ‘ 
g 
8 ! 
of 
6 ‘of d 
3 
4 


IS IS IT 1G 2b 235° 25°27 5° 29° 31 


33° 35 


Fic. 1. Scatter diagram of a wild sample from Riverside showing the relationship 
between the number of florets per spikelet (vertical axis) and the ratio of spikelet 
length/glume length (horizontal axis). Long diagonal lines indicate awns 1.5-—7 mm. 
long, short diagonal lines indicate awns 0.5—1 mm. long. 


A progeny of 15 plants, grown in the greenhouse and derived from one 
one pollinated awnless plant growing in the Riverside population, shows 
a similar lack of correlation between awn length and the floret number 
and spikelet/glume ratio (fig. 2). Furthermore, these plants, all from the 
same female parent, segregated for the awn-awnless character. Seven 
plants had an awn 2—6 mm. long, two plants had an awn 1 mm. long, and 
six plants had no awn. 

The two progenies from the Kings Canyon (fig. 3) also show a lack of 
correlation between awn length, floret number and the spikelet/glume 
ratio. The distribution of plants on the scatter diagram indicate, too, that 
the two progenies represent but one population. Furthermore, both prog- 
enies segregated for the awn-awnless character as follows: 


1963 | VASEK & FERGUSON: LOLIUM 81 


PARENT OFFSPRING 

Awnless Awn 0.5-1.5 mm. Awn 2.0-6.0 mm. 
Awnless 2 11 8 
Awned 4 fi 20 


Unpublished data of the authors further indicate that awn length, 
floret number and spikelet/glume ratio are not correlated with spike 
length, spikelet length, number of spikelets per spike, number of spikes 
per plant and flowering response to cold treatment, long photoperiod and 
normal day length. 


| 
l2 per | mult 


Fic. 2. Scatter diagram of the progeny of an open pollinated, awnless seed parent 
from Riverside (coordinates as in fig. 1). 


re per, mult 


Fic. 3. Scatter diagram of the progeny of an awnless seed parent (open circles) 
and an awned seed parent (solid circles) from Kings Canyon National Park (coordi- 
nates as in fig. 1). 


82 MADRONO [Vol. 17 


Although our samples are few, our data clearly indicate that segrega- 
tion for awned vs. awnless may occur in the progeny of a single plant, 
regardless of whether that plant was awned or awnless. Furthermore, the 
progenies of individual plants exhibit continuous variation in floret num- 
ber and spikelet/glume ratio. Thus, these characters do not distinguish 
two species. Nor could we find any other basis for distinguishing two 
species. 

Separation of two species on the basis of the traditional taxonomic 
characters thus appears to be arbitrary and our studies suggest only one 
species is involved. A similar situation may have confronted Jenkin 
(1931) who concluded, after considerable experience breeding various 
grasses, that there was no evidence of incompatibility or of sterility in 
F, of L. perenne and (at that time) L. perenne var. multiflorum. Jenkin’s 
results were not different from those obtainable in intraspecific crosses. 
This suggests that L. perenne and L. multiflorum are, in reality, the same 
species. 

However, we have not tested the ‘key’ characters used by Fernald 
(1950), namely whether the unexpanded leaves are folded or inrolled, 
whether the glumes exceed or do not exceed the contiguous floret, and 
whether the rachis is smooth or roughened. Presumably Jenkin did not 
use these characters either. Therefore, a possibility exists that our ma- 
terial, and perhaps that studied by Jenkin, may have been all L. perenne 
or all L. multiflorum. If so, the problem of identification remains acute 
in view of the inadequacy of descriptions currently found in west Ameri- 
can manuals. Should the ‘key’ characters used by Fernald prove to be 
capable of distinguishing two valid taxa, the problem of inadequate de- 
scription is no less acute for these ‘key’ characters appear to be merely 
added to the conventional, inadequate descriptions. 


In conclusion, the treatment of Lolium perenne and L. multiflorum, as 
currently found in many manuals, is unsatisfactory. A thorough test of 
the characters used by Fernald, followed by revision of the species de- 
scriptions, is in order and thorough population analyses may reveal the 
existence of only one species. 


University of California 
Riverside, California 


LITERATURE CITED 


ABRAMS, L. 1940. Illustrated flora of the Pacific States, Vol. I. Stanford Univ. Press. 
FERNALD, M.L. 1950. Gray’s Manual of Botany. American Book Co., New York. 


Hircucock, A.S. 1950. Manual of the grasses of the United States. U.S. Dept. Agr. 
Misc. Publ. 200. Washington. 


JENKIN, T. J. 1931. The interfertility of Lolium perenne and Lolium perenne var. 
multiflorum. Bull. Welsh Pl. Breeding Sta. Ser. H., No. 12:121-125. 


Muwnz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley. 


1963 | FRYXELL: MALVACEAE 83 


CLEISTOGAMY IN THE MALVACEAE 
PAuL A. FRYXELL 
INTRODUCTION 


Facultative cleistogamy is a phenomenon frequently overlooked. This 
was made clear by Biloni (1957) who grew Pavonia sepium St. Hil. under 
circumstances permitting daily observation of the plants for several 
years. He published a note on the flowers they produce (Biloni, 1945) 
without suspecting that, in addition to these showy chasmogamous flow- 
ers, they also produce abundant cleistogamous flowers. The observation 
of fruits maturing in the spring before flowering was apparent suggested 
the occurrence of cleistogamy, which he subsequently found does occur 
(Biloni, 1957). 

If such intensive observation of living plants by a competent botanist 
fails immediately to disclose the occurrence of cleistogamy, botanists 
working primarily with dried plant materials may be expected to overlook 
it also. Consequently, only a few examples are to be found in the litera- 
ture of cleistogamy in the Malvaceae. They are as follows: 


SPECIES TRIBE AUTHORITY 
Pavonia hastata Cav. Ureneae Heckel, 1879 
P. sepium St. Hil. Ureneae Biloni, 1957 
Malva parviflora L. Malveae Uphof, 1938 
Hibiscus trionum L. Hibisceae Lassimonne, 1929 


These few examples are distributed among three tribes, suggesting that 
facultative cleistogamy is of widespread occurrence in the family. 


OBSERVATIONS 

A number of species of the Malvaceae, all in the tribe Hibisceae, have 
been under cultivation and regular observation for several years. Five 
species have shown some tendency to produce cleistogamous flowers 
under the environmental conditions in which they have been grown. Com- 
parative observations have shown that several other closely related spe- 
cies do not produce cleistogamous flowers. The five species are discussed 
individually as follows. 

Hibiscus denudatus Benth. was grown from seed collected in the Cape 
Region of Baja California by I. L. Wiggins. Underd greenhouse con- 
ditions that appeared adverse to the optimum growth of this species, it 
produced both cleistogamous and chasmogamous flowers, the former in 
greater abundance. 


[Note added after manuscript had gone to press. ] 

A recent paper by Brown (Brown, Meta S. Anomalous flowering Gossypium 
australe F. Muell. Jour. Hered. 53:139-141. 1962.) refers to the same phenomenon 
noted here and illustrates a cleistogamous flower at anthesis, dissected to show the 
reduction of parts. 


84 MADRONO [Vol. 17 


Cienfuegosia argentina Gurke var. hasslerana (Hochr.) Hassler f. B 
escholtzioides Hassler was grown from seed collected in Argentina by 
Manuel Gutierrez. Under greenhouse conditions that appear to be fa- 
vorable to its growth, this species produces chasmogamous flowers only 
rarely, while literally hundreds of fruits result from cleistogamous flow- 
ers (fig. 1, c and d). Chasmogamous flowering occurs in the open. Dis- 
section of cleistogamous buds shows a marked reduction of flower parts 
with, for example, only five or six anthers. 


Fic. 1. Flowers and fruits of Cienfuegosia: a-b, C. drummondi (A. Gray) Lew- 
ton: a, young fruit developing from cleistogamic flower with persisting “cap” (two 
bracts and one calyx lobe removed), x 1; 6, chasmogamous flower, X 34. c-d, C. 
argentina Girke var. hasslerana (Hochr.) Hassler: c, young fruit developing from 
cleistogamic flower with persisting “cap” (two calyx lobes removed), x 3; d, nearly 
mature fruit developing from cleistogamic flower with persisting “cap,” * 3%. 


Cienfuegosia drummondu (A. Gray) Lewton was grown from several 
seed accessions from southern Texas obtained through C. F. Lewis 
and M. Lukefahr. In the greenhouse and in the open it produces both 
chasmogamous (fig. 1, 0:) and cleistogamous flowers (fig. 1,@) in approxi- 
mately equal proportions. The fruits developing from the two types of 
flowers cannot be distinguished in size or number of seeds. 


Gossypium australe F. von Muell. was grown from seed obtained from 
J. H. Saunders, Khartoum, though the native habitat of this species 
is north-central to north-west Australia. Under conditions apparently 


1963 | FRYXELL: MALVACEAE 85 


favorable to its growth and reproductive activity, it has set numerous 
fruit, but its first chasmogamous flower was produced only after some 
twenty fruits had set from cleistogamous flowers. Later flowering is of 
both types. Chasmogamous flowers tend to develop at the first node of 
a fruiting branch with cleistogamous flowers developing at subsequent 
nodes. 


The fruits developing from the two types of flowers of G. australe can 
be distinguished by the size of the calyx. The calyx lobes of the cleisto- 
gamic fruit exceed the bracteoles by only 2—3 mm. (fig. 2,@), while in the 
chasmogamic flower and fruit the calyx lobes exceed the bracteoles by 
9-10 mm. (fig. 2, 0). In either case the bracteoles are of essentially 
equal size. 


Gossypium bicku Prokhanov | Notoxylinon pedatum (Bailey) Lewton | 
was grown from seed collected in central Australia by George Chip- 
pendale. This species is closely allied to G. australe (Fryxell, in manu- 
script) and shares with it the propensity for producing cleistogamic 
flowers. Indeed, a cleistogamic “cap” has been observed on the syntype 
of G. bicku (E. W. Bick 82, Sept.-Oct., 1910, BRI). 


Fic. 2. Flowers and fruits of Gossypium australe F. Muell.: a, fruits developing 
from cleistogamic flowers at various stages of development with “cap” persisting, 
x 1%; b, chasmogamic flower bud on day of anthesis, « 1; c-d, the same flower 
as in b, a few hours later, x 34. 


86 MADRONO [Vol. 17 


Should the ‘“‘cap” (the dried corolla of the cleistogamic flower) persist 
through the development of the fruit, its presence is, of course, diagnostic 
for cleistogamy for that fruit for all the species discussed. In all cases, 
also, the first flowers produced are cleistogamic; chasmogamous flowers 
are produced after the plant has attained additional growth. 


DISCUSSION 

Facultatively cleistogamous plants are known to depend upon envi- 
ronmental conditions for the degree to which their cleistogamy is ex- 
pressed (Brown, 1952; Ernst-Schwarzenbach, 1956; Harlan, 1945; Up- 
hof, 1938). Therefore, one cannot state with any assurance that a given 
species produces cleistogamous flowers in any particular proportion except 
with reference to some specific set of environmental conditions. Of the 
five species under discussion here, none was growing in its native habitat. 

I do not wish to imply that differences in frequency of cleistogamy are 
not real, because they doubtless are. But without controlled experiments 
in which environmental variables are carefully manipulated (as was done, 
for example, by Brown), discussions of the frequency of cleistogamy are 
not productive. Nevertheless, comparisons with related taxa in which 
cleistogamy is absent can be of value. 

In addition to the two species of Cienfuegosia described herein as cleis- 
togamic, two others have been observed sufficiently to state that they 
do not produce cleistogamic flowers under the various conditions in which 
they have been grown. They are C. heterophylla (Vent.) Garcke and 
C. hildebrandtu Garcke. 

Similarly, in Gossypium, extensive observation of many species of this 
genus has heretofore failed to bring to light examples of cleistogamy.” 
This genus, of course, has been intensively studied by botanists for many 
years. However, G. australe and G. bicki, the species described here as 
facultatively cleistogamic, have become available in culture only recently. 
Gossypium australe originally was placed in Gossypium by von Mueller 
in 1858. Subsequently it was assigned by various authors to Fugosia, 
Cienfuegosia, Hibiscus, and Notoxylinon (Lewton, 1915). More recently 
it was returned to Gossypium on morphological grounds by Prokhanov 
(1947) and for cytological reasons by Saunders (1961). Gossypium bicku 
was first described as Fugosia pedata by Bailey in 1910. It was included 
in Notoxylinon by Lewton (1915) and more recently transferred to Gos- 
sypium by Prokhanov (1947). The inclusion of these two species in 
Gossypium is considered sound by the present author, although the occur- 
rence in them of cleistogamy is unusual for Gossypium and must be given 
attention in classifying and understanding the genus (Fryxell, in manu- 
script). In both G. australe and G. bicki from their native habitat the 


2A mutant strain of G. barbadense L. is extant that is described as ‘“cleisto- 
gamic” but it does not have cryptic floral development with a marked reduction of 
floral parts. The mutant strain merely has a normal chasmogamous flower that fails 
to open fully. 


1963 | FRYXELL: MALVACEAE 87 


author has observed cleistogamic “caps” on herbarium specimens, indi- 
cating that cleistogamy is not confined to artificial growing conditions. 

I can say little concerning the taxonomic value of cleistogamy as an aid 
to understanding the large and complex genus Hibiscus, because of my 
limited knowledge of that genus. However, the examples described of 
differential incidence of cleistogamy below the level of genus in the genera 
Cienfuegosia and Gossypium point to this phenomenon as an additional 
means of understanding the subgroups of these genera. 


SUMMARY 


The incidence of cleistogamy in the Malvaceae is reviewed, and the 
occurrence of cleistogamy is noted in the following species of the Hibis- 
ceae: Hibiscus denudatus Benth., Cienfuegosia argentina Giurke var. 
hasslerana (Hochr.) Hassler, C. drummondiu (A. Gray) Lewton, Gossy- 
pium australe F. von Muell., and G. bickit Prokhanov. 


University of Arizona Cotton Research Center 
U.S. Department of Agriculture 
Tempe, Arizona 


LITERATURE CITED 


Bitoni, J. S. 1945. In: Suelo Argentino 40:55-56. 


. 1957. Observaciones sobre las flores cleistogamas de la Malvacea Pavonia 
sepium Saint Hilaire. Darwiniana 11:286-289. 


Brown, W. V. 1952. The relation of soil moisture to cleistogamy in Stipa leucotricha. 
Bot. Gaz. 113:438-444. 


ERNST-SCHWARZENBACH, M. 1956. Kleistogamie und Antherenbau in der Hydro- 
charitaceen Gattung Ottelia. Phytomorph. 6:296-311. 


FrYXELL, P. A. The genus Gossypium in Australia. Manuscript. 


Haran, J. R. 1945. Cleistogamy and chasmogamy in Bromus carinatus Hook. & Arn. 
Am. Jour. Bot. 32:66-72. 


HECKEL, E. 1879. De l’état cleistogamique de la Pavonia hastata. Comptes Rendus 
des Seances de l’Acad. de Paris 89:609. 


LASSIMONNE, S. 1929. Cleistogamie d’Hibiscus trionum L. Bull. Soc. Bot. Franc. 75: 
1014-1015. 


LewrTon, F.L. 1915. The Australian Fugosias. Jour. Wash. Acad. Sci. 5:303-309. 


PROKHANOV, Y. 1947. [The conspectus of a new system of cotton (Gossypium L.) ] 
Bot. Zhur. SSSR 32:61-78. (In Russian and Latin.) 


SAUNDERS, J. H. 1961. A re-examination of a species of the genus Notoxylinon. Em- 
pire Cotton Growing Rev. 38:103-105. 


Upuor, J. C. Th. 1938. Cleistogamic flowers. Bot. Rev. 4:21-49. 


88 MADRONO [Vol. 17 


SWALLENIA, A NEW NAME FOR THE CALIFORNIA GENUS 
ECTOSPERMA (GRAMINEAE) 


THOMAS R. SODERSTROM AND HENRY F. DECKER 


In May of 1949, Annie M. Alexander and Louise Kellogg collected a 
most unusual grass forming dense tussocks on sand hills in Eureka Valley, 
Inyo County, California. Specimens were sent to Jason R. Swallen, Smith- 
sonian Institution, for identification. It was found to be distinctive in so” 
many characters that without doubt it represented an undescribed genus. 
Swallen named it Ectosperma, but he recently pointed out to the authors 
that the name is invalid because it had been used in 1803 for an algal 
genus. We take this opportunity to rename the grass Swallenia, in honor 
of this distinguished American agrostologist who has contributed so great- 
ly to our knowledge of the grasses of the New World. 


Swallenia Soderstrom & Decker, nom. nov. Ectosperma Swallen, Jour. 
Wash. Acad. 40:19. 1950, non Ectosperma Vaucher, 1803 |= Vaucheria 
DC. |; 

Type species: Swallenia alexandrae (Swallen) Soderstrom & Decker, 
comb. nov. Ectosperma alexandrae Swallen, loc. cit. 


In his paper describing the genus, Swallen placed it in the tribe Festu- 
ceae on the basis of gross morphology, indicating that the glumes, almost 
as long as the spikelet, might point to affinities with the tribe Aveneae. 
He added that “‘the characters are so distinctive that it is difficult to de- 
termine its actual relationship in the tribe.” Pilger (1954) also included 
it in the Festuceae, aligning it with the genera Melica, Schizachne, Lyco- 
chloa, Vaseyochloa, Anthochloa, Neostapfia, and Ramosia, in subtribe 
Melicinae. More recently, Stebbins and Crampton (1961) have placed 
it in the tribe Aelurop[o|deae, along with Aeluropus, Distichlis, Monan- 
thochloé, Jouvea, and Vaseyochloa, taking into consideration additional 
morphological and anatomical characters. A fuller discussion of the rela- 
tionship of Swallenia to these latter genera will appear in a forthcoming 
paper by the present authors. 


Department of Botany, 
Smithsonian Institution, Washington, D.C. 


Department of Botany & Bacteriology, 
Ohio Wesleyan University, Delaware, Ohio 


LITERATURE CITED 


PILGER, R. 1954. Das System der Gramineae unter Ausschluss der Bambusoideae. 
Bot. Jahrb. 76(3) :281-384. 


STEBBINS, G. L. & BEECHER CRAMPTON. 1961. A suggested revision of the grass genera 
of temperate North America. Jn Recent Advances in Botany (IX Int. Bot. Con- 
gress, Montreal, 1959) 1:133-145). 


1963 ] REVIEWS 89 


REVIEWS 


Ferns and Fern Allies of Chihuahua, Mex.ico. By Irvinc W. KwnopiocH and 
Donovan S. CorreELL; illustrated by PHOEBEJANE HoRNING and JANE ROLLER. xiv 
+ 198 pp., 57 plates. Contr. Tex. Res. Found. Bot., Vol. 3. Texas Research Founda- 
tion, Renner, Texas, 1962. $10.00. 

This attractively prepared volume merits special attention for it is the first com- 
prehensive modern treatment of the rich pteridophyte flora of Mexico. The authors 
have both collected extensively in the state of Chihuahua, with the particular pur- 
pose of augmenting the previous collections and phytogeographic knowledge. To- 
gether, they have collected all but 18 of the species. They are to be congratulated on 
the thoroughness of their work and the accurate presentation of their results. 

Although 120 species are known from Chihuahua, and an additional 6 from adja- 
cent areas are treated, the authors recognize that the fern flora is still incompletely 
known. Additional species will be discovered as collecting continues in many less 
accessible areas, especially the isolated mesic barrancas of the western Sierra Madre 
Occidental. However, the basic explorations have been completed. The authors are 
to be commended in bringing out their work at this time so that it may fill a long- 
standing need. 

The introduction surveys the collecting activities in the state of Chihuahua, the 
vegetation, geology, physiography, climate and the ecology and distribution of the 
pteridophytes. There is also a discussion of the typical structure and life-history of 
ferns, their cultivation and economic uses. A list of the known chromosome numbers 
of Chihuahuan species is a useful supplement. The principal text presents the perti- 
nent nomenclature, a careful and complete description and the ecology and distribu- 
tion of the 138 taxa recognized. Each species is illustrated; a number of them ade- 
quately for the first time. Keys are provided as an aid in identification and these, in 
combination with the excellent illustrations and the frequent discussions of characters 
make the book one of unusual utility. The appendix is devoted to comments on the 
species to be expected in Chihuahua, a gazetteer, a glossary of terms, a bibliography 
and an wdex. The gazetteer will be useful to many botanists concerned with the 
location of often obscure place-names in Chihuahua. 

From my own point of view, it is refreshing that the classification adopted is 
a highly realistic one. I refer particularly to the recognition of the Polypodiaceae in 
the broad sense, to the generic treatment of the Cheilanthoid ferns and to the use 
_of the category variety (rather than species) for important but subordinate taxa. 
Such practice is a departure from the tendency of many authors to accept, uncriti- 
cally, some of the more recent but unsubstantiated views on classification. 

The pteridophyte flora of Chihuahua, as one expects is primarily a xeric one. This 
is emphasized by the predominance of the genera characteristic of arid lands in the 
Americas. Cheilanthes is represented by 23 species, Notholaena by 15, Selaginella 
by 11 xeric ones and Pellaea by 9. The truly mesic element consists of about 12 
species and it is confined to the barrancas of the Sierra Madre Occidental. This is a 
tropical element which mostly reaches its northwestern limit in Chihuahua. Repre- 
sentative species of this element are: Dennstaedtia distenta, Trichomanes radicans, 
Hymenophyllum tunbridgense, Adiantum Poireti, Thelypteris rudis and Dryopteris 
parallelogramma. This book will be useful for a considerably larger area than the 
single state it covers. Fifty-nine of the species also occur in southern California, 
Arizona or New Mexico and 110 of them occur elsewhere in Mexico. Of these, 85 
extend southward to central Mexico and 61 to southern Mexico or beyond. Eleven 
species are known in Mexico only from Chihuahua. Five of these are endemic and 
the remainder occur in the United States. 

The ferns and fern allies of Chihuahua is an authoritative treatment; it can be 
recommended to all botanists with an interset in ferns or with a broader interest 
in the flora of arid regions. It will also be an important reference for those concerned 
with any portion of the flora of Mexico or the southwestern United States——RoLLa 
M. Tryon, Gray Herbarium, Harvard University. 


90 MADRONO [Vol. 17 


The Morphology of Pteridophytes. By K. R. Sporne. 192 pages, many figures. 
1962. Hutchinson & Co., Ltd., London. 12 s 6 d. 


It is difficult not to become enthusiastic about this little book. In less than 200 
pages Sporne reviews the main structural features of pteridophytes, discusses 
alternation of generations, and presents a concise and up-to-date account of the 
Psilophytopsida, Psilotopsida, Lycopsida, Sphenopsida and Pteropsida. The final 
chapter contains some general conclusions among which we find an admirably bal- 
anced discussion of the influential but fading Telome Theory, reference to Bower’s 
Enation Theory, and a brief account of the bearing of cytological and experimental 
data on the distinctions between the sporophyte and gametophyte generations in 
spore-bearing plants. This last chapter is especially welcome. It is rare, indeed, to 
find a book in structural botany which does not abruptly end in the descriptive 
details of a particular organ. 


The taxonomic framework of the book is squarely based on Reimer’s system in 
the 1954 edition of Engler’s “Syllabus der Pflanzenfamilien.” One of the most valu- 
able aspects of Sporne’s treatment is his acquaintance with fossil plants, and his 
careful integration of it with the existing knowledge of living plants. But this is not 
a fossil book; a casual glance at the bibliography shows much very recent work. 
The illustrations, mostly redrawn from other works, are consistently good although 
often too crowded on the page. 

The most amazing thing about “The Morphology of Pteridophytes’” is the incred- 
ibly low price, a mere two dollars. Surely this is the biggest bargain in botanical 
books today. Sporne’s book will appeal not only to those of us who lounge in the 
shaded halls of descriptive botany, but also to more adventurously inclined botanists 
with even the slightest appreciation of things ferny. 

A most valuable addition to morphological literature; a gem.—Jos KurjT, De- 
partment of Biology and Botany, University of British Columbia, Vancouver, Canada. 


Vascular Plants of the Pacific Northwest. By C. LEo HircHcock, ARTHUR CRON- 
QuIST, MARION OWNBEY, AND J. W. THompson. Illustrated. University of Washing- 
ton Press. Part 3, pp. 1-614. 1961. $13.50. 


Publication of this third part of the projected five-part Vascular Plants of the 
Pacific Northwest marks the passing of the halfway point in this notable series. 
In an earlier review (Madrofio 16:74-76) I made several general comments and criti- 
cisms largely still relevant to the present part, which covers Saxifragaceae through 
the Garryaceae. The Englerian sequence is followed through the Umbelliferae, 
though the placement of the Garryaceae following this family is a departure in keep- 
ing with recent evidence concerning the affinities of this former amentiferan. 


In this portion of the series, most groups were treated by C. L. Hitchcock; Rosa 
and the Umbelliferae were prepared by Arthur Cronquist. The conservative taxo- 
nomic approach evident earlier is continued here. Such conservatism, while attrac- 
tive in offering a greater assurance of arriving at a name for a taxon, runs the risk 
of obscuring important facts regarding variation patterns in the broadly-conceived 
species. To a large extent, however, this problem is alleviated by pointing out such 
patterns in the discussion of the species. Perplexing genera such as Lupinus and 
Astragalus are handled well. There is a key to fruiting plants and another to flower- 
ing material of Astragalus, the treatment of which was prepared with the assistance 
of Rupert Barneby, whose A. amni-amissi appears to be the only new species de- 
scribed in the families covered by this part of the series. The treatment of the lupines 
is one of the most workable to appear in years. It is refreshing to learn that at last 
Rosa, mistreated by taxonomists for so long, has had her real species delimited! 
A number of species and a few genera have disappeared as casualties of lumping; 
Pseudocymopterus and Pteryxia have been swallowed by Cymopterus. 

The larger type in this part, the use of italics, and glossier paper considerably 
improve the appearance of the printed page. The inclusion in the keys, though not 


1963 | NOTES AND NEWS 91 


in the text, of species which occur just outside the area covered by the flora will 
increase its utility in peripheral regions. Finally, in this era of delay, the authors are 
to be congratulated on keeping the intervals between publication of the parts so 
short —ROBERT ORNDUFF, Department of Botany, Duke University, Durham, North 
Carolina. 


NOTES AND NEWS 

NEw DISTRIBUTIONS FOR FouR GRASSES IN OREGON.—NARDUS STRICTA L.—This 
species, which is sparingly introduced in the northeastern United States, has appar- 
ently never before been reported from the Pacific coast. It is principally European 
but is said to be native in Greenland and Newfoundland as well (Fernald, Gray’s 
Man. Bot., 132, 1950; Hubbard, Grasses, 319, 1954). Our specimens were collected 
by J. D. Vertrees in the Fort Klamath region of Klamath County, where the species 
is spreading rapidly in certain fields. Its date of introduction is uncertain. H. A. 
Schoth, of Oregon State University, has told us that he first saw this grass about 
1950, on the Grant Brown ranch in the vicinity of Upper Klamath Lake. It was then 
already well established around fields where Alopecurus was being cultivated. Accord- 
ing to Clapham, Tutin and Warburg (FI. Brit. Isl., 1498, 1957), the species is apo- 
mictic and becomes especially abundant in overgrazed areas. In this and other modern 
taxonomic treatments (cf. Stebbins and Crampton, Recent Advances in Botany, 140, 
1961), Nardus stricta is made a monotypic tribe, the Nardeae, rather than being 
placed artificially in the Hordeae (Hitchcock, Man. Grasses U.S., ed. 2, 277, 1950). 


MOLINIA CAERULEA (L.) Moench—Purple Moor-grass, another widespread species 
of Europe and western Asia, was collected on the coast in Lincoln County, 2 miles 
south of Newport, October 8, 1962 (Chambers 1982). It is occasionally adventive 
in the northeastern United States (Dix, Bartonia 23, 41-42, 1945) where it seems to 
have been introduced both in ballast and in seed lots of lawn and pasture grasses. 
In the locality where it has invaded Oregon, there are only a few plants, but each one 
forms a tall, dense clump and is quite conspicuous. Probably these individuals have 
persisted for many years but have not multiplied to any extent. This would be a 
situation comparable to that observed by Dix in Wayne County, Pennsylvania, where 
the species was locally abundant but limited to a single abandoned field. 


SIEGLINGIA DECUMBENS (L.) Bernh.—Also new to the flora of Oregon, Heath Grass 
was discovered at the same time and place as Molinia (Chambers 1981). Sieglingia 
is a common European plant having about the same distribution as Nardus and is 
native, as well, to Newfoundland (Fernald, op. cit., 129). Its few other occurrences 
in the New World are as an adventive in Washington and California (Hitchcock, 
op. cit.,307). The species is reported to be principally cleistogamous, and this has 
been verified in our specimens from Lincoln County, in which tiny anthers are visible 
in contact with the stigmas even on mature fruits. It is much more abundant than 
Molinia at this locality but is less conspicuous because of its similarity in appear- 
ance to sedges and other grasses. 


The habitat of Molinia and Sieglingia in Oregon is a marshy flat just inland from 
low dunes near the mouth of Henderson Creek. Some associated species are Poten- 
tilla pacifica How., Gentiana sceptrum Griseb., Ranunculus fammula L., and Sisyrin- 
chium californicum (Ker.) Dryand. In the surrounding dunes and sandy flats there 
is evidence of former disturbance—roads, a drainage ditch, dilapitated buildings— 
partially reclaimed by mats of Arctostaphylos uva-ursi (L.) Spreng. and other native 
vegetation. What we have been able to learn about the history of this spot suggests 
that these adventive grasses may have been here for many years, perhaps from before 
the turn of the century. 


The following information was supplied to James M. Howes, of Newport, by 
Jack Fogarty, a long-time resident of Lincoln County. In 1884 the land at the mouth 
of Henderson Creek was owned by B. L. Arnold, who was president of what was then 


92 MADRONO (Vol. 17 


the newly established Oregon Agricultural College, in Corvains. After being cleared 
and drained, the marsh was planted with seeds of various forage grasses and legumes 
provided by Arnold and was called an “experiment station.” It was also at this time 
that the state Agricultural Experiment Station was begun at the College; however, 
the plantation on the coast must have been very short lived, as it was sold to a land 
promoter about 1887. We have not been able to locate a reference to the coastal 
“experiment station” in any early records at Oregon State University. The first annual 
report of the Station was published in 1889 and dealt only with the research then 
being started at Corvallis. The abandoned roads and buildings already mentioned 
are the remnants of the Yaquina Bay Life Saving Station, built in 1896 and operated 
for almost 25 years. Judging from the extent to which the vegetation has recovered, 
there has not been any major disturbance since the close of this station. It is possible 
that Sieglingia and Molinia were introduced here when the land was first brought 
into cultivation and have spread only slightly, if at all, since that time. However, 
without further evidence, one can not rule out an alternative origin from ballast in 
nearby Newport. 


HELEOCHLOA ALOPECUROIDES (Pill. & Mitterp.) Host—Spreading along the Wil- 
lamette River from its probable point of introduction at Portland (Hitchcock, op. 
cit., 433), this species is now quite common on gravelly and sandy banks near Cor- 
vallis. Raven (Leafl. West. Bot. 8:200, 1957) has noted its occurrence in eastern 
Washington as well. Collections at hand include: Seldon s.n., October 15, 1952, 
Benton County, Corvallis Sand and Gravel Company; Dennis 2343, October 29, 1961, 
Linn County, gravel pits just east of Corvallis; Masterson s.n., December 12, 1950, 
Multnomah County, Government Island. In a recent treatment (Lorch, Bull. Res. 
Counc. Israel 11D:94—-96, 1962) this species is placed in Crypsis as C. alopecuroides 
(Pill. & Mitterp.) Schrad. 

Cited specimens are deposited in the herbarium of Oregon State University. We 
are grateful to Thomas R. Soderstrom, James M. Howes, H. A. Schoth, and Harriet 
L. Moore for their assistance in the preparation of this report—Kernton L. CHam- 
BERS and La Rea J. DENNIS, Department of Botany and Plant Pathology, Oregon 
State University, Corvallis. 


CALYPSO BULBOSA IN THE SANTA Cruz MOUNTAINS, CALIFORNIA.—This species 
was recently discovered in the vicinity of Big Basin Redwoods State Park in Santa 
Cruz County. State Park Attendant Herman E. Schlerf found the plants and I iden- 
tified them as Calypso bulbosa (L.) Salisb. (Schlerf & Crandall s.n., May 4, 1963, 
DS). The calypsos numbered about 75 and grew in duff on weathered sandstone in 
a madronfo-tanbark oak-redwood association. This species was previously not known 
south of the vicinity of Mount Tamalpais in Marin County some 50 miles to the 
north—Tuomas A. CRANDALL, Big Basin Redwoods State Park, California. 


NoTES ON TETRACOCCUS AND CHILOPSIS IN BAja CALIFORNIA, MExico.—In Feb- 
ruary, 1960, while seeking shelter with Frank Vasek from a strong wind in an un- 
named wash just west of the highest elevation (ca. 2000 ft.) along the road between 
San Felipe and the Sierra San Pedro Martir, I came upon a well developed plant 
of Tetracoccus hallii Brandegee. During December, 1961, Steven A. Kaune and I 
visited the same ironwood and palo verde wash and after considerable scouting found 
another shrub up the wash from the first (Kaune 204, DS). It is quite possible that 
other plants will be found in the vicinity. 

Another interesting discovery made while camping in a broad, sandy wash emerg- 
ing from a large canyon of the east escarpment of the Sierra San Pedro Martir was 
a number of large specimens of Chilopsis linearis (Cav.) Sweet. Two trees had crown 
breadths of 73 and 67 feet, respectively. Another tree had a basal trunk 4 feet in 
diameter and a crown 40 feet across. This I believe is the largest trunk diameter 
recorded for this species —EpmMuUND C. JAEGER, Riverside Municipal Museum, River- 
side, California. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume I, 1916-1929 . . . . . $5.00 
Volume II, 1930-1934. . . . . 5.00 
Volume III, 1935-1936 PS Ae SLO 
Volume IV, 1937-1938. . . . . 5.00 
Volume V, 1939-1940 . . . . . 5.00 
Volume VI, 1941-1942 ee S00) 
Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 Pe ee 00) 
Volume X, 1949-1950 . . . . = . ~~ 7.00 
Volume XI, 1951-1952 Meer A OMe 30.0) 
Volume XIT, 1953-1954 . . . . 8.00 
Volume XITI, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 gee ort ee E1000 
Volume XVI, 1961-1962 . . .  . 12.00 


Single numbers: 


Volumes Land dis.) 2 3.5) 4 ee. 50 
Volumes II through XIV. . .._.. 1.00 
Volume ey, 5.76...) ae ae 125 
VolumecX Vic. Bertier. he ae 1.50 


The subscription price of MApRONo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


ADRONO 


VOLUME 17, NUMBER 4 OCTOBER, 1963 


Contents 
PAGE 
SOME CORDILLERAN PLANT SPECIES NEW FOR THE 


SIERRA NEVADA OF CALIFORNIA, J. Major and 
S.A. Bamberg 93 


NATURAL AND ARTIFICIAL HYBRIDS OF BESSEYA AND 
SYNTHYRIS (SCROPHULARIACEAE), A. R. Krucke- 
berg and F. L. Hedglin 109 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 116 


ARTIFICIAL INTERGENERIC HYBRIDS OF HELIANTHUS 
AND VIGUIERA, Charles B. Heiser, Jr. 118 


CHROMOSOME NUMBERS IN THE ComposiITaE. VII. 
ADDITIONAL SPECIES FROM THE SOUTHWESTERN 
UNITED STATES AND Mexico, A. M. Powell and 
B.L. Turner 128 


Reviews: Philip A. Munz, California Spring Wildflow- 
ers, California Desert Wildflowers, and California 
Mountain Wildflowers, Roxana S. Ferris, Death Val- 
ley Widflowers, Leonid Enari, Ornamental Shrubs 
of California (Helen K. Sharsmith) ; M. Walter Pes- 
man, Meet Flora Mexicana (Frederick G. Meyer) 140 


NotTES AND News: Howarp E. McMINN 115 
TRAVEL TO THE 10TH INTERNATIONAL BOTANICAL 
CONGRESS, EDINBURGH, AUGUST 3-12, 1964 127 


NOTE ON DAMAGE TO THE HOOKER Oak, Kingsley R. 
Stern; ETHEL BAILEY HiccINns, George E. Lindsay; 
CALIFORNIA BOTANICAL SOCIETY 143 


A WEST AMERICAN JOURNAL OF BOTANY 


UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


EpcArR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 
HERBERT F, COPELAND, Sacramento College, Sacramento, California 
Joun F. Davinson, University of Nebraska, Lincoln 
MitpreD E. Matuias, University of California, Los Angeles 24 
RoBERT ORNDUFF, University of California, Berkeley 
MaArIon OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
IrA L. WiccIns, Stanford University, Stanford, California 


Editor—JoHN H. THomAs 
Dudley Herbarium, Stanford University, Stanford, California 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Herbert L. Mason, Department of Botany, University of California, 
Berkeley. First Vice-President: Paul C. Silva, Department of Botany, University of 
California, Berkeley. Second Vice-President: Robert F. Hoover, California State 
Polytechnic College, San Luis Obispo. Recording Secretary: Mary L. Bowerman, 
Department of Botany, University of California, Berkeley. Corresponding Secretary, 
Margaret Bergseng, Department of Botany, University of California, Berkeley. 
Treasurer: Douglas M. Post, Biology Department, San Francisco State College, San 
Francisco, California. 


1963 | MAJOR & BAMBERG: SIERRA NEVADA PLANTS 93 


SOME CORDILLERAN PLANT SPECIES NEW FOR THE 
SIERRA NEVADA OF CALIFORNIA 


J. Major AND S. A. BAMBERG 


In any floristic study, two aspects of the flora can be the center of 
interest: 1, local endemism; 2, phytogeographical relationships to other 
regions. This paper is concerned with the second, i.e., with some taxa 
found in Convict Creek basin on the east slope of the Sierra Nevada, 
having a unique ecological setting, and disjunct from the Rocky Moun- 
tains, the eastern Great Basin ranges, or the Cascades. The basin con- 
tains at least five geographically disjunct plants, namely Arctostaphylos 
uva-ursi,' hitherto unreported for the Sierra Nevada, and Kobresia myo- 
suroides (Vill.) Fiori & Paol., Scirpus pumilus Vahl, Salix brachycarpa 
Nutt., and Draba nivalts Liljebl. var. elongata Wats. new for California. 
The disjunct presence of Pedicularis crenulata at the mouth of Convict 
Canyon has been known for some time (Munz, 1959). 

On the advice of G. L. Stebbins we selected the Convict Creek Basin 
as a starting point for our studies of Sierra Nevada alpine plant ecology. 
We spent 9-12 June, 6-13 July, and 12-31 August, 1962 in the basin. 
This paper is a first report on our findings. 

To a number of people we are very grateful for herbarium data: M. E. 
Lewis, U. S. Forest Service, Ogden; R. F. Thorne and M. J. Henrickson, 
Rancho Santa Ana Botanic Garden, Claremont; C. L. Porter, University 
of Wyoming; L. D. Potter and W. Martin, University of New Mexico; 
W. P. Cottam, S. Flowers and I. B. McNulty, University of Utah; J. R. 
Murdock, Brigham Young University; J. T. Howell, California Academy 
of Sciences, San Francisco; R. Bacigalupi, University of California; and 
W. A. Weber, University of Colorado. J. T. Howell has been a generous 
advisor, and G. L. Stebbins has been an appreciated source of encourage- 
ment and expert help in the field. We appreciate support from National 
Science Foundation Grant 23317. 


EcoLoGiIcAL UNIQUENESS OF THE CONVICT CREEK BASIN 


Most of the high Sierra Nevada is a monotonous expanse of glaciated, 
gray granodiorite. It does have enclaves of metamorphic rocks, and Con- 
vict Creek basin and the Mount Baldwin area is one of the most inter- 
esting of these and is ecologically unique in several ways. A half mile 
wide strip of coarse-grained, gray marble extends north from Mount 
Baldwin in the southeast corner of the basin past Laurel Mountain on 
the north rim. 

Convict Creek basin is on the desert side of the Sierra Nevada just 
south of Mammoth Mountain, between Leevining and Bishop. Figure 1 


1 Nomenclature corresponds to Munz (1959) except where an author is named. 


MaproNo, Vol. 17, No. 4, pp. 93-144. November 15, 1963. 


94 MADRONO [Vol. 17 
118° 50 W 


118°55 W 
: — 7 038 
LEGEND: =~ ' 


—-—-=— Routes taken on foot. 


ARS Ridges . P y 2 
ish and Game <a 
Bd Racks Experimental \ 2 
1 Station e. 
—— Posses 4, 
> 
ri) Perennial snow. 
Rae 
scale 
¢? 
x4 10, 95é 
( 
( 
37°35 N 
Bev ourel Mtn. 
6, 11,812 o 
JZ 8,250 \% 
5. \\ 2 
CG? r) \2 
G on « McGee Mtn. 
4 Nv 10,871 
Pe ’ x { aps McGee Moraines ~ 
) \ AITSD ty 
Slooay vIn. \ — Il’ 
; ae } t] 5 S —_ 
Ss S A 
G1 } 
4 'S 
4 o 
D 
11,975 6 Gy 
“ Cloverleaf ’ y 
- Sy Cake ) st Mt. Aggie 
e . ' 
rn g y NY BN 58) aa 
: k , Wy lw Springs 
L y 9 Horsetoil 
—s!) 855 ) se Falls 
ae : 
AN = 
~~ XS) Mt. Baldwin 
( 12,614 
\? 
12,2779 © ‘a 
Scheelore 
Mine (ruin) 
(\11,000 
Q5 
4 
Tully Hole 
’ ) 
Cecil Lake Beale ~ eh) ; 
C.% Red Slate min. SEF =o Ww 
13,163 (= —S= 
—_ 
eetitote (7S McGee Pos 0 os 
hn 37° 30 N 
1 u 0 ] 2 Miles 
1 5 0 ] 2 3 4 5 Km 


Fic. 1. Map of the Convict Creek basin, Sierra Nevada. From the U. S. Geological 
Survey topographic map of the Mount Morrison quadrangle, published 1953, 1/62500, 
80 ft contour interval. 


shows salient features and our routes of travel. A geographical discus- 
sion of the area with photographs of the lakes is contained in Reimers, 
Maciolek, and Pister’s bulletin (1955). 

Convict Lake, dammed by terminal moraines at the mouth of Convict 
Creek, is at an elevation of 7580 ft. Cirque floors in the upper part of 
the basin contain lakes and generally lie at elevations of 10000 to 10800 
ft. The highest elevation on the rim of the basin is Red Slate Mountain 
at 13163 ft. Quite extensive remnants of old, subdued surfaces exist above 
12000 ft. 


| 


1963 | MAJOR & BAMBERG: SIERRA NEVADA PLANTS 95 


Geologically the area is very complex. Most of the surface of the basin 
is underlain by dark red, metamorphic hornfels, marble, and vari-colored 
sandstones. The northwest corner of the basin is the usual granodiorite. 
Faulting has been extensive. The geology was mapped and discussed by 
Mayo (1934), and more detailed information for field use was obtained 
through the courtesy of D. C. Ross of the U. S. Geological Survey. 

The basin is not extensively forested, for glaciation has removed soil 
from much of the area, and slopes are so steep that much of the winter’s 
snow avalanches. Detritus from the easily weathered bedrock forms ex- 
tensive scree slopes where bedrock itself is not exposed. Rock glaciers 
are common from the outcrops of more resistant rocks. Much of the 
basin is above the low forest limit at about 10600 ft. Timberline Krumm- 
holz (Pinus albicaulis) is about 1200 ft higher on the granodiorite. 

Of the ecological factors commonly considered as influencing where 
plants occur (Major, 1951), the Convict Creek basin differs little or not 
at all from most of the rest of the Sierra Nevada at comparable altitudes 
in climate, relief, grazing and fire history, lengths of time available for 
plant succession, or available fauna and flora—although the last factor 
evidently deserves more attention than it has yet received. The basin 
basically differs from most of the Sierra Nevada in its large area under- 
lain by calcareous soil parent materials. Some 1371 acres are underlain 
directly by the Mount Baldwin marble. An additional 114 acres is under- 
lain by marshy, meadow sediments predominantly derived from the 
marble. In addition, the widespread hornfels has calcareous lenses, and 
dust accumulated on the snow in winter is calcareous. 

These calcareous substrates immediately affect plants. The mountain 
climate in this basin is insufficiently moist to produce acid humus layers 
over the calcareous parent material, and thus offer possibilities for an 
acidiphile regional climax vegetation to form the zonal plant cover as 
in Braun-Blanquet and Jenny’s (1926) classic case from the Swiss Alps. 
We should note here that Ellenberg (1953) disputes Braun-Blanquet and 
Jenny’s conclusion that the climax Caricetum curvulae forms on strictly 
calcareous rock. He believed from experience in the eastern Alps of 
Austria that admixture of siliceous rock to the calcareous parent material 
was necessary. We could find no evidence that an acidiphile climax vege- 
tation would form on the marble of the Convict Creek basin. However, 
the Sierra Nevada ecosystem differs so much from that of the Alps in 
other respects—notably in climate and flora—that comparisons are 
difficult. 

Associated with the calcareous bedrock are general aridity and con- 
trasting, local, and contiguous areas continuously wet from seepage water. 
These continuously wet seepage areas can also be well-drained. 

Convict Creek basin in general is strikingly less lush than Dana 
Meadows or upper Rock Creek basin located a few miles north and south, 
both of which are in relatively impermeable granodioritic basins. The 
Convict Creek basin itself probably receives as much precipitation as 


96 MADRONO [Vol. 17 


other east-side canyons, but it disposes of it differently. Much water per- 
colates directly through fault-shattered bedrock, and especially through 
solution channels in the marble, into underground channels beyond the 
reach of plant roots. Convict Creek as a matter of fact is doubled in size 
by a large spring at the 8250 ft level; other small springs are common 
on the cliffs; and in McGee Creek immediately to the south the springs 
at Horsetail Falls are notable (fig. 1). A deep topographic sink in shat- 
tered hornfels and marble just south of Laurel Mountain does not hold 
pemanent water in spite of the great quantities of snow which are blown 
into it. 

Not only is water lost by underground seepage, to be concentrated 
and regulated in flow in springs or seeps, but the marble weathers into 
a very coarse soil with very low field capacity. Some such ‘‘sand flats” 
of comminuted calcareous rock forming only an incipient soil may 
appear almost devoid of plants. The impression of aridity on other 
vegetated, flat areas is extreme. The presence of extensive steppe plant 
communities at 10100 to 10700 ft lacking annual plants, of widely- 
spaced bunch-grass physiognomy, with much bare soil surfaces, and 
consisting of such species as Stzpa comata, Oryzopsis hymenoides, Chry- 
sothamnus nauseosus, Artemisia tridentata, Lygodesmia spinosa, Brickel- 
la oblongifolia var. linifolia, and Muhlenbergia asperifolia (in warm 
seeps), does nothing to lessen that impression. Such areas form an ex- 
treme contrast both physiognomically and ecologically with the adjacent, 
lush, green, bright-flowered areas of cold-water seepage where numerous 
rare and disjunct boreal plants occur. 


AN ECOLOGICAL NOTE ON FLORISTICS AND TAXONOMY 


We shall discuss the geographical distribution of the disjunct taxa, 
with notes on their ecology. The best extant guide to their ecology is 
their geographical distribution. The guide becomes more precise the more 
precisely the distribution can be described. The only definitive statement 
on plant distribution is a dot map. This is true both for rare and for 
common plants. Obviously coalescence of dots is a function both of rarity, 
map scale, and frequency of collection. Our effort here is to provide some 
background for dot maps of the species discussed. We welcome more 
information. Because none of the literature statements on the geographi- 
cal distribution of the disjunct plants mentioned is as accurate as possible, 
we have attempted to modernize such statements. Our debt to Hultén’s 
Alaska flora (1941-1950) is obvious, but by using subsequently pub- 
lished local floras even his descriptions can be improved. A further rea- 
son for elaborating on the geographical distribution of our species is that 
the botanical literature mentions these plants and describes their dis- 
tribution repeatedly. The described distributions all differ. Thus, the 
literature is a source of confusion and not of help. It is desirable to try 
to bring some order into this uncorrelated confusion. 


1963 | MAJOR & BAMBERG: SIERRA NEVADA PLANTS 97 


DISJUNCTION OF ARCTOSTAPHYLOS UVA-URSI 


This species is widespread in the northern hemisphere, being circum- 
polar, boreal, and montane to the south. Meusel (1943) described it as 
amphiboreal montane-continental, and grouped it with plants of similar 
chorology such as Allium schoenoprasm L., Goodyera repens (L.) R. Br., 
Populus tremula L.—P. tremuloides, Chimaphila umbellata, Pyrola se- 
cunda, P. uniflora L., and Linnaea borealis. 

In North America A. uva-ursi is found in continental western Green- 
land and from Newfoundland to southern Hudson Bay, southern Kee- 
watin, over Great Bear Lake, to Alaska and south in the mountains to 
Virginia and to northern Indiana and Illinois, Missouri, northern Minne- 
sota, North and South Dakota (Black Hills) and south in the western 
Cordillera to New Mexico, Arizona, Nevada, Oregon, and coastal north- 
ern California. In northern Eurasia it is found from Iceland and northern 
Scandinavia across the continent to Okhotsk and Sakhalin south of the 
treeless arctic for the most part, and south to Ireland and northern 
England, the coniferous forest region of central Europe, in the moun- 
tains of northern Portugal (Hegi, 1927) and southern Spain, eastern 
France, the central Appenines, Albania, Macedonia, and Bulgaria, in 
the Himalayas and Altai (Hegi, 1927), the Caucasus, across northern 
Russia south to the Voronezh region and eastwards across the Urals, and 
scattered in Siberia in the coniferous forest belt south to Lake Baikal 
(Hultén, 1941-1950) plus other pertinent floras). Lipschitz (1961) has 
described the Caucasus plant as a new species, A. caucasica Lipsch., and 
believes it is a Tertiary relict in the flora of the Caucasus. More details 
on the Eurasian distribution of A. uva-ursi are available in Hegi (1927) 
and in Kirchner, Loew, and Schroter (1923). 

Arctostaphylos uva-ursi in California is scattered down the coast to 
Point Reyes north of San Francisco Bay, ison San Bruno Mountain south 
of the Bay (Munz, 1959) and has recently been reported even further 
south at Point Sur in Monterey County (Roof, 1961). It is not on cal- 
careous substrates in these sites. 

Many of the California coastal specimens are aberrant morphologi- 
cally, even for such a variable taxon as A. uva-ursi. They do not all appear 
to be the Rocky Mountain form whereas the plants in the Convict Creek 
basin are. Lenz in working on this problem at the Rancho Santa Ana 
Botanic Garden. 

Specimens from mountain elevations (5665 ft) in Humboldt County 
(Kildale 10660, DS) and under Pinus ponderosa in eastern Del Norte 
County (Applegate 5248, DS) seem to be the widespread A. nevadensis 
of the Sierra Nevada, the Coast Ranges, the Cascades, and the Blue 
Mountains. 

Arctostaphylos uva-ursi is common in the Rocky Mountains and also 
just across the Great Basin, occurring in the Wasatch Mountains of Utah 
and the Ruby Mountains of Nevada (Holmgren, 1942) over 250 mi to 
the northeast of the Sierra Nevada stations. It is not in the Toiyabe 


98 MADRONO [Vol. 17 


Mountains (Linsdale, Howell, and Linsdale, 1952) halfway to the Rubies 
nor in the Deep Creek Mountains between the Rubies and the Wasatch 
(McMillan, 1948). As a mountain plant Merkle (1951) lists it for 
Mary’s Peak in the Oregon Coast Ranges, but Baker (1956) did not 
record it from Iron Mountain in the Rogue River Range some 130 mi 
to the south although some of its normal associates are found there. It 
does occur northeast and west of Klamath Lake in southern Oregon 
under lodgepole pine only a little distance north of the California state 
line (Applegate 3770, DS). To the southwest from Convict Creek, Kear- 
ney and Peebles (1951) doubted its reported occurrence in Arizona. 
However, it was collected by E. K. Douglas in 1934 at 9000 ft in Tsailee 
Canyon, Navajo County (UNM). The species is scattered in the moun- 
tains of northern New Mexico not only along the southern extensions 
of the Colorado Rockies east and west of the Rio Grande River south 
to Santa Fe but also near the Arizona border in the northwestern corner 
of New Mexico (Wooton and Standley, 1915). 

This is the most widespread of the ‘‘new” plants in Convict Creek 
basin (Major & Bamberg 851, 856, 865, 933, 990, DAV). It occupies 
the widest range of altitudes and ecological conditions. It occurs as low 
as 8250 ft at the stream crossing in Convict Creek canyon under blond 
Betula occidentalis and with such plants as Equisetum laevigatum, 
Sphenosciadium capitellatum, Habenaria hyperborea, H. dilatata var. 
leucostachys, Carex hasset, Sisyrinchium halophilum, Solidago multira- 
diata, Castilleja miniata, and Taraxacum officinale. It is in seepage areas 
at 10000 ft or so on more or less physically disintegrated marble with 
Kobresia myosuroides, Potentilla fruticosa, Carex pseudoscirpoidea, 
Salix orestera, S. brachycarpa, Thalictrum alpinum, Botrychium simplex, 
Trisetum spicatum, Solidago multiradiata, Castilleja miniata, and Epilo- 
bium latifolium. It is quite common in almost pure mats on both marble 
and hornfels in seepage areas of cliffs. It is on the ledges of steep cliffs 
where snow accumulates under either Pinus flexilis or P. albicaulis and 
on hornfels knobs along the shore of Bright Dot Lake where snow is 


probably blown off in winter. It even occurs on marble scree just below 
Lake Mildred. 


DISJUNCTION OF KOBRESIA MYOSUROIDES 


This is another circumpolar species classed by Meusel (1943) as 
amphiarctic-alpine-continental with plants of similar chorology such as 
Carex rupestris Bell. ex All. (including C. drummondiana Dewey), C. 
atrofusca Schkuhr (lacking in the North American mountains), Draba 
fladnizensis, Saxifraga cernua L., S. hirculus L., Sibbaldia procumbens, 
Potentilla fruticosa, and Sedum rosea. Meusel’s distribution map can 
be corrected and expanded in America using Porsild’s (1957) and other 
floras for other regions. In the New World K. myosuroides extends from 
Greenland west through Labrador and arctic and subarctic Canada to 
Great Bear Lake and Alaska. It occurs through the Canadian Rockies 


1963] MAJOR & BAMBERG: SIERRA NEVADA PLANTS 99 


with considerable gaps as known at present to northern New Mexico 
and also occurs isolated in northeastern Oregon. In Eurasia it is known 
from Iceland across to Kamchatka and the Chukotsk Peninsula with a 
gap from Scandinavia to the Taimyr Peninsula except for an occurrence 
in the Urals. It has southern, alpine outliers in the Pyrenees, Alps, Ap- 
penines, Balkans, Carpathians, the Asiatic mountains from Dzungaria 
eastwards through northern China and the mountains of southern Siberia 
and Mongolia to Korea and central Honshu in Japan. Closely related 
species occur in the Caucasus and Pamirs to the Himalaya and Tien Shan 
Mountains. 

Kobresia myosuroides is considerably more disjunct at Convict Creek 
than even Arctostaphylos uva-urst. It is also more limited ecologically. 
To the east it occurs first on the slopes of Mount Emmons in the Uinta 
Mountains of Utah (Graham, 1937; Hermann, 1934; Murdock, 1951) 
at the headwaters of the Uinta River and in adjacent areas according to 
Lewis. In Colorado it is abundant in the alpine tundras, and it extends 
south into New Mexico (Wooton and Standley, 1915). To the north of 
the Sierra Nevada it is recorded from alpine summits of the Wallowa 
Mountains in Oregon (Peck, 1961). In other words, the Sierra Nevadan 
stations for K. myosuroides are disjunct by over 500 mi from the closest 
stations to the east and 560 mi to the north. 

Ecologically the K. myosuroides (Major & Bamberg 867, 950, 952, 
1297, 1376, 1464, 1472, DAV; 1498, COLO) occupies a more limited 
range of altitudes and sites in the Convict Creek basin than Arctostaphy- 
los uva-urst. It occurs from about 9700 to 10600 ft in two kinds of 
habitats: 1, seepage areas with or without Arctostaphylos uva-ursi on 
distintegrated marble (soil pH about 8) and with many of the plants 
listed under the higher altitude Arctostaphylos uva-ursi stands, plus 
Parnassia palustris var. californica, Aquilegia formosa, and Habenaria 
hyperborea; and 2, meadows with good drainage but surface still only 
a few decimeters above the water table, either on marl or hornfels (pH 
7.5-8.0), with Salix brachycarpa, Carex pseudoscirpoidea, Juncus bal- 
ticus, Deschampsia caespitosa, Solidago multiradiata, Thalictrum alpi- 
num, Pedicularis attolens, Gentiana holopetala, and Parnassia palustris 
var. californica. Many of the plants typical for the springs and seepage 
areas are missing from these meadows, but the stand surveys recording 
both kinds of communities can probably be arranged in a floristic and 
therefore ecological continuum. 

In general, K. myosuroides is known as a calcicole or a species indif- 
ferent to substratum calcareousness (Schroter, 1926). It is a calcicole 
in Convict Creek basin. It does occur on hornfels substrates as well as 
on marble. However, most areas in the basin are treated with calcareous 
dust blown by the wind from the extensive bare areas of marble and 
marble detritus and accumulated in the winter’s snowfall, and so soils 
from hornfels adjacent to calcareous areas remain basic. On the other 
hand, this Kobresia occurs abundantly and dominantly on James Peak 


100 MADRONO [Vol. 17 


in Colorado on soils of pH (4.6)—5.4—5.6—(6.0) (Cox, 1933) and in the 
Uinta Mountains of Utah at pH 5.6-6.8 (Murdock, 1951). It is a 
climax dominant in the alpine tundras of the Uintas and in Colorado 
(op. cit., Marr, 1961; Weber 1961). Thus, one would expect K. myo- 
suroides to occupy the most highly developed, leached soils in a moun- 
tain climate which provides water in excess of current needs so soil 
leaching occurs. In the Alps for example the subclimax Kobresia (as the 
Elynetum) occupies soils of pH 6(5—7) developed over limestone, but 
it is replaced by the climax Curvuletum on more acid, more highly de- 
veloped soils (Braun-Blanquet and Jenny, 1926). Regionally, zonal 
alpine tundra soils in Colorado dominated by K. myosuroides are (Retzer, 
1956) uniformly acid except where derived from limestone. Acidities may 
be as strong as pH 4.5 although most are in the range of pH 5-6. It is 
possible that K. myosurotdes has one range of edaphic demands in north- 
ern Eurasia and the Sierra Nevada and another in the southern Rocky 
Mountains and the Alps, but competition could equally well explain 
the apparently anomalous tolerances. 


DISJUNCTION OF SCIRPUS PUMILUS 


Only a single station for this diminutive plant (Major 1298, DAV) was 
found in the Convict Creek basin. This species is disjunct from Colorado 
on the east or from Montana to the northeast and is circumboreal in dis- 
tribution although rare everywhere and widely disjunct in its other North 
American and Eurasiatic stations. Its ecology is enigmatic. Raymond 
(1957), Hylander (1945), and Hultén (1958) have discussed the tax- 
onomy, nomenclature, and distribution of S. pumdlus. 

Meusel (1943) classed S. pumilus chorologically as a plant of arctic- 
alpine continental distribution, specifically eurasiatic-alpine continental, 
along with Anemone narcissiflora L., Androsace chamaejasme Wulfen, 
Gentiana algida Pallas, and Aster alpinus L. The taxa selected from this 
chorological group of Meusel’s as familiar all occur also in western 
North America although they are lacking in the East. Therefore they 
could be part of Meusel’s amphiarctic-alpine continental group already 
mentioned under Kobresia mvosuroides, namely those with a gap in 
eastern North America. 

Scirpus pumilus is widespread but very scattered and usually described 
as rare in the Alps from the Dauphine and Cottian Alps to Savoy, the 
high Valais, eastern Graubunden to the upper Adige and on the Eisack 
(Isarco) near Brixen (Bressanone) in the South Tyrol in moist, cal- 
careous, often open sites. Localities (Hegi, 1939) in the Tessin are 
mentioned which more or less connect the eastern and western groups 
of stations in Switzerland. The many stations enumerated by Braun- 
Blanquet and Riibel (1932-1936) would make a small continuous area 
in southeastern Graubunden on the small scale of Hultén’s map. A still 
more eastern station, unmentioned in the floras available to us, is mapped 
in eastern Austria. This may be the unmapped station at Delnice just 


1963 | MAJOR & BAMBERG: SIERRA NEVADA PLANTS 101 


east of Fiume in Jugoslavia mentioned by Hermann (1956). Scirpus 
pumilus is on boggy, calcareous soils in the northern Carpathians of 
Slovakia (Raymond, 1957). Five stations are now known from north- 
ernmost Norway (Nordhagen, 1963). Only on a large scale map can 
these be differentiated from the two stations mapped. In Norway the 
plant occurs locally in wet sites both at sea level and near the birch 
altitudinal limit on dolomite and also on the boundary between the 
subalpine and low alpine regions on calcareous schist (Nordhagen 1963). 
The plant is rare in alpine bogs in the western Caucasus to southern 
Ossetia (Grossgeim, 1949). Boissier (1881) long ago recorded the plant 
from the high Persian Mountains, and both Raymond and Hultén add 
stations. From as far south as 30°N latitude the plant’s area is more or 
less continuous, although actually stations are widely scattered within 
it, into Afghanistan, the Karakorum, and western Tibet. A combination 
of the two maps would give a somewhat altered picture of distribution 
in the western Himalayas. Very isolated occurrences are shown in the 
trans-Volga lowland, in the southernmost Urals, and farther north along 
the Tobol River. Kotov’s (1943) double station near Ufa on meadow 
solonchak soils of the floodplain of the Dema River in Bashkiria is not 
shown on either map. It may be the mentioned trans-Volga station mis- 
placed. The same, or perhaps another station, is mentioned by Hermann 
(1956) at Chkalov (Orenburg) on the Ural River south of Ufa. Other 
lowland stations also occur along the Irtysh River near Omsk and north- 
eastward, west of Tomsk. These lowland stations must be in the saline 
meadows referred to in Soviet floras. The dot shown on both maps on 
the Syr Darya River some 200 km east of the Aral Sea is not mentioned 
in the floras of the USSR or Kazakhstan. It is improbable in this desert, 
but then many of the stations recorded in southern Siberia would seem 
inhospitable to a high altitude or boreal plant. Stations are missing from 
the maps according to the flora of Kazakhstan (Pavlov, 1958) in the 
forest-steppe of the Kokchetavski district and to the southwest in the 
presently internally drained Turgai steppe district where runoff from 
continental glaciation once passed. The Zaisan and Altai districts are 
correctly shown as having the plant, but to the southwest the Dzungarian 
Alatau should have it also. The western and central Tian Shan have the 
plant; the area northwest of Chimkent along the upper Syr Darya should 
also, as Raymond’s map does although his dots extend far into the 
Kyzl Kum desert. Here again the two maps need to be reconciled. To 
the southwest of the Tien Shan the area should be more or less con- 
tinuous, on the map scales used, in the Pamirs and Hindu Kush. In 
Tadzhikistan an irregular area between 40-37°N and 6712-7414°E has 
43 stations listed (Ovchinnikov, 1963). The stations are actually scat- 
tered, occurring in an ecologically wide variety of sites from 1800 to 
4200 m elevation. From the Altai both maps have the area extending 
north along the upper Ob River. The adjacent Angara-Sayan area has 
the plant in several places. On the south side of the Mongolian Altai the 


102 MADRONO [Vol. 17 


plant occurs at Uinchi (46°N, 92°E) in the foothills and in the sink 
of the Bodonchi River where the water vanishes in the desert (Egorova, 
1959). In Mongolia the plant is described by Grubov (1955) as very 
rare, occurring in the Kahangai forest steppe, far to the east in the Mon- 
golian steppe, and adjoining the more or less continuous area toward 
Lake Baikal (Popov, 1957). The area extends north of Lake Baikal, 
southeast toward eastern Mongolia, and into the Chahar province of 
China. An isolated station in northern Szechuan is kept as spp. distig- 
maticus Kuk. 

In North America at the mouth of the Gulf of St. Lawrence the sta- 
tions on the Mingan and Anticosti Islands are well-known (Raymond, 
1957). They are on calcareous substrates. In western Canada are sta- 
tions in Alberta in calcareous bogs (Moss, 1959). The stations in the 
vicinity of Banff, all at valley elevations of 4600 to 5000 ft, have been 
known for some time. A new one is Porsild’s (1959) in a fen on cal- 
careous clay to the north near Jasper along with about a dozen of our 
Convict Creek basin plants. Fernald (1931) listed our plant for two 
British Columbia stations, one of which is actually in Alberta. The plant 
does occur in British Columbia but not near Banff, evidently (Hultén, 
1958). Porsild and Crum (1961) describe the first record for British 
Columbia at Liard Hot Springs where it is “‘very local in wet calcareous 
mud.” Except for the maritime Norwegian stations, this station is the 
farthest north. The plant occurs in western Montana in meadows (Booth, 
1950). There is a disjunct plains station in an alkaline bog just east of 
Saskatoon in Saskatchewan (Fraser et al., 1954) which is very reminis- 
cent of many of the Siberian stations north of the high mountains. The 
widely disjunct station in Colorado is a good one according to Weber 
(1961a) even if the plant has been found only once. Our station in Cali- 
fornia is not that mentioned by Abrams (1923). 

The California disjunction of S. pumilus is very large—about 750 mi 
to the single Colorado locality or to those in western Montana. It is sur- 
passed however by the Norwegian disjunction. 

In the Convict Creek basin S. pumilus occurs in some seepage areas 
or wet meadows at 10200 to 10600 ft elevation with Kobresia mvosu- 
roides, Thalictrum alpinum, Carex pseudoscirpoidea, Epilobium lati- 
folium, Salix brachycarpa, Pedicularis attollens, Habenaria hyperborea, 
Danthonia intermedia, Parnassia palustris var californica, Carex hasset, 
and Aquilegia formosa. These two kinds of permanently wet areas were 
described in discussing the ecology of Kobresia myosuroides. 

There is universal agreement that S. pumilus is everywhere a hygro- 
philous calcicole. The combination in many very continental areas leads 
to describing the plant’s habitat as saline meadows. This tiny plant, which 
is always rare, scattered, and never abundant, combines in a most pecu- 
liar way on alpine or boreal distribution and association with boreal or 
even arctic-alpine plants with occurrence in lowland, steppe habitats. 

Ecological segregation between S. clementis and S. pumilus is as sharp 


1963 | MAJOR & BAMBERG: SIERRA NEVADA PLANTS 103 


in the Convict Creek Basin as that between S. caespitosus L. and S. pumi- 
lus is elsewhere. The former is restricted to subirrigated meadows or bogs 
on granitic alluvium or peat and the latter to subirrigated meadows or 
bogs on calcareous alluvium. Scirpus caespitosus var. delicatulus Fern. 
(Fernald, 1950) of “Calcareous gravels, shores and cliffs” may perhaps 
be interpreted as a form occupying an azonal habitat where competition 
with zonal plants is weakened on a unique soil parent material (Gankin 
and Major, 1963). 


DISJUNCTION OF SALIX BRACHYCARPA 


Salix brachycarpa is a cordilleran and boreal American willow related 
to the circumpolar, arctic-montane S. glauca L. and its associated com- 
plex. The systematics of this group is not a subject of universal agree- 
ment. We have relied mostly on the geographical, taxonomic, and nomen- 
clatural treatments in Raup (1959), Hultén (1941-1950), Argus (1957), 
and Fernald (1950). Other floras have been consulted when pertinent. 
Raup (1959) gives rough outline maps of the northern, American species’ 
distributions, and these maps should adequately represent current in- 
formation on geographical distribution. Ecological notes agree that all 
the taxa in this complex occupy moist or wet, often boggy soils. Few data 
on vegetation associated with these species in North America have been 
found and very little on soil requirements. 

The distribution of S. glauca in Eurasia is from Iceland through the 
Scandinavian mountains, arctic and subarctic USSR including Novaya 
Zemlya, and south in the mountains including the Urals, Sayan and 
Altai both in the USSR and in Mongolia, the mountains around Lake 
Baikal and of Dahuria and Khangai in Mongolia, and farther east the 
Okhotsk and Zee-Bureinsk regions. In North America it occurs from 
arctic Alaska across to Greenland and Ungava and south in Alaska and 
the Rocky Mountains to Lake Athabaska in Saskatchewan and to the 
Waterton Lakes Park region and northern Washington and Montana 
(Raup, 1947). There is no agreement, however, as to the southern limit 
of the species. In Saskatchewan Fraser e¢ al. (1954) have it only to Lake 
Athabaska but Raup to the southern part of the province. Moss (1959) 
has it in only northwestern Alberta; but \’orsild (1959) says it is com- 
mon near Banff. Salix brachycarpa extends southward from Canada to 
the Wenatchee Mountains of central Washington on serpentine, the 
Wallowa Mountains of northeastern Oregon. It is also known in Idaho 
from the Tetons, Bear River Range, and Sawtooth Mountains, in 
Wyoming in the Medicine Bows, Wind Rivers, and Gros Ventre to 
western Yellowstone National Park, Absarokas, and Beartooth, in Utah 
from two localities, and in the mountains of Colorado. 

Salix brachycarpa at Convict Creek (Major & Bamberg 894, 937, 
1279, 1378, DAV) is evidently disjunct by 560 mi from the Wallowa 
Mountains to the north, by 310 mi from the southern Utah station north- 
west of the Pink Cliffs on the south edge of the Markagunt Plateau north 


104 MADRONO [Vol. 17 


of Zion National Park, by 440 mi from the locality at Alta in the Wasatch 
Mountains east of Salt Lake, and by about 530 mi to the Idaho localities. 

In Convict Creek basin S. brachycarpa is abundant from about 9900 
to 10600 ft. It is found in summer-moist seepage areas which are well- 
drained and in the well-drained parts of the alluvial, marly flat above 
Lake Mildred. It is limited to calcareous soils. Its associates have been 
mentioned under the higher altitude Arctostaphylos uva-ursi stands and 
both the seepage and meadow Kobresia myosuroides stands. 

In the Medicine Bow Mountains of Wyoming Salix brachycarpa was 
found in wind-protected parts of an alpine fellfield dominated by Carex 
rupestris, Geum rossu, and Polygonum viviparum (Bliss, 1956) on acid 
(pH 5.0) soils frequently drier than the 15 atm. percentage and developed 
from quartzites. 


DISJUNCTION OF DRABA NIVALIS VAR. ELONGATA 


In 1957 Stebbins found specimens of this variety on an ascent of 
Mount Baldwin and recognized it as new for the Sierran flora. With his 
help it was again found at several localities in the Convict Creek basin 
during the summer of 1962. 

Draba nivalis var. elongata occurs northwards through Wyoming, from 
the isolated LaSal Mountains of southeastern Utah, from the Wasatch 
and Uinta Mountains of Utah north and west through the Deep Creek 
Mountains to the Ruby Mountains of Nevada, northwards from central 
Idaho, northwards from the Wallowa Mountains of Oregon and the 
central Cascades of Washington. The disjunction to the Convict Creek 
localities is thus 560 mi to the north and over 250 mi to the northeast. 
The disjunction to the north is identical to that of Kobresia myosuroides 
but to the northeast is equal to that of Arctostaphylos uva-urst. 

Ecologically this taxon occupies quite a different habitat in the Con- 
vict Creek Basin as compared with Arctostaphylos uva-ursi and Kobresia 
myosuroides. Draba nivalis var. elongata (Major & Bamberg 1296, 1432, 
DAV) was found at about 10300 to 10800 ft and again at 11750 ft on 
wet, disintegrated marble scree or in wet crevices in the marble. The 
sites are all on north-facing slopes, steep, and covered by snow until 
very late in the season. The highest site was covered by over 2 m of 
snow in the middle of July but was snow-free the last week in August, 
although at 11:30 a.m. the ground was still frozen in the shade. The 
associated vegetation is very open with scattered individuals of such 
plants as Oxyria digyna, Crepis nana, and Arenaria rossi. 


DISJUNCTION OF PEDICULARIS CRENULATA 


Munz (1959) described the disjunct occurrence of Pedicularis crenu- 
lata in the Sierra Nevada as ‘‘near streams, ca. 7000 ft, Convict Creek, 
Mono County; to Wyoming, Colorado.” The species was collected here 
in 1925, 1927 and 1933 by Peirson (JEPS) and recently studied by 
Sprague (1962) in regard to pollination. Her reported station having 


1963 ] MAJOR & BAMBERG: SIERRA NEVADA PLANTS 105 


‘(25-30 white-flowering plants” appeared to us to be a common kind of 
mountain meadow along the outlet of Convict Lake and 500 ft higher 
in elevation than Peirson’s earlier stream-side collections. All these 
meadows are grazed by the local pack stock and have been heavily cattle- 
grazed. They have more the aspect of mountain meadows in the Great 
Basin or Rockies than of the Sierra. We could not find the plant in 1962, 
but there is no reason to believe it does not still occupy the area. 

The distribution of P. crenulata is not precisely known. Tidestrom 
(1925), Rydberg (1922), and Sprague (1962) list it for Nevada. It was 
collected at Duck Creek at 7300 ft in the Schell Creek Range near Ely, 
Nevada (Jones, June 30, 1893, JEPS). It occurs in Wyoming and Colo- 
rado to New Mexico. Harrington (1954) listed it from the western half 
of Colorado at 7000 to 9500 ft. It extends at least from the Gunnison 
River southeast into the Sangre de Cristo Range to the headwaters of 
the Pecos River, north into the Elk Mountains, east to the Pike’s Peak 
region, and northeast to the Continental Divide, thence north along the 
ranges bordering the divide and in the Park Range, into Wyoming in the 
Medicine Bow Mountains and along the upper tributaries of the North 
Platte River, and against along the upper Green River at Sublette and 
Daniel, on the Black’s Fork of the Green River at Fort Bridger, and in 
Yellowstone National Park in Montana. Its area evidently does not 
include the Tetons and Jackson Hole (Shaw, 1958; Reed, 1952). It 
seems to be disjunct in the Sierra Nevada by 250 mi from the Nevada 
locality, disjunct in Nevada by 280 mi from the lower Green River 
station or 370 mi from the nearest of the numerous central Colorado sta- 
tions, disjunct in New Mexico by up to 90 mi from the southern Colorado 
stations, disjunct along the lower Green River by 160 mi and along the 
upper Green River by 220 mi from the southern Wyoming stations, and 
disjunct in Yellowstone National Park by 130 mi from the upper Green 
River. Possibly a more complete herbarium and field search would fill 
in some of these gaps. 

Could it be a recent introduction in some of these places, and specifi- 
cally at Convict Creek? The note by Coulter and Nelson (1909) might 
suggest this: “‘. . . in grassy mountain meadows; spreading rapidly in 
irrigated meadows and becoming a pest.” Or this behavior may have 
simply reflected the response im situ of an unpalatable plant to current, 
turn of the century abuse of the vegetation by overgrazing of domestic 
livestock. Its persistence and considerable area of occupancy at Convict 
Creek does not suggest a recently adventive plant. And there is at Con- 
vict Creek “‘a remarkable correlation between pollinator and flower... 
in the parallel size and curvature of the head and thorax of Bombus 
fervidus and that of the galea of P. crenulata. They more obviously com- 
plement each other in these respects than do any other observed species 
of Pedicularis and their pollinators” (Sprague, 1962). This correlation 
does not suggest an adventive plant even though this Bombus does not 
extend to the Rocky Mountains. 


106 MADRONO [Vol. 17 


The habitat of P. crenulata is low-altitude, subirrigated mountain 
meadows bordered by willows and aspens. Deschampsia caespitosa is a 
common associate, with Carex spp., Rosa spp., and Smilacina stellata 
under the shrubs. The bordering xeric, well drained community may 
have Artemisia tridentata, and Elymus cinereus. 


DISCUSSION 


The Convict Creek basin is both ecologically and floristically unique 
in the Sierra Nevada. Fortunately it adjoins quite typical Sierra Nevada 
vegetation on granitic substrates in the same drainage basin. This juxta- 
position will allow contiguous comparisons to be made between the 
unique and the usual Sierra Nevada plant communities and their ecology. 
Perhaps the local uniqueness will allow correlation with other western 
North American mountain vegetation. This western mountain vegetation 
forms a unit whose geographically separated parts will be understood 
only in relation to the whole. Vegetation studies, including their ecologi- 
cal relationships, will enlighten floristic, including historical, relation- 
ships and vice versa. 

While the Convict Creek basin is ecologically unique, it should be 
mentioned that the ecologically unique substrate to be found in the area, 
namely marble and its mechanically weathered products, is not the only 
substrate on which the floristically unique plants are found. All the 
six taxa mentioned occur on soils derived both from calcareous parent 
materials and also from noncalcareous metamorphics. It is true that 
none of their stations are on granodiorite, and that Convict Creek horn- 
fels soils are often calcareous because of snowpack accumulation of 
windblown calcareous dust. But we are very poorly informed on the 
chemistry of formation of Sierra Nevadan soils on granodiorite (pedocals 
form on granodiorite in many climates), and other areas of calcareous 
metamorphic rocks exist in the Sierra Nevada yet none have been re- 
ported to harbor these six taxa. There is no one-to-one correspondence 
here between edaphic specialization and a floristic distinctiveness. 

This last conclusion is bolstered by the extra-Sierra Nevadan edaphic 
behavior of these six taxa. Judging from the ecological literature, all the 
six disjunct taxa are indifferent outside the Sierra Nevada so far as 
calcicoly is concerned. Unfortunately, herbarium labels are usually de- 
ficient in such data, floras often ignore the problem, and insufficient 
systematic ecological and descriptive vegetation work has been done. 

The six disjunct plants discussed above form a kind of equiformal pro- 
gressive area in Hultén’s sense (1937). They are North American cor- 
dilleran plants with boreal and arctic, and hence circumpolar, affinities. 
Because most are so wide ranging, they cannot be easily ascribed to any 
known or hypothesized centers of dispersal or persistence. And Hultén 
specifically excluded from his discussion species such as Pedicularis 
crenulata, known only from south of the ice of continental Pleistocene 
glaciation. Therefore we prefer not to discuss Hultén’s actual disposition 


1963 ] MAJOR & BAMBERG: SIERRA NEVADA PLANTS 107 


of these species. He looked on them from the viewpoint of the boreal, 
circumpolar biota. Our center of focus must be the cordilleran flora. This 
flora obviously cannot be divorced from its boreal and arctic, circum- 
polar affinnities, but it has recruited many members from adjacent low- 
land floras. 

The meaning of the equiformity and progessiveness of distributional 
areas exhibited by our six species must be left to further, systematic 
floristic studies, but it is difficult to believe that we have here either a 
simple coincidence of distributional areas or a distribution pattern based 
solely on ecological demands of the six species concerned. 


University of California, Davis 
Department of Botany 


LITERATURE CITED 


ABRAMS, L. 1923. Illustrated flora of the Pacific States. Vol. 1. Stanford Univ. Press. 

Arcus, G. 1957. The willows of Wyoming. Univ. Wyo. Publ. 21:1-63. 

Baker, W. H. 1956. Plants of Iron Mountain Range, Oregon. Am. Midl. Nat. 56:1-53. 

Buiss, L. C. 1956. A comparison of plant development in microenvironments of arctic 
and alpine tundras. Ecol. Monogr. 26:303-337. 

Botssigr, E. 1881. Flora orientalis. Vol. 5. H. Georg, Geneva. 

BootH, W. E. 1950. Flora of Montana. Vol. 1. Montana State Coll., Bozeman. 

BRAUN-BLANQUET, J. and H. JENNY. 1926. Vegetationsentwicklung und Boden bildung 
in der alpinen Stufe der Zentralalpen (Klimaxgebiet des Caricion curvulae). 
Denkschr. Schweiz. Naturf. Ges. 63:175-349. 

and E. RUBEL. 1932-1936. Flora von Graubunden. 4 Lfg. Verott. Geobot. 

Inst. Riibel Heft 7. 

Coutter, J. M. and A. NEtson. 1909. New manual of botany of the central Rocky 
Mountains (vascular plants). American Book Co., N.Y. 

Cox, C. F. 1933. Alpine plant succession on James Peak, Colorado. Ecol. Monogr- 
3:299-372. 

Ecorova, T. V. 1959. Additions to the cyperaceous flora of the Mongolian Peoples’ 
Republic. [In Russian.] Bot. Mat. Gerb. Bot. Inst. Komarov 19:79-82. 

ELLENBERG, H. 1953. Fuhrt die alpine Vegetations- und Bodenentwicklung auch aut 
reinen Karbonatgesteinen zum Krummseggenrasen (Caricetum curvulae) ? Ber. 
deutsch. Bot. Ges. 66:241-6. 

FERNALD, M. L. 1931. Scirpus pumilus in the Rocky Mountains. Rhodora 33:23-4. 

. 1950. Gray’s manual of botany. American Book Co., N. Y. 

Fraser, W. P., R. C. Russert, G. F. LEpINGHAM, and R. T. CoupLanp. 1954. An 
annotated list of the plants of Saskatchewan. Univ. Saskatchewan, Saskatoon. 

GankIn, R. and J. Major. 1963. Biology of Arctostaphylos myrtifolia Parry. Ecol- 
ogy (in press). 

GraHaAM, E. H. 1937. Botanical studies in the Uinta Basin of Utah and Colorado. 
Ann. Carnegie Mus. 26. 

GrossGEIM, A. A. 1949. Manual of Caucasus plants. [In Russian.] Gos. Izd. Soviet 
Nauka, Moscow. 

HarrincTon, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver. 

Hec1, G. Illustrierte Flora von Mittel-Europa. Vol. 2, 1939; Vol. 3 (1), 1957; Vol. 
5(3), 1927. Hanser, Munich. 

HERMANN, F. 1956. Flora von Nord- und Mitteleuropa. Gustav Fischer, Stuttgart. 

HERMANN, F. J. 1934. A new Cardamine from the Uinta Mountains, Utah. Rhodora 
36:409-412. 

Hortmcren, A. H. 1942. Handbook of the vascular plants of northeastern Nevada. 
Utah State Agric. Coll., Logan. 


108 MADRONO [Vol. 17 


Hu ten, E. 1937. Outline of the history of arctic and boreal biota during the Quater- 
nary period. Bokforlags Aktiebolaget Thule, Stockholm. 
. 1941-1950. Flora of Alaska and Yukon. Lunds Univ. Arsskr. 
. 1958. The amphi-atlantic plants and their phytogeographical connections. 
Kungl. Svensk Vetenskapsakademiens Handl. Fjarde Ser. 7:1-340. 
HyYyLaAnpvErR, N. 1945. Nomenklatorische und systematische Studien iiber nordische 
Gefasspflanzen. Uppsala Univ. Arsskr. 1945 :1-377. 
Kearney, T. H. and R. H. PEEBLEs. 1951. Arizona Flora. Univ. Calif. Press, Berkeley. 
KircHner, O., E. Loew, and C. ScHROTER. 1923. Ericaceae in Lebensgeschichte der 
Blutenpflanzen Mitteleuropas. 4(1), Lfg. 23/24. Ulmer, Stuttgart. 
Kortov, M. I. 1943. New materials on the flora of Bashkiria. I. [In Russian.] Bot. 
Zhurnal 28:117-122. 
LiyspaLe, M. A., J. T. Howe tt, J. M. Linspare. 1952. Plants of the Toiyabe Moun- 
tains area, Nevada. Wasmann Jour. Biol. 10:129—200. 
Liescuitz, S. Yu. 1961. Remarks on a new Caucasian Arctostaphylos—A. caucasia 
Lipsch. [In Russian.] Bot. Mat. Gerb. Bot. Inst. Komarov 21:289-291. 
Major, J. 1951. A functional, factorial approach to plant ecology. Ecology 32:293- 
412. 
Marr, J. W. 1961 Ecosystems of the east slope of the Front Range in Colorado. Univ. 
Colo. Stud. D. 8:1-134. 
Mayo, E. B. 1934. Geology and mineral deposits of Laurel and Convict Basins, South- 
western Mono County, California. Calif. Jour. Mines and Geol. 30:79-88. 
McMirian, 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. 
MERELE, J. 1951. An analysis of the plant communities of Mary’s Peak, western 
Oregon. Ecology 32:618—640. 
MeEuseEL, H. 1943. Vergleichende Arealkunde. Borntrager, Berlin. 
Moss, E. H. 1959. Flora of Alberta. Univ. Toronto Press. 
Muwnz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley. 
Murpock, J. R. 1951. Alpine plant succession near Mount Emmons, Uinta Moun- 
tains, Utah. M. S. thesis, Brigham Young Univ., Provo. 
NorDHAGEN, R. 1963. Om Crepis multicaulis (Led.) og dens utbredelse i Norge, 
arktisk Russland og Asia. Blyttia 21:1—42. 
OvcHINNIKoV, P. N., ed. 1963. Flora of Tadzhikistan. [In Russian.] Vol. 2. Akad. 
Nauk USSR, Moscow. 
Paviov, N. V., ed. 1958. Flora of Kazakhstan. [In Russian.] Vol. 2. Izd. Akad. Nauk 
Kazakhskoi SSR, Alma Ata. 
Peck, M. E. 1961. A manual of the higher plants of Oregon. Binfords and Mort, 
Portland. 
Popov, M. G. 1957. Flora of Central Siberia. [In Russian.] Vol. 1. Akad. Nauk USSR, 
Moscow. 
Porsitp, A. E. 1957. Illustrated flora of the Canadian Arctic Archipelago. Natl. Mus. 
Canada Bull. 146. 
—. 1959. Botanical excursion to Jasper and Banff National Parks, Alberta. 
Natl. Mus. Canada, Ottawa. 
and H. A. Crum. 1961. The vascular flora of Liard Hotsprings, British 
Columbia, with notes on some bryophytes. Natl. Mus. Canada Bull. 171:131-197. 
Raup, H. M. 1947. The botany of southwestern Mackenzie. Sargentia 6:1-275. 
. 1959. The willows of boreal western America. Contr. Gray Herb. 185:3-95. 
Raymonp, M. 1957. Notes sur le Scirpus pumilus Vahl. Contr. Inst. Bot. Univ. 
Montréal 70:83-93. 
Reep, J. F. 1952. The vegetation of the Jackson Hole Wildlife Park, Wyoming. 
Am. Midl. Nat. 48:700-729. 
Rermers, N., J. A. MacroLex, and E. P. Pister. 1955. Limnological study of the 
lakes in Convict Creek Basin, Mono County, California. Fishery Bull. Fish and 
Wildlife Service 56:437-503. 


1963] KRUCKEBERG & HEDGLIN: BESSEYA AND SYNTHYRIS 109 


ReETzeER, J. L. 1956. Alpine soils of the Rocky Mountains. Jour. Soil Science 7:22-32. 

Roor, J. B. 1961. The manzanitas of California’s Point Sur region. Leafl. West. Bot. 
9:188-196. 

RypBeErG, P. A. 1922. Flora of the Rocky Mountains and adjacent plains. New York. 

SCHROTER, C. 1926. Das Pflanzenleben der Alpen. Raustein, Zurich. 

SHAw, R. J. 1958. Vascular plants of Grand Teton National Park. Am. Mid]. Nat. 
59:146-166. 

SPRAGUE, E. F. 1962. Pollination and evolution in Pedicularis. Aliso 5:181—209. 

Tipwestrom, I. 1925. Flora of Utah and Nevada. Cont. U.S. Nat. Herb. 25. 

Weser, W. A. 1961 Handbook of plants of the Colorado Front Range. 2nd ed. Univ. 
Colorado Press, Boulder. 

—_———. 1961a. Additions to the flora of Colorado. III. Univ. Colo. Stud. D. 7:1-26. 

Wooton, E. O. and P. C. STANprEY. 1915. Flora of New Mexico. Cont. U. S. Nat. 
Herb. 19. 


NATURAL AND ARTIFICIAL HYBRIDS OF BESSEYA 
AND SYNTHYRIS (SCROPHULARIACEAE) 


A. R. KRUCKEBERG AND F. L. HEDGLIN! 


_ Within the Scrophulariaceae is a group of five genera well set apart from 
others in the family (Pennell, 1933). All are characterized by basal, 
petiolate leaves, scapose inflorescences, and weakly zygomorphic (veron- 
ica-like) flowers. Two of the genera, Synthyris and Besseya occur in 
western North America, from sea-level to timberline. Species of Synthyris 
are either woodland inhabitants or occur as elements of the snow-flush 
flora of high montane slopes. Besseya species are less mesic in habitat; 
the most common species, B. rubra, occurs in open yellow-pine forests 
or the bunch grass-forb-shrub vegetation type. 

The intriguing distribution of species in Besseya and Synthyris as well 
as in their Eurasian relatives has prompted a long-range study of the 
clan. Morphological, cytological, and breeding criteria will be sought to 
determine the relationships of the highly disjunctively distributed species. 


NATURAL HYBRIDS 


Early in the study it was called to our attention that B. rubra (Dougl.) 
Rydb. and S. missurica (Raf.) Penn. underwent sporadic hybridization in 
the Clearwater River drainage of west central Idaho. The first collections 
of the hybrid were made by Fred Warren, a student of Harold St. John 
at Washington State College. Living plants of this initial collection are 
still growing in Carl English’s garden in Seattle and have been examined 
by the authors. 


1 The authors acknowledge with gratitude the aid given by Carl S. English, Jr., 
who provided us with living plants, C. Leo Hitchcock, for counsel during the study, 
and to the several collectors who furnished buds and living plants. Photographs by 
Audio-Visual Services, University of Washington. A portion of the research was 
supported by funds granted the senior author from the State of Washington Initiative 
No. 171 Fund. 


110 MADRONO [Vol. 17 


In March of 1958, following directions given by English, the authors 
located an area near Kamiah, Idaho, where the hybrids occurred. The 
site had been disturbed by logging about ten years earlier and signs of 
spring cattle-grazing were noticed. Second-growth ponderosa pine and 
douglas fir were the dominant trees, and Ribes, Symphoricarpos, Rosa, 
and Holodiscus constituted the major shrub vegetation. The terrain was 
dominated by a broad, gently rounded ridge running northeast to south- 
west. Besseya rubra grew on the open, grassy crown of the exposed ridge, 
while S. missurica occurred on the cool, shaded northwest-facing slope, 
among shrub thickets. The few hybrid plants were found here and there 
in a zone between the areas of the two species, closest to the lower edge 
of the area occupied by B. rubra, and following the contour on the 
northwest-facing slope a few feet below the edge of the ridge. Both species 
as well as the hybrids were in bud and full bloom. 

No insects were noticed visiting the plants while the authors recon- 
noitered the area, though hymenopterons have been observed on other 
species of Synthyris elsewhere. The pollination necessary for hybridiza- 
tion was probably performed by insect vectors. 

Inflorescences and buds were taken for later study prior to digging 
hybrid plants for transplanting to the greenhouse at the University of 
Washington. Several inflorescences of the parental species were also 
collected. 

Comparison of the hybrid and parental plants revealed the hybrids to 
be intermediate in several characters (table 1, and fig. 1a). Most striking 
of all was the corolla of the hybrids—perhaps half the size of showy- 
flowered S. missurica. Besseya rubra is apetalous (Hedglin, 1959). 


FERTILITY AND CYTOLOGICAL BEHAVIOR OF NATURAL HYBRIDS 


Pollen Stainability: 19.5 %—Hybrid; 88.6%—S. missurica; 89.2% —B. rubra. 

Hybrid seed: None from selfing the hybrids. 

Backcross seed: A very few; 2—3 backcross progeny recovered. 

Meiosis: n = 12 pairs in both species and commonly 12 pairs in hybrid. Of 426 meiotic 
figures examined, 186 (42.7%) were abnormal (one or more lagging chromo- 
somes). Metaphase II figures with laggards: 70/102 (70%) (fig. 2a,b). 


ARTIFICIAL HYBRIDS 


Attempts to synthesize the hybrid from its putative parents met with 
easy and rapid success. Greenhouse hybrids, made in both directions, 
flowered the second year after crossing. Though somewhat more variable, 
the synthetic hybrids were a good match for the Kamiah natural hybrids. 
Some of the F, plants approached Synthyris in morphology, especially 
in having showier corollas than the natural hybrid; others were good 
intermediates (fig. 1b). 

Pollen stainability: 7.5 to 40.0% stainable pollen. 
Meiosis: 2, rarely numerous laggards at metaphase I & II; S. missurica « B. rubra 


156/231 (67.5%) irregular figures at metaphase I; B. rubra < S. missurica 
134/278 (46.5%) irregular figures at metaphase I (fig. 2c,d). 


1963| KRUCKEBERG & HEDGLIN: BESSEYA AND SYNTHYRIS 111 


Fic. 1. Inflorescences and single basal leaves of parents, natural hybrid, and arti- 
ficial hybrid, B. rubra and S. missurica: 1a, B. rubra (left), natural hybrid (center), 


and S. missurica (right) ; 1b, B. rubra (left), artificial hybrid (center), and S. mis- 
surica (right). 


112 MADRONO [Violea7 


TABLE 1. COMPARISON OF THE NATURAL HYBRID WITH ITS PARENTS 


FEATURE S. missurica HYBRID B.rubra 
leaf shape reniform, cordate ovate, cordate elliptic-ovate, truncate 
leaf pubescence — glabrous glabrate with age villous-pubescent 
scape pubescence nearly glabrous white-villous, gla- villous-puberulent 

brate below 
number of bracts 3 to 7 5 to 12 12 to 15 
bract shape widely obovate, rhombic to obovate, lower obovate to elliptic, crenate- 
slightly toothed slightly petioled, toothed dentate, lower petioled 

sepal shape lanceolate, entire lance-ovate, irreg. margins ovate, irreg. toothed 
sepal size 3mm long 4mm long 6 mm long 
corolla size 6-7 mm long 4 (6)'mm long absent 
corolla color blue purple 
stamen color blue purple red 


1 One plant had flowers with the corolla 6 mm long and light blue. 


OTHER HYBRIDS 


At the same time that the artificial intergeneric hybrid was made, 
crosses between other species of Synthyris were tried. To date only the 
larger-leaved woodland species have been used as parents: the high- 
montane, laciniate-leaved species rarely flower in cultivation. Syvnthyris 
platycarpa, S. missurica, S. reniformis, S. schizantha, and S. stellata have 
been intercrossed; all are n = 12. To date, only the crosses, S. missurica 
x S. reniformis and S. missurica X S. platycarpa have given F, hybrids. 
Flowering plants of S. missurica & S. reniformis are shown (fig. 3); the 
hybrids approach reniformis in stature and inflorescence pattern, but 
definitely reflect the influence of missurica. Of the nine hybrid plants, 
seven had no stainable pollen. Meiotic figures, though not severely ab- 
normal, had a fairly high frequency (ca. 50%) of aberrant figures (one 
or more laggards and/or univalents, fig. 2d,e). 


Other interspecific hybrids will be attempted as the opportunity 
permits. 


CHROMOSOME NUMBERS IN BESSEYA AND SYNTHYRIS 


Chromosome counts have been made on a number of taxa in Besseya 
and Synthvris (table 2). All taxa but one thus far sampled are diploids, 
with the gametic number of 12 chromosomes. The one exception, Besseya 
plantaginea, is a tetraploid. We eventually plan to obtain counts on all 
North American taxa as well as on members of the European and Asian 
genera, Wulfenia, Lagotis, and Picrorhiza. Cytological material of any 
of these will be much appreciated. 


CONCLUSIONS 


The occurrence of the natural hybrid, S. missurica « B. rubra, ina 
habitat of disturbed conditions, offers opportunity for introgression, either 


1963] KRUCKEBERG & HEDGLIN: BESSEYA AND SYNTHYRIS 113 


E 


Fic. 2. Abnormal meiotic behavior of hybrids of Synthyris and Besseya: 2a, meta- 
phase II with two lagging univalents, B. rubra * S. missurica (natural hybrid) ; 
2b, pollen tetrad with four spores and one microcyte (natural hybrid) ; 2c, meta- 
phase I with 5 univalents off metaphase plate, B. rubra x S. missurica (artificial 
hybrid) ; 2d, pollen tetrad with four spores and two microcytes (artificial hybrid) ; 
2e, metaphase I with eight univalents off metaphase plate, S. reniformis X S. mis- 
surica; 2{, pollen tetrad with four spores and two microcytes, S. reniformis xX S. 
missurica. 


114 


MADRONO 


TABLE 2. CHROMOSOME NUMBERS IN BESSEYA AND SYNTHYRIS 


SPECIES 


Bessevya bullii (Eat.) Rydb. 


B. plantaginea (James) Rydb. 


B. rubra (Dougl.) Rydb. 


Synthyris missurica (Rat.) 
Penn. 


S. pinnatifida Wats. var. 
pinnatifida 


S. pinnatifida var. canescens 
(Penn.) Cronq. 


S. platycarpa Gail & Penn. 


S. reniformis (Dougl.) Benth. 


S. stellata Penn. 


S. missurica X B. rubra 


LOCALITY AND VOUCHER 
Wolf Lake, Jackson Co., Michigan. 
Gillett 1148* 


White Mountains, Apache Co., Arizona 
Kruckeberg 4583* 


E of Kooskia, Idaho Co., Idaho. 
Hedglin 17* 


Pattee Canyon, Missoula Co., Montana. 


Preece 2265 

Clearwater River, Idaho Co., Idaho. 
Hedglin 16* 

Tony Grove Lake, Cache Co., Utah. 
Hedglin 31 


Bloomington Lake, Bear Lake Co.., 
Idaho. Hedglin 35* 


White Cloud Mountains, Custer Co., 
Idaho. Kruckeberg 4536* 


W of Challis, Custer Co., Idaho. 
Hedglin 39 

Indian Hill, Idaho Co., Idaho. 
Kruckeberg 4109* 


Marin Co., California. McMillan 1931 
(McMillan, 1949) 


New Era, Clackamas Co., Oregon. 
Hedglin 5* 

Drain, Douglas Co., Oregon. 
Hedglin 1 


Pierce Co., Washington. 
Kruckeberg s.n. 


[Vol. 17 


n NUMBER 


12 


24 


Columbia River Gorge, Multnomah Co., 12 


Oregon. Hedglin 7* 


Kamiah, Lewis Co., Idaho. 
Hedglin 18* 


* Prepared slides available (WTU). 


locally or extensively. Fertility, though low, is sufficient for slow infiltra- 
tion of genetic material into one or the other parental species. As yet 
there is no field evidence for introgression and it appears that as yet the 
intergeneric hybrid is of rare occurrence and is not in the process of 
contaminating either of the parents. The fact that this natural inter- 
generic hybrid and its artificial counterpart is more fertile than the 
wholly sterile hybrid between S. reniformis and S. missurica suggests 
that species in Synthyris and Besseya are genetically congeneric. Should 
the natural hybrid lead to introgressant populations, it would seem even 


1963] KRUCKEBERG & HEDGLIN: BESSEYA AND SYNTHYRIS 115 


Fic. 3. Inflorescences and single basal leaves of S. reniformis (left), hybrid (cen- 
ter), and S. missurica (right). 


more natural to retain species of Besseyva in Synthyris—as was done up 


to the time of Rydberg (1903). 


Department of Botany 
University of Washington, Seattle 
Federal Emergency Science School 

Lagos, Nigeria 


LITERATURE CITED 


Hepc in, F. L. 1959. A survey of the genus Synthyris. Master’s degree thesis (unpub- 
lished). Univ. of Washington, Seattle. 

McMILtran, C. 1949. In Documented chromosome numbers of plants. Madrono 10:95. 

PENNELL, F. W. 1933. A revision of Synthyris and Besseya. Proc. Acad. Phila. 
85:77-106. 

RypBerc, P. A. 1903. Some generic segregations. Bull. Torrey Club 30:271-281. 


NOTES AND NEWS 


Howarp E. McMinn.—Professor McMinn passed away at his home in Oakland, 
California, on August 25, 1963, after a lengthy illness. From 1918 to 1957 Mr. McMinn 
was professor of botany at Mills College. During his long career he published several 
books, the best known being “An Illustrated Manual of Pacific Coast Trees” and “An 
Illustrated Manual of California Shrubs,’ and comprehensive monographs of the 
genera Ceanothus and Diplacus. A detailed biographical account will be published in 
a forthcoming number of Madrono. 


116 MADRONO [Vol. 17 
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 
(See Madronio 9:257-258. 1948) 
SPECIES NUMBER COUNTED BY COLLECTION LocaLiTy 
Arcytophyllum 2n = 36 W.H. Lewis Monroe 40 6 km n of Andohu- 
thymifolium (R. & P). K K aylas, Apurimac, 
Standl. Peru 
2n ==36 W. H. Lewis Monroe 41 6 km n of Andohu- 
K SMU aylas, Apurimac, 
Peru 
2n-== $6 W.H. Lewis Monroe 21 20 km s of Huncayo, 
K K, SMU Junin, Peru 
2ni==36 W.H. Lewis Cerrate 3259 Ancash, Peru 
K K 
2n==.36 W. H. Lewis Monroe 18 40 km n of Quito, 
K K, SMU, TEX Ecuador 
Baptisa n= 9 B. L. Turner Turner 4694 Lowndes Co., 
calycosa Canby TEX TEX Georgia 
perfoliata nh ==9. B. L. Turner Turner 4689 Columbia Co., 
(Ee) RR: Br: TEX TEX Georgia 
sim plicifolia n= 9 B. L. Turner Turner 4659 Franklin Co., 
Croom TEX TEX Florida 
Brachycome n= 18 D.C.D.DeJong Whibley 775 55 m see of Ooldea, 
ciliaris (Labill.) MSC MSC South Australia, 
Less. var. ciliaris Australia 
iberidifolia He D.C.D.DeJong De Jong 1254 Cultivated, Mich- 
Benth. MSC MSC igan State Univer- | 
sity, East Lansing 
n='9 D.C.D.DeJong De Jong 1255 Cultivated, Mich- | 
MSC MSC igan State Univer- 
sity, East Lansing 
trachycar pa n= 27 D.C.D.DeJong Johnson 1688 Near Moura, Queens- | 
F. Muell. MSC MSC land, Australia 
Clerodendron Ha? L. Nevling, Jr. Howard & Nevling 5 mi w of Ponce, 
aculeatum A 15377, A Puerto Rico 
Griseb. 
Dalea n= 7 B. L. Turner Turner & Powell Hidalgo, Mexico 
citriodora Cav. TEX 1110, TEX 
schotti var. n= 10 B. L. Turner Turner 4776 Imperial Co., 
puberula (Parish) TEX TEX California 
Munz | 
Daphnopsis neo L. Nevling, Jr. Howard & Nevling Near Aibonito, | 
americana ssp. A 15399, AN Puerto Rico 
caribaea 
(Griseb.) Nevl. | 
philip piana n= 9 L. Nevling, Jr. Howard & Nevling Bosque Toro Negro, | 
Krug & Urban A 15428, A Puerto Rico 
Desmanthus n= 14 E. B. Smith Smith 214 Douglas Co., | 
illinoensis KANU KANU Kansas , 


(Michx.) MacM. 


1963 | 


SPECIES 


Garrya 
elliptica Dougl. 


Hedyosmum 
arborescens Sw. 6 


Leucothoe 
fontanesiana 
Sleumer 


Lythrum 
alatum Pursh 


Montia 
perfoliata 
(Willd.) Howell 
stbirica (L.) 
Howell 


Orobanche 
grayana var. 
nelsoni? Munz 


Parietaria 
pennsylvanica 
Muhl. 


Petalostemum 
feayi Chapm. 


| Petasites 
frigidus var. 
palmatus ( Ait.) 
Cronaq. 


Phytolacca 
decandra L. 


~ Polygonum 

scandens L. 

_ Ribes 

Sanguineum var. 
glutinosum 


(Benth.) Loud. 


laxiflorum Pursh 


CHROMOSOME NUMBERS 


NUMBER 
<9) See 11h; 
n= 8 
hn = [1 
neo 
=H —— 18 
2n = 18; 
A perm 1851 
*n — 12 
=H 12 
45) cet LP 
*2n = 2477 
n= 8 
Nie 
*2n = 30), 
n= 18 
n= 17 
2n = 8, 
2n = 8), 


COUNTED BY 


G.S. Van Horn 
HSC 


L. Ruidenberg 
GH 


L. Ruidenberg 
GH 


E. B. Smith 
KANU 


W. H. Lewis 
K 


D. E. Anderson 
HSC 


D. E. Anderson 
HSC 


W. H. Lewis 
W. H. Lewis 


W. H. Lewis 
K 


D. E. Anderson 


B. L. Turner 
TEX 


D. E. Anderson 
HSC 


E. B. Smith 
KANU 


E. B. Smith 
KANU 


D. E. Anderson 
HSC 


D. E. Anderson 
HSC 


LT] 


COLLECTION LOCALITY 
Van Horn 25 Humboldt Co., 
UC California 
Howard & Nevling Near La Mina, Lu- 
15327, A quillo National For- 
est, Puerto Rico 
Green 20-63 Cultivated, 
AAH, Arnold Arboretum, 
no. 71345K Weston, Mass. 
Smith 180 Douglas Co., 
KANU Kansas 
Lewis 5851 Leeds (Moortown), 
K, SMU Yorkshire, England 
Anderson 1930 Arcata, Humboldt 
UC Co., California 
Anderson 2239 Lake Earl, Del 
UC Norte Co., California 
Lewis 5811 Wimbledon, Surrey, 
K England 
Lewis 5810 Shipley, Baildon, 
K Yorkshire, England 
Lewts 5809 Leeds, Yorkshire, 
K England 
Anderson 2287 Big Lagoon Park, 
UC Humboldt Co., 
California 
Smith 203 Douglas Co., 
KANU Kansas 
Turner 4673 Lake Co., 
TEX Florida 
Anderson 2346 Arcata, Humboldt 
UC Co., California 
Smith 204 Douglas Co., 
KANU Kansas 
Smith 134 Douglas Co., 
KANU Kansas 
Anderson 1907 Humboldt Co., 
UC California 


Anderson 1969 
UC 


Humboldt Co., 
California 


* Prepared slide available. 


118 MADRONO [ Vol. 17 


ARTIFICIAL INTERGENERIC HYBRIDS OF HELIANTHUS 
AND VIGUIERA!’ 


CHARLES B. HEISER, JR. 


Helianthus and Viguiera are closely allied and their relationship has . . 
been discussed at some length by Blake (1918). He distinguished the two 
genera on the basis of the pappus, Helianthus having two deciduous awns 
rarely with intermediate squamellae and Viguiera having persistent awns 
and squamellae or lacking a pappus entirely. He also wrote of plants hav- 
ing a “habit” of Helianthus or ‘habit” of Viguiera but nowhere was this 
habit defined. Thus there is apparently no single character other than 
pappus which is fairly consistently different. Truly intermediate species 
are few in North America, but are represented by H. similis and H.niveus, 
according to Blake. In South America the situation is somewhat different — 
and it seems probable that the 19 South American species of Helianthus 
are more closely related to the Vigwiera of that region than they are to 
the North American members of Helianthus (Heiser, 1957). 

In his treatment of Vzguiera, Blake, realizing that presence or absence 
of pappus was hardly a generic character, eliminated the genus Gymno- 
lomnia which was defined by its lack of pappus, and disposed of its species 
by placing 25 of them into various sections of Viguiera, and assigning the 
remaining eight to other genera. Of particular concern here are those 
former species of Gymnolomnia which Blake placed in the section Helio- 
meris of Viguiera. This group which he considered ‘‘a compact group of 


closely related species, well distinguished by habit and involucre” is 


clearly set off from other sections of the genus and at the same time is 
clearly morphologically distinct from Helianthus. The section comprises 
Six species, five found in western North America, and the sixth, the curi- 
ously isolated V. porteri, known only from De Kalb, Rockdale, and 
Walton counties, Georgia. Chromosome numbers are known for four of 
the species, V. multiflora, V. longifolia, and V. ovalis, all n = 8; and 
V. porteri, n = 17. The latter number is the same as the basic number 
in Helianthus. . 

In 1957, 1958 and 1959 reciprocal pollinations of V. porteri were made 
with H. angustifolius, a species from the southeastern United States, 
which shares a number of morphological features with V. porter and has 
an overlapping blooming period. No hybrids were secured the first two 
years but three were secured in the third year with V. porteri as the female 
parent. In the following year hybrids were attempted with diploid repre- 
sentatives of the various “‘sections” of Helianthus. No seed set resulted 
from reciprocal pollinations of H. agrestis, H. annuus, H. carnosus, H. 


1 This study was carried out with aid of a grant from the National Science 
Foundation. Special thanks are due to my research assistant, Alvin Reeves and to 
those mentioned in Table 2 who supplied seeds. Figures 1-16 were drawn by Nancy 
Clark. The chromosome count for V. longifolia was made by David Kramer. 


1963 | HEISER: HELIANTHUS AND VIGUIERA 119 


laciniatus, and H. microcephalus with V. portert but a small number of 
seed was secured from crosses with H. canus, H. debilis and H. niveus 
with V. porteri again being the female parent. At the same time crosses 
of another species of Viguiera, V. dentata (n=17, sect. Chloracra) were 
attempted with H. atrorubens, H. canus, H. debilis, H. divaricatus, H. 
grosseserratus, H. laciniatus, H. microcephalus, H. neglectus, H. niveus, 
and H. occidentalis. None of these resulted in seed set, nor did crosses of 
V. portert with V. dentata and V. multiflora. Two hybrids, however, were 
secured from a cross of V. porteri and V. adenophylla (n=17, sect. 
Chloracra) with V. porteri as the female parent. 7 


All of the hybrids between V. porteri and the four species of Helianthus 
superficially appear similar to V. porteri but careful examination shows 
that they are more or less intermediate for many characters or more 
nearly approach the condition found in Helianthus. Some of the more 
conspicuous morphological features of the hybrids and their parents based 
on plants grown in the greenhouse are summarized in Table 1 in which 
the average value is given for the numerical characters. Attention should 
also be called to the differences observed in the trichomes found on the 
corolla of the various species and the hybrids. These are illustrated for 
V. portert (figs. 3-5), H. debilis (figs. 14-16) and their hybrid (figs. 8-11). 
Whether or not the extremely different type of bulbous trichome seen in 
V. portert will prove of value as a “generic character” is not yet known 
but deserves additional study. A cursory examination of specimens of 25 
different species of Helianthus revealed this type of trichome to be lack- 
ing. One of the most striking differences distinguishing V. porteri from 
the four species of Helianthus, as can be seen in Table 1, is the number 
of phyllaries. The involucre in V. porteri is best described as uniseriate 
whereas in Helianthus a second series is usually partially developed. It 
is worth noting that apparently all other sections of Viguiera are charac- 
terized by multiseriate involucres. The disk is low conic or slightly convex 
in V. porteri, nearly flat in the four species of Helianthus and interme- 
diate in the hybrids. Viguiera porteri has fewer disk flowers per head 
than do the species of Helianthus which, of course, can be accounted for 
in part by the smaller disk, but more significant is the fact that the flow- 
ers are much more tightly packed in the species of Helianthus. Differences 
in stem and leaf pubescence are also found, but except to note the con- 
spicuous trichomes at the leaf base in V. porteri (fig. 1) no attempt will 
be made to treat these in detail. Voucher herbarium specimens of the 
hybrids and their parents as well as those of the other species discussed 
here are deposited in the herbarium of Indiana University (table 2). 

The intergeneric hybrids were all extremely vigorous, produced an 
abundance of flowers, and were nearly or completely sterile. The pollen 
stainability with cotton blue of the seven hybrids of V. porteri « H. de- 
bilis ranged from 1 to 7% with a mean of 4%. The seven hybrids with 
H. canus gave a range of 0 to 13% with a mean of 4%. The V. porteri « 


[Vol. 17 


a 


MADRONO 


120 


911}ua 


SCA8S 
pol 


pol 
SV 


dIOU 10 Z 
MO][9A 


CXL 


GC 
cl 
9X1 


MO][9A 


cl 
oun f 


[etuussed 


: 


SNAQAU * H 


eR ielec) 


OI XOL 
pel 


MOT[9A 


Le 


(ia) 
MoO][I4 


1X8 


OI 
OT 
8XLI 


MOT[9A 
OT 
Ayn f 
UuMOUY 
JaA4 JOU 
¢ 


ux 


9} e[N11as 
0} 9} 8119S 


LLXS9 
ajdind 


ajdind 


8r 


vI 
LAL 


MO][9A-93UL1O 
8I 


Ain 
[etuuased 


s 
Si1Q2P HH 


SdIwd4AH Yay AGNv 


97e[N119s 


07 X06 
WII pal 


YVIM MOTTaA 


pol 
ae 


T-0 
ajdind 
JUIe] 
7X11 


12 
el 
LX9I 


9} PIPIUL19zUI 


cT 
A[nf 


jetuuased 


s 


px d 


9} e[NA19s 


pS X £6 
gjdind 


gidind 


SY 


02 
ST 


OT X02 


MO][9A-I93ULIO 


vT 
A[nf 
jenuue 


4 


SnuDd* H 


‘aSS SOHINVITAY 


31a 


7 X06 
pal 


pal 
6¢ 


0 
MO]T[AA 


ZALXOL 
sl 


els 
8XLZ 


9} CIPIWI9zUI 


el 
An f 
jenuue 


A) xd 


‘1WaLYOd VudINOTA AO NOSIYVdWO,) “T AIEVE 


aatqua 


IT X021 
ajdind 


ajdind 


gu 
e} 
9X91 


MO][9A-93ULIO 


vr 
"ydas 
[etuusied 


¢ 


snyofzy 
~SNBUD "HH 


aitqua 


cLXSel 
MO][9A 


pai 


ve 


fo) 
LXS1 


9} BIPIUIII}UT 


cl 
"ydas 
[etuusied 
2 
‘0 x ii 


ehialic) 


wWUI 8 XOS 


MO][9A 
MOT[9A 
WUL $7 


0 
MO][9A 


wu T XZ 


el 
WIUI g 


wu {X71 


MO][AA 
8 
"ydas 
jenuue 


s 
1494404 "A 


UISIVUL Jea] 


SIABI] JOM PUR “’T 
SBVUISI}S JO IO[OD 


SdqO] P[[O109 
YSIP FO 10[O) 
B][0109 YsIp 
jo yysuaT 
sajeos sndded jo ‘oN 


diy yeyp Jo 10]09 


sotiey[Ayd 
Jo YIPIM pue yyZ2ue] 
sateypAyd Jo ‘ON 
“WeIp ASIC 


YIPIM puke yysuaT 
SABI 10 IO[OD 
SABI JO JOqQUINN 
d}ep sUuIWOO[g 
uoljeing 


sjueid jo Jaquinnyy 


1963 | HEISER: HELIANTHUS AND VIGUIERA 124 
TABLE 2. SOURCE OF MATERIALS 
SPECIES LocaLiTy CoLLECTOR NUMBER’ 
Helianthus agrestis Poll. Polk Co., Florida Stoutamire 1745 
H. angustifolius L. Marion Co., Texas Martin 2009 
H. annuus L. St. Louis, Missouri Heiser 6001 
H. atrorubens L. Orange Co., N. Carolina Heiser H551 
H. canus (Britt.) Woot. Pima Co., Arizona Heiser 4707 
& Stand. 
H. carnosus Small Volusia Co., Florida Heiser 3184 
H. divaricatus L. Monroe Co., Indiana Heiser H553 
H. debilis Nutt. ssp. debilis Volusia Co., Florida Heiser 4665 
H. grosseserraus Mart. Greene Co., Indiana Heiser H502 
H. laciniatus Gray Zacatecas, Mexico Jackson 2493 (H535) 
H. microcephalus T.& G. Monroe Co., Indiana Heiser (6024A) 
H. neglectus Heiser Ward Co., Texas Clewell & H561 
Torres 
H. niveus (Benth.) Brandg. Ensenada, Baja Cali- Heiser 5849 
fornia, Mexico 
H. occidentalis Ridd. Lake Co., Indiana Dale (N1) 
Viguiera adenophylla Blake San Luis Potosi, Stoutamire 2813 
Mexico 
V.dentata (Cav.) Spreng Bernalillo Co., New Mexico Heiser V4693 
V. longifolia (Robins. & Cochise Co., Arizona Goodding 443-61 
Greenm.) Blake (62V22) 
V. multifiora (Nutt.) Blake Bernalillo Co., New Mexico Heiser V4994 
V. ovalis Blake Cochise Co., Arizona Goodding 416-5la 
(62V25) 
V. porter? (Gray) Blake DeKalb Co., Georgia Duncan (Vp) 


1 The number in parenthesis is the culture number used for some plants grown at 


Bloomington. 


H. niveus hybrids gave counts of 9, 19, and 29%, and those with H. an- 
gustifolius were 5, 8, and 20%. Three-fourths of the stainable grains in 
the last plant were quite large and had four instead of the usual three 
germ pores. That these grains are polyploid seems likely and is suggested 
by the occurrence of two celled ‘“‘tetrads” (fig. 24). 

Over 100 heads of field grown hybrids of V. porteri & H. angustifolius 
and V. portert * H. debilis which were examined failed to yield a single 
seed. Sister crosses of these hybrids as well as those of the other two com- 
binations also failed to produce seed. All possible backcrosses, except 
that of V. portert & H.canus to V. porteri which did not overlap in their 
blooming period, were attempted and all were barren with the exception 
that six apparently filled achenes were secured in the cross of V. porteri 
x H. debilis with H. debilis as the female parent. These seeds failed to 
germinate. 

Meiosis was examined in two or more of the plants in all of the hybrid 
combinations. Acetocarmine squash preparations were utilized and these 
were made permanent by means of the Venetian turpentine technique. 


[ Vol. 17 


~ 


MADRONO 


122 


[es 
ei ili eae a 
: Geka iy) 


SS 


No 
a 
7 WEE Saar 
&, ee ao 


1963 | HEISER: HELIANTHUS AND VIGUIERA 123 


In V. portert X H. angustifolius over 100 slides were made and only a 
small number of cells were found at diakinesis. These could not be ana- 
lyzed in detail but up to ten pairs of chromosomes were observed and 
chains were always present. At late diakinesis and metaphase I (figs. 22, 
23) from six to 26 univalents were observed. Micronuclei were observed 
at the tetrad stage (fig. 25). In V. portert K H.niveus in 25 cells studied 
at diakinesis the number of bivalents ranged from one to ten with a mean 
of five. Fewer irregularities were noted in the other two hybrid combina- 
tions. In V. portert « H. canus 89 cells were studied at diakinesis and 
the following configurations were found: nine cells with 17,, 11 with 
16y; 2;, 30 with 15;; and a chain of IV, nine with 15,, and a circle of IV, 
six with 15y; ly 11, one with 15;; 4;, 16 with 14, lyy 2;, three with 14, 
lyy, two with 14; ln 31, and two with 13y; ly 4;. At metaphase unival- 
ents were uncommon, although one to three were seen in some cells. Good 
stages of diakinesis were rare in V. porteri & H. debilis but four cells 
were seen with 17 bivalents (fig. 17). Univalents were also evident at 
this stage (figs. 18, 19) and from two to ten univalents were observed at 
metaphase (fig. 20). Bridges were observed at anaphase in all of the 
hybrids and were sometimes accompanied by a fragment (fig. 21). 

The two hybrids secured between V. porteri and V. adenophylla were 
less vigorous than the intergeneric hybrids. One died before flowering and 
the other plant produced only a small number of flowers and these were 
smaller than those of either parent. Pollen stainability was 5%. Very few 
cells could be found for study of meiosis. Eight of these, however, showed 
34 univalents, with 17 large chromosomes and 17 small chromosomes 
(fig. 27), and from a study of the chromosomes of the parents it appears 
that the 17 large chromosomes came from V. porteri (fig. 26). The pos- 
sibility that some chromosome pairing occurs in the hybrid cannot yet 
be ruled out. 


DISCUSSION 


The difference in pairing observed in the Helianthus-Viguiera hybrids 
is of interest in connection with the presumed relationships of the species 
of Helianthus involved. Two general groups of North American sunflow- 
ers may be recognized, the “‘annuals” and the ‘“‘perennials” (Heiser, Mar- 
tin, and Smith, 1962). Hybrids within either of these groups are generally 
readily secured and usually show some fertility whereas hybrids between 
the two groups are impossible to obtain or are highly sterile. Both H. 
canus and H. debilis are placed in the annual group, and hybrids of these 
two species show moderate to good chromosome pairing with V. porterz, 


Fics. 1-16. Leaves, disk corollas, and trichomes of V. porteri, V. porteri x H. 
debilis, and H. debilis; 1-5, V. porteri; 6-11, V. porteri < H. debilis; 12-16, H. debilis ; 
1, 6, 12, leaves, X 93; 2, 7, 13, disk corollas, « 5; 3, 8, 14, trichomes from apex of 
achenes; 4, 10, 11, 16, trichomes from tube of corolla; 5, 9, 15, trichomes from throat 
of corolla. All trichomes, x 18. 


124 MADRONO (Vole? 


8 - 


Fics. 17-21. Meiotic stages in V. portert * H. debilis, X 1200. For explanation 
see text. 


whereas the hybrids of Viguzera with H. angustifolius and H. niveus show 
little or no pairing. Helianthus angustifolius is a member of the peren- 
nial group. Helianthus niveus has hitherto been grouped with the annuals 
on overall morphological similarity but it is perhaps significant that thus 
far it has been impossible to secure crosses of H. niveus with the annuals. 

It may also be of significance that hybrids of V. porteri are more readily 
secured with Helianthus than with other species of Viguiera, and more- 
over, that the one intrageneric hybrid of Viguiera, V. porteri * adeno- 
phylla, is highly sterile and indicates considerable difference in the chrom- 
osomes of the two parents. Thus the evidence we have at present, although 


1963 | HEISER: HELIANTHUS AND VIGUIERA 12 


Ut 


24 25 


Fics. 22-25. Meiotis stages in V. porteri « H. angustifolius, * 1200; 22, 23, 
mostly univalents; 24, tetrad stage, showing two- and four-celled types; 25, pollen 
“tetrad” with micronuclei. 


admittedly very limited in view of the large size of Vigutera, perhaps 
suggest a closer relationship of V. porteri to Helianthus than to Viguzera. 

The securing of hybrids between Viguiera and Helianthus may bring 
up the question whether or not we are dealing with distinct genera. To 
some (Simpson, 1961) the production of a hybrid between two genera 
is sufficient basis for merging them. Hybrids between Euchlaena and Zea 
are known, and Reeves and Mangelsdorf (1942) have transferred the 
species of Euchlaena to Zea. Their basis for doing so, however, was on 
morphological grounds. The hybrids secured between Lycopersicon and 
Solanum by Rick (1951; 1960) are also of interest in this connection. 
Correll (1958) pointed out that the species of Solanum involved in these 
hybrids are morphological intermediates between the two genera, and the 
two have recently been united (Macbride, 1962) or reunited, for Linnaeus 
originally placed the tomato in Solanum. Kruckeberg (1962) pointed out 
that ‘“‘the ability to make successful hybridizations between species of 
related genera is not sufficient to cause the joining of those species into 
a single genus.”’ 


126 MADRONO [Vol. 17 


e.- a 

Y OOS as c (fe 
-_> 

= ade * ~~ Fe; e 


VJ rs 
26 Ye Din te 28 


Fics. 26-28. Camera lucida drawings of chromosomes of V. porteri, V. porteri 
x V. adenophylla, and V. adenophylla, < 1200; 26, V. porteri; 27, V. porteri < 
V. adenophylla; 28, V. adenophylla. 


In view of the weakness of the “generic” character it may indeed even- 
tually prove desirable to unite Helianthus and Viguiera, but for the sake 
of convenience, which may well enter in since there is no completely satis- 
factory definition of genera, there would appear to be good reason not to 
combine them. Moreover, there are certain other possibilities that need 
exploration—(1) should V. porteri be considered a monotypic genus? 
(2) should it alone be transferred to Helianthus? (3) should the section 
Heliomeris of Viguiera, either with or without V. porteri, be given generic 
status. 

On the basis of the present study it might be argued that V. porter 
should be placed in the genus Helianthus. It has a haploid chromosome 
number of 17, the same as the diploid sunflowers, whereas the three other 
members of the section Heliomeris are n=8, although it should be pointed 
out that chromosome numbers are not yet known for all members of this 
section. Its unique geographical position might more readily be explained 
if it were considered a Helianthus rather than a Viguiera. However, at 
present I still feel that on morphological grounds it is closer to the group 
with which Blake places it, so until more detailed morphological stuides 
are made there appears to be no justification for transferring it to Helian- 
thus or for erecting a monotypic genus for it. Certainly the placing of the 
section Heliomerts into the genus Viguiera needs reexamination. The pos- 
sibility of restoring these species to a genus of their own deserves con- 
sideration.” Thus these intergeneric hybrids are of interest in stimulating 
an examination of the existing classification, but by themselves are hardly 
adequate basis for making taxonomic changes. 


2 Since this was written it has been found that T. D. A. Cockerell (Notes on the 
flora of Boulder County, Colorado. Torreya 8:177-183. 1918) had already reached 
the same conclusions and had made formal transfers of these species, including 
V. porteri, to Heliomeris. 


1963 | HEISER: HELIANTHUS AND VIGUIERA WA 


SUMMARY 


Artificial hybrids of Viguiera porteri (n=17) with four species of Heli- 
anthus, H. angustifolius, H. canus, H. debilis, and H. niveus (all n=17) 
are described. Chromosome behavior was analyzed at meiosis and two 
of the hybrid combinations show a high number of univalents whereas 
the other two show fairly good pairing. All of the hybrids are sterile. 
Viguiera porteri has also been hybridized with V. adenophylia, producing 
a sterile hybrid showing 34 univalents at meiosis. It is pointed out that in 
some respects V. porteri appears to be more closely related to Helianthus 
than to other members of Viguiera. Problems in the classification of V. 
porteri are discussed, but no taxonomic transfers appear to be warranted 
until additional studies can be undertaken. 

Department of Botany 


Indiana University 
Bloomington, Indiana 


LITERATURE CITED 


BLAKE, S.F. 1918. A revision of the genus Viguiera. Contr. Gray Herb. 54:1-205. 


CorRELL, Donovan S. 1958. A new species and some nomenclatural changes in Sola- 
num, section Tuberarium. Madrono 14:232-236. 


HEISER, CHARLES B. 1957. A revision of the South American species of Helianthus. 
Brittonia 8:283-295. 


, W. C. Martin, and D. M. SmirH. 1962. Species crosses in Helianthus: 
I. Diploid species. Brittonia 14:137-147. 


KRUCKEBERG, A. R. 1962. Intergeneric hybrids in the Lychnideae (Carophyllaceae) 
Brittonia 14:311-321. 


MaAcsrIpDE, J. FRANCIS. 1962. Flora of Peru. Fieldiana Bot. 13 (pt. 5B) :158-163. 


REEVES, R.G. and P.C. MANGELSpoRF. 1942. A proposed taxonomic change in the 
tribe Maydeae (family Gramineae). Am. Jour. Bot. 29:815-817. 


Rick, CHARLES M. 1951. Hybrids between Lycopersicon esculentum Mill. and 
Solanum lycopersicoides Dun. Proc. Natl. Acad. U.S. 37:741-744. 


. 1960. Hybridization between Lycopersicon esculentum and Solanum penel- 
lii. Proc. Natl. Acad. U.S. 46:78-82. 


Stmpson, G.G. 1961. Principles of Animal Taxonomy. Columbia Univ. Press, New 
York. 


NOTES AND NEWS 


TRAVEL TO THE 10TH INTERNATIONAL BOTANICAL CONGRESS, EDINBURGH, AUGUST 
3-12, 1964.—The Botanical Society of America has appointed a committee to receive 
applications and recommend grants toward the cost of travel to the forthcoming 
congress. Professional botanists who are citizens of the United States are eligible to 
apply. They need not be members of the Botanical Society. The deadline for receipt 
of applications is February 15, 1964 and awards will be announced on or about 
April 1. Application forms may be obtained from Ralph E. Cleland, Chairman of the 
Travel Grants Committee, Department of Botany, Indiana University, Bloomington, 
Indiana. 


128 MADRONO [Vol. 17 


CHROMOSOME NUMBERS IN THE COMPOSITAE. 


VII. ADDITIONAL SPECIES FROM THE SOUTHWESTERN 
UNITED STATES AND MEXICO 


A. M. PowELl AND B. L. TURNER 


The present contribution is essentially a continuation of several papers, 
the latest of which (Turner et al., 1962) dealt with chromosome counts 
from species of southern Mexico; except for the 25 species from the 
United States, the counts reported in the present paper were obtained 
from a wide area of Mexico. 

Chromosome counts were made from pollen-mother-cell squashes as 
outlined by Turner and Johnston (1961). Voucher specimens (table 1) 
are deposited at The University of Texas Herbarium; these all were col- 
lected during the years 1961-62. The tribal and subtribal arrangements 
listed in Table 1 follow those of Hoffman (1894). The identifications are 
our Own, except for several which were kindly made by Arthur Cronquist 
and Kittie Parker. This study has been supported by National Science 
Foundation Grant 1216. 

EUPATORIEAE. Ageratum (x = 10). Chromosome counts for the 
species of this genus are consistent with the basic number obtained by 
other workers. Both diploid and tetraploid collections of several species 
of Ageratum have been reported. The three species listed in Table 1 are 
all diploid (n = 10); collections of these same species were reported by 
Turner et al. (1961a) as tetraploids (n = 20). 

Conoclinium greggu (n = 10). This genus is often included as a sec- 
tion of Eupatorium (Fernald, 1950; Hoffmann, 1894), but is treated as 
a separate genus by Small (1933) and others. The only other count for 
the genus has been that of Grant (1953) who reported C. coelestinum as 
diploid with 2n — 20. 

Eupatorium (x= 10, 17). E. prunellifolium (n= 52 univalents). 
Turner et al. (1961a) reported a meiotic count of 50 asynaptic chromo- 
somes for this species. Basic chromosome numbers of x = 10 and 17 have 
been reported for the section Eximbricata to which this species belongs 
(Grant, 1953). Thus, E. prunellaefolium is either a triploid on a base 
of x = 17, the two numbers 50 and 52 being aneuploid derivatives from 
2n — 51, or else it is on a base of x = 10, the present count being an 
aneuploid derivative from what might be an apomictic pentaploid. The 
authors prefer the former interpretation since at least one triploid species 
on a base of x = 17 is known for the section Eximbricata (Grant, 1953), 
while asynaptic pentaploids are unknown for the genus; however, Turner 
et al. (1961a), reported the chromosome number of EF. patzcuarenstis 
as n = 25 bivalents; the latter is apparently a pentaploid on a base of 
Xia 0 (er 2 = 50). 

Stevia (x = 11, 12, 17). Previous published counts for this multibasic 
genus have been on a base of x = 11 (Darlington and Wylie, 1956) and 


1963 | 


10 


o “oe aoe a? 
ae Py 


AND TURNER: COMPOSITAE 129 


e “. ° F ~2 
aoe 


F os Xe 
Se" @ as 


Ge 3 
@ PX o 
oe" ¢s wo 
e @ 
- © 
5 6 


fa. f gw 


n &@ So 


ae 
11 ad 12 Ow 


Fics. 1-12. Meiotic chromosomes of species of Compositae, & ca. 2000: 1, Eupa- 
torium prunellaefolium (n = 52 univalents) ; 2, Stevia elatior (n = 34 univalents) ; 
3, S. lucida (n = 12); 4, S. salicifolia (n = 12); 5, S. viscida (n = 11); 6, Aphano- 
stephus ramosus (n= 4-4 1 univalent) ; 7, Croptilon divaricatum (n = 4) ; 8, Grin- 
delia oxylepis var. eligulata (n= 6); 9, Gutierrezia glutinosa (n = 4); vO, Haplo- 
pappus gracilis (n= 2); 11, Isocoma veneta (n=6); 12, Milleria quinqueflora 


(n= 15). 


130 MADRONO [Vol. 17 


x = 17 (Turner ef al., 1961a). Chromosome numbers for all seven species 
listed in Table 1 are first reports. Three of the species had asynaptic 
chromosomes at meiosis I (n = 34 univalents) while one species, S. pur- 
purea, had meiotic configurations showing a variable number of bivalents 
and univalents. 

ASTEREAE. Croptilon divaricatum (n = 4). This count agrees with 
an earlier report for the species by Turner and Ellison (1960); however 
Jackson (1959) reported a count of 2n = 10 from a population in Kansas. 
Shinners (1950) recognizes several varieties for the species; both of the 
n = 4 counts were more from the typical variety (var. divaricatum) ; 
Jackson’s count was apparently obtained from the variety hookerianum 
as treated by Shinners. 

Chromosome numbers for A phanostephus (n = 4), Baccharis (n = 9), 
Conyza (n= 9, 27), Erigeron (n= 9, 18), Grindelia (n = 6), Gutier- 
rezia (n— 4), Haplopappus (n= 2), Heterotheca (n= 9), Isocoma 
(n= 6) and Townsendia (n= 9) are consistent with previous basic 
chromosome counts established for these genera (Darlington and Wylie, 
1956; Raven et al., 1960; Cave, 1960). 

INULEAE. The meiotic counts of Gnaphalium (n = 7, 14) listed in 
Table 1 are consistent with the basic number (x = 7) established for the 
genus by other workers (Darlington and Wylie, 1956). 

HELIANTHEAE. MIL erinae. This subtribe contains 16 genera; 
the count for Milleria quinquefolia (n = 15) is the first genus in the 
group to be counted. 

ZINNINEA. Chromosome counts for the species of Sanvitalia and Zinnia 
are consistent with counts previously reported for members of these taxa. 
Zinnia angustifolia is apparently dibasic, consisting of populations with 
n— 12 andn= 11 (Turner e¢ al., 1961a, 1962; present paper). 

VERBESININAE. Geraea canescens (n = 18). This is the first chromo- 
some count for the genus. Chromosome counts for the four species of 
Simsia (x = 17), listed in Table 1 were all diploid with n = 17: the genus 
is apparently unibasic, in that at least 7 of its approximately 25 species 
are diploid with this n number (Turner e¢ al., 1961b). Gray (1849) in- 
cluded Geraea in Simsia but Blake (1913) recognized the genera as 
distinct. 

Chromosome counts for Melanthera (x = 15) and Pervmenium (x = 
15) are consistent with previous basic numbers established for these 
genera (Turner et al., 1961b, 1962). Viguiera deltoidea (n = 18) was 
also reported as n = 18 by Heiser (1960). Chromosome counts for the 
two species of Zaluzania agree with the established base numbers of 
x = 16 and 18 (Turner, e# al., 1961a). 

CoREOPSIDINAE. Bidens pilosa (x = 12, 14). A single collection of 
Bidens pilosa var. radiata was found to be diploid with n — 14 bivalents. 
At least 8 collections for this widespread variety have been found to be 
diploid with n = 12 (including the seven counts listed in table 1). The 
meiotic figure with n= 14 appeared unequivocal (fig. 16). although 


———— 


1963 | POWELL AND TURNER: COMPOSITAE 131 


, Vv ~~ 
=, a2, Lage : hh c sty 
° ’ t 15 Ne 
13 14 


Fics. 13-24. Meiotic chromosomes of species of Compositae, X ca. 2000: 13, Zinnia 
cf. leucoglossa (n= 11); 14, Geraea canescens (n= 18); 15, Bidens cf. reptans 
(n= 11); 16, B. pilosa var. radiata (n= 14); 17, Chrysanthellum mexicanum 
(n= 8); 18, Baileya multiradiata (n = 16); 19, Hymenoxys acaulis (n = 14); 20, 
Laphamia lindheimeri (n= 18); 21, Psilostrophe villosa (n= 17); 22, Bartlettia 
scaposa (n= 11); 23, Pseudoclappia arenaria (n=18+1); 24, Senecio filifolius 
(ni —20). 


132 MADRONO [Vol. 17 


some of the cells observed might have been interpretable as n = 13. 
Approximately 22 species of Bzdens have been studied chromosomally; 
they form an aneuploid series with numbers of n = 10, 11, 12 and 14. 

Chrysanthellum mexicanum (n = 8). This is the first chromosome 
count for this genus and is the lowest basic number reported for the sub- 
tribe Coreopsidinae; the only other taxon with a base of x = 8 is the 
multibasic species Thelesperma filifolium with numbers of n = 8, 9, and 
11 (Melchert, personal communication). Chromosome counts of Coreop- 
sis (n = 12), and Cosmos (n = 12) are consistent with basic numbers 
previously reported (Darlington and Wylie, 1956). A single collection 
of Cosmos bipinnatus was found to be diploid with n = 11, this being a 
new basic number for the genus. Crowe (1954) had reported a count 
of 2n = 24 for this species. 


GALINSOGINAE. Chromosome counts listed in Table 1 for the genus 
Bebbia (n = 9) are first reports. 

Calea (x = ca. 16, 18). Turner e¢ al. (1961b; 1962) reported counts 
of x =ca. 16 and x = 18 for this genus. The count for C. urticifolia 
(n = 19) adds yet another chromosome number to the genus. Calea 
scabra (n = 32), is apparently a polyploid on a base of x = 16. 

Sabazia (x = 4). First reports for three species of Sabazia (x = 4) 
are listed in Table 1. Two of these taxa, S. cf. liebmannii (n = 24) and 
S. cf. michoacana (n = 24) are apparently dodecaploids, and one taxon, 
Sabazia sp. (n = 8), is a tetraploid. The only previous reports for the 
genus are n = 4 for S. humilis and n = 18 for Sabazia sp. (Turner and 
Johnston, 1961; Turner e¢ al., 1962). 


HELENIEAE. Hymenoxys odorata (n = 11, 15). This species appar- 
ently consists of races with at least 2 chromosome numbers (table 1). 
Turner (1963) reported several collections from Northeastern Mexico 
as n= 15; collections from the United States however have all been 
diploid with n= 11 (Speese and Baldwin, 1952; Raven and Kyhos, 
1961; present paper). The morphological differences, if any, which mark 
the races do not appear significant. 


Hymenoxys acaulis (n = 14). Previous reports for the genus Hymen- 
oxys have all been on a base of x = 11 or 15. The present count was 
unequivocal and was observed in numerous cells. While a newly discov- 
ered chromosome number for a taxon that was believed to be constant 
for such a character might prove vexing, it none-the-less can be expected 
to occur from time to time in various taxa when a more extensive sampling 
is undertaken. Several such examples in the experience of the junior 
author could be cited and such reports are becoming increasingly common 
in the literature. 

Psilostrophe villosa (n = 17). Previous species counts for this genus 
have all been diploid with n = 16. The present count appeared unequi- 
vocal (fig. 21) and was obtained from a number of cells from different 
bud material taken from the same population. 


1963] POWELL AND TURNER: COMPOSITAE 133 


Laphamia lindheimeri (n = 18). Raven and Kyhos have reported an- 
other species (L. congesta) as diploid with n = 32. 

Chromosome numbers for Baileya (n= 16), Bahia (n = 8, 12, and 
11), Chaenactis (n= 5, 8), Chrysactinia (n= 15), Flaveria (n= 18), 
Gaillardia (n= 17), Dyssodia (n = 13, ca. 26), Nicolletia (n = 10), 
Pectis (n= 12, ca. 24), Porophyllum (n = 12), Sartwellia (n= 18), 
and Tagetes (n = 24) are consistent with basic numbers previously re- 
ported (Ellison, 1961; Raven and Kyhos, 1961; Turner and Johnston, 
1961; Johnston and Turner, 1962; Turner e¢ al., 1961a; 1961b). 


SENECIONEAE. Bartlettia (n = 11). This count is a first report for 
the genus. 

Pseudoclappia arenaria (n = 18 + 1); Clappia suaedifolia (n = 16). 
Rydberg (1923) placed Pseudoclappia in the tribe Senecioneae, subtribe 
Senecioninae. He considered its relationship with the genus Clappia but 
noting, in particular, the resin glands which are found on the involucre 
and upper leaves of this latter taxon, he placed the two genera in separate 
tribes relegating Clappia to the tribe Tageteae. 

Rydberg retained Clappia in the subtribe Jauminae for his treatment 
in the North American Flora. In his 1923 discussion, he gave as a 
reason for this earlier disposition the following: “. . . I was not so well 
acquainted with the variations displayed in the subtribe Tagetinae, 
and let it remain in the position given by Gray and Hoffman [svc.], 
though I felt that it was out of place.” Raven and Kyhos (1961) also 
remarked on the anomalous position of Clappia if placed in the subtribe 
Jauminae, although they suggested no alternative position. 


We are inclined to agree with Rydberg’s later disposition of Clappia 
as well as his observations concerning the position of Pseudoclappia. We 
would go further however and suggest that there is a relatively close 
relationship between Clappia and Pseudoclappia and that while they 
might belong to different tribes, they probably do tie the Tagetinae with 
the Senecioneae in a phyletic sense and it might be both more expedient 
and more natural, to include the tribe Tageteae as a subtribe within the 
Senecioneae. 

In this connection, it seems worthwhile to consider the small genus 
Varilla, which is also found in northeastern Mexico and adjacent Texas. 
Its position in the Compositae is also anomalous, being placed in the 
Heliantheae by Gray and others, probably because of its chaffy receptacle. 
Clappia, Pseudoclappia and Varilla are alike in their succulent habits, 
distribution, and ecology (all are characteristic of saline or brackish flats 
of northeastern Mexico and Texas). While the morphological similarities, 
particularly in the case of Varilla may be more superficial than real, we 
believe that serious consideration should be given the possibility of a 
closer relationship among themselves than has heretofore been assumed. 


MUTISIEAE. Trixis (x = 27). The chromosome number for Trixis 
californica (n = 27) agrees with previous reports (Turner et al., 1962). 


134 


MADRONO 


[Vol. 17 


CICHORIEAE. Chromosome counts for Hieracium (n= 9), Mala- 
cothrix (n = 7), and Pinaropappus (n = 18) agree with the basic num- 
bers established for these genera by other workers (Darlington and 
Wylie, 1956; Stebbins e# al., 1953). 


TABLE 1. SPECIES OF COMPOSITAE EXAMINED FOR CHROMOSOME NUMBER 


SPECIES 


Ageratum albidum (DC.) 
Hemsl. 

A. cf. corymbosum Zucc. 

A. salicifolum Hemsl. 

Conoclinum greggiit Gray 


Eupatorium prunellifolium 
H.B.K. 
Stevia elatior H. B. K. 


S. lucida Lag. 

S. purpurea Pers. 
S.rhombifolia H. B. K. 
S. salicifolia Cav. 


S. serrata Cav. 


S. viscida H.B.K. 


Aphanostephus cf. pachy- 
rrhizus Shinn. 
A. ramosus (DC.) Gray 


Baccharis sordescens DC. 


Conyza bonariensis (L.) 
Cronq. 

C. coronopifolia H. B. K. 

C. gnaphaloides H.B.K. 

Cro ptilon divaricatum 
(Nutt.) Raf. 

Erigeron affinis DC. 


E. delphinifolius var. oreo- 
philus (Greenm.) Cronq. 


1 Collection numbers are those of Powell and Edmundson unless otherwise 


indicated. 


LOCATION AND VOUCHER 


n NUMBER 


EUPATORIEAE 


59 ms of Tehuacan, Oaxaca. 6661 


45 mi w of Sahuayo, Jalisco. 841 

6 mi w of Guadalajara, Jalisco. 866 

60 mi e of San Pedro de las Colonias, 
Coahuila. 522 

27 mis of Mexico City, Mexico. 745 


Along route 190, 40 mi e of junction of 
125-190, Oaxaca. 669 

53 mis of Tehuacan, Oaxaca. 665 

15 min of Cuernavaca, Morelos. 375 

Llano Grande, Mexico. Rock M—352 

4 mie of Majalca, Chihuahua. Ellison 
148 

Along route 55, 3 mi from junction with 
route 57. 583 

10 mi w of Toluca, Mexico. 794 

10 mi e of Puebla, Puebla. 623 

Huehuetenango, Guatemala. King 3412 

6 mi w of Guadalajara, Jalisco. 868 


ASTEREAE 
8 mi ne of Tepeaca, Puebla. 633 


2 mi w of Hidalgo, Michoacan. 815 


5 min of Chilpancingo, Guerrero. John- 
ston 6002 
54 min of Acapulco, Guerrero. 782 


12 min of Cuernavaca, Morelos. 720 

10 mie of Puebla, Puebla. 628 

Eldorado, Union Co., Arkansas. Turner 
4854 

9 mis of Victoria, Tamaulipas. King 4537 

14 mi sw of Chihuahua City, Chihuahua. 
978 


10 


10 
10 
10 


52 (fig. 1) 
univalents) 

17 (fig. 2) 

34 univalents) 

12 (fig. 3) 

17 (2-8,,, 30-18,) 
17 (34 univalents) 
12 (fig. 4) 


17 (34 univalents) 


17 (34 univalents) 
17 (34 univalents) 
17 (34 univalents) 
11 (fig. 5) 


4 


4+ 1 univalent 
(fig. 6) 
9 


ca. 27 
9 
ca.9 


4 (fig. 7) 


9 
y) 


1963] 


SPECIES 


POWELL AND TURNER: COMPOSITAE 


LOCATION AND VOUCHER 


1335 


n NUMBER 


Erigeron sp. 
E. sp. 
Grindelia inuloides Willd. 


G. oxyle pis var. eligulata 
Steyerm. 
G. robinsonii Steyerm. 


Gutierrezia glutinosa 
(Schauer) Sch. Bip. 

Haplopappus gracilis 
(Nutt.) Gray 


Heterotheca inuloides Cass. 


Isocoma heterophylla 
(Gray) Greene 

I. veneta (H.B.K.) 
Greene 


Townsendia mexicana Gray 


Gnaphalium cf. lepto- 
phyllum DC. 


G. sp. 


Milleria quinqueflora L. 


Parthenium fruticosum 
Less. 


Sanvitalia ocymoides DC. 
S.cf. procumbens L. 


S. sp. 


Zinnia angustifolia 
HoBoK. 

Z. haageana Regel 

Z. cf. leucoglossa Blake 

Z. tenella Robinson 


14 mi sw of Chihuahua City, Chihuahua. 
797 

22 mie of San Luis Potosi, San Luis Po- 
tosi. 550 

29 mi s of toll gate, n of Mexico City, 
Mexico. 585 

12 mis of Saltillo, Cohuila. 527 


3 mis of San Luis Potosi, San Luis Po- 
tosi. 553 
Cuahutemoc, Chihuahua. 986 


53 mis of Parral, Chihuahua. 963 


14 mi sw of Chihuahua City, Chihuahua. 
980 

10 mi s of Apam, Hidalgo. 608 

2 mi nw of Pecos, Reeves Co., Texas. 
Ellison 174 

Along route 85, ca. 20 min of route 30, 
Mexico. 596 

3 mis of San Luis Potosi, San Luis Po- 
tosi. 554 

12 mis of Saltillo, Coahuila. 530 


INULEAE 
20 mi n of route 30, Mexico. 601 


12 mi n of Cuernavaca, Morelos. 721 
10 mi ne of Oaxaca, Oaxaca. 677 


HELIANTHEAE. MILLERINAE 


45 mi w of Sahuayo, Jalisco. 845 


MELAMPODINAE 


3 mi e of the San Fernando-Santander 
highway on road to Loreto, Tamauli- 
pas. Johnston 5592 


ZINNINAE 


12 mis of Saltillo, Coahuila. 525 

4 mis of Villa Hidalgo, San Luis Potsoi. 
547 

Along route 190, 40 mi e of junction of 
125-190, Oaxaca. 671 

20 mi s of Guadalajara, Jalisco. 861 

5 mi w of Morelia, Michoacan. 826 

30 mi w of Tequila, Jalisco. 893 

30 mi w of Tequila, Jalisco. 892 


18 


2 (fig. 10) 


9 
6 


6 (fig. 11) 
6 


9 


15 (fig. 12) 


ca. 16 


11 


i set 

11 (fig. 13) 

11+ 1lor2 
fragments 


136 


SPECIES 


Geraea canescens T.& G. 


Melanthera angustifolia 
A. Rich. 
Perymenium flexuosum 
Greenm. 
P.hypoleucum Blake 
Simsia foetida (Cav.) Blake 


S. lagasciformis DC. 
S. cf. megace phala Sch.-Bip. 
S. triloba Blake 
Viguiera deltoidea var. 
parishii (Greene) 
Vasey & Rose 
Zaluzania globosa Sch.-Bip. 


Z. montagnifolia Sch.-Bip. 
Z.robinsonii Sharp 


Bidens cf. aurea (Ait.) Sherff 
B.cf.reptans (L.) G. Don. 


B. pilosa var. bimucronata 
(Turez.) Schulz 


B. pilosa var. minor (Blume) 
Sherff 

B. pilosa L. var. pilosa 

B. pilosa var. radiata 
Sch.-Bip. 


MADRONO 


LOCATION AND VOUCHER 


VERBESININAE 


Dateland, Yuma Co., Arizona. Turner 
4754 

16 mi e of Brawley, Imperial Co., Cali- 
fornia. Turner 4762 

Along route 144, 21 mi sw of junction of 
140-144, Vera Cruz. 642 

12 min of Cuernavaca, Morelos. 731 


53 mis of Tehuacan, Oaxaca. 659 

3 mi s of Huehuetenango, Guatemala. 
King, 3415 

Along route 85, 20 mi n of route 30, 
Mexico. 599 

Along route 85, 20 mi n of route 30, 
Mexico. 603 

22 mi ne of Tepeaca, Puebla. 640 

Cuahutemoc, Chihuahua. 998 

14 mi w of Zamora, Michoacan. 840 

4 mie of Jacumba, San Diego Co., Cali- 
fornia. Turner 4773 


Along route 85, 20 mi n of route 30, 
Mexico. 597 

Tehuacan, Puebla. King 2645 

31 mis of Galeana, Nuevo Leon. John- 
ston 5857 


COREOPSIDINAE 


2 mi w of Hidalgo, Michoacan. 819 
22 mi ne of Tepeaca, Puebla. 638 
22 mi ne of Tepeaca, Puebla 641 
Cuahutemoc, Chihuahua. 992 


10 mi w of Toluca, Mexico. 793 

38 mis of San Luis Potosi, San Luis Po- 
tosi. 572 

17 mis of Huatusco, Vera Cruz. 643 

14 mi sw of Chihuahua, Chihuahua. 977 


54 min of Acapulco, Guerrero. 778 
65 min of Acapulco, Guerrero. 787 


Along route 57, 29 mis of toll gate, n of 
Mexico City, Mexico. 586 

24 min of Cuernavaca, Mexico. 738 

10 mi w of Toluca, Mexico. 792 

8 mi e of Morelia, Michoacan. Ellison 
101 

8 mi ne of Tepeaca, Puebla. 635 

22 mi ne of Tepeaca, Puebla. 637 

17 mi s of Huatusco, Vera Cruz. 644 


[Vol. 17 


n NUMBER 


18 
18 (fig. 14) 
15 


ca. 30 


ca.17 


ca. 16 


12 Ghig. 15) 


12 


12 
12 
14 (fig. 16) 


12 
12 
12 


1963 | 


SPECIES 


POWELL AND TURNER: COMPOSITAE 


LOCATION AND VOUCHER 


Chrysanthellum mexicanum 
Greenm. 

Coreopsis cvclocarpa Blake 

Cosmos bipinnatus Cav. 

C. parviflorus (Jacq.) Pers. 


Bebbia juncea Greene 


Calea scabra (Lag.) Rob. 
C. urticifolia (Mill.) DC. 


Sabazia cf. iebmannii Klatt 
S.cf. michoacanna Rob. 


S. sp. 


Baileva mu!tiradiata Torr. 


Bahia glandulosa Greenm. 
B. schaffineri S. wats. 


B.xvlopoda Greenm. 
Chaenactis car phoclinia var. 
attenuata (Gray) Jones 

C. steviordes H.& A. 


Chrvsaciinia pinnata Wats. 


Dyssodia pinnata (Cav.) 
Rob. 


D. porophylloides Gray 


D. setifolia Lag. 


Flaveria ramosissima Klatt 
Gaillardia pinnatifida Torr. 


G. pinnatifida var. linearis 
(Rydb.) Bidd. 

Hymenoxvys acaulis (Pursh) 
Parker 


5 mi w of Morelia, Michoacan. 831 


6 mi w of Guadalajara, Jalisco. 863 
4 mi w of El Palmito, Sinaloa. 925 
Cuahutemoc, Chihuahua. 990 


GALINSOGINAE 


35 mi w of El Centro, Imperial Co., 
California. Turner 4778 

20 mi nw of Yuma, Mohave Co., Ari- 
zona. Turner 4784 

12 min of Cuernavaca, Morelos. 719 

10 mi se of Jalopa, Vera Cruz. Johnston 
4797 

10 mi ne of Oaxaca, Oaxaca. 674 

25 mi w of Oaxaca. Oaxaca. 706 

Along route 190, 40 mi e of junction of 
125-190, Oaxaca. 673 

4 mi w of El Palmito, Sinaloa. 920 


HELENIEAE 

6 mis of Van Horn, Culberson Co., 
Texas. Turner 4739 

14 mi w of Durango, Durango. 939 

4 mie of San Luis Potosi, San Luis Po- 
tosi. 548 

Along route 115, 5 mis of junction of 
routes 115-130, Hidalgo. 605 

12 mi e of Glamis, Imperial Co., Califor- 
nla. Turner 4765 

25 mi w of Deming, Luna Co., New 
Mexico. Turner 4745 

23 mis of Sabinas Hidalgo, Nuevo Leon. 
Johnston 5458 


Intersection of 136 and 57, Hidalgo. 
Ellison 92 

10 mi w of Toluca, Mexico. 803 

35 mi w of El Centro, Imperial Co., 
California. Turner 4780 

15 mis of Saltillo, Coahuila. 536 

28 mi ne of San Luis Potosi, San Luis 
Potosi. Ellison 62 

39 mis of Saltillo, Nuevo Leon. 542 

Intersection of 97 and 2009, Floyd Co., 
Texas. 179 


Cuahutemoc, Chihuahua. 989 


Palo Duro Canyon, Randall Co., Texas. 
Turner 4847 


PS) 


n NUMBER 


8 (fig. 17) 


16 (fig. 18) 


12 
8 


11 + 1 fragment 


se 


13 


17 


14 (fig. 19) 


138 


SPECIES 


Hymenoxys cf. linearifolia 
Hook. 
H. odorata DC. 


Nicolletia edwardsii Gray 
Laphamia lindheimeri Gray 


Pectis depressa Fern. 
P. latisquama Sch.-Bip. 
P. saturejoides (Mill.) 


Sch.-Bip. 

P. cf. texana Cory 

P. tenella DC. 

Porophyllum coloratum 
(H.B.K.) DC. 

P. ervendbergit Gray 

P. scoparium Gray 

Psilostrophe cooperi (Gray) 
Greene 

P. villosa Rydb. 


Sartwellia mexicana Gray 


Tagetes cf. elongata Willd. 


Bartlettia scaposa Gray 
Clap pia suaedifolia Gray 


Pseudoclap pia arenaria 


Rydb. 
Senecio filifolium Nutt. 


S. monoensis Greene 


Varilla texana Gray 


MADRONO 


LOCATION AND VOUCHER 


5 mi se of Valentine, Jeff Davis Cou 


Texas. Turner 4737 


5 mi se of Valentine, Jeff Davis Co.. 
Texas. Turner 4735 

25 mi ne of Yuma, Mohave Co., Ari- 
zona. Turner 4785 

5 mis of Quitaque, Briscoe Co., Texas. 
Melchert 177 

12 mi s of Saltillo, Coahuila. 528 

60 mi e of San Pedro de las Colonias, 
Coahuila. 523 

Edge Falls, 5 mis of Kendalia, Kendall 
Co., Texas. Johnston 6494 

16 mi w of Acapulco, Guerrero. 768 

10 mi e of Puebla, Puebla. King 3557 


19 mi w of Oaxaca, Oaxaca. 702 


8 mi nw of Royalty, Ward Co., Texas. 
Melchert 252 


7 min of La Gloria, Nuevo Leon. John- 
ston 4587A 


7 mie of Tierra Blanca, Oaxaca. 711 


13 mi w of Orizaba, Vera Cruz. John- 
ston 4779 


15 mi e of Ciudad del Maiz, San Luis 
Potosi. Johnston 5666A 


37 mis of Monclova, Coahuila. 571 


15 mi e of Tucson, Pima Co., Arizona. 
Turner 4750 


12 mi n of Matador, Motley Co., Texas. 
Melchert 183 


30 mis of Matehuala, San Luis Potosi. 
Ellison 59 


Ixtlan de Juarez, Oaxaca. King 3508 


SENECIONEAE 


23 mi e of Aldama, Chihuahua, 1019 


25 mi e of General Bravo, Nuevo Leon. 
Johnston 5324 

2 mi n of Pecos, Reeves Co., Texas. 
Melchert & Powell 256 

30 mi w of Chihuahua City, Chihuahua. 

1007 

4 mie of San Luis Potosi, San Luis Po- 
tosl. 552 

10 mi sw 20 of Congress, Yavapai Co.. 
Arizona. Turner 4789 

22 mi w of China, Nuevo Leon. John- 
ston 5012 


| Vol. 17 

n NUMBER 
ae 
Vl 
pat 
11 


15 
10 


18 (fig. 20) 


12 
ca. 24 
12 


11 (hig. 22) 
16 


1963 | POWELL AND TURNER: COMPOSITAE 139 


SPECIES LocaTION AND VOUCHER n NUMBER 
MUTISIEAE 
Trixis californica Kell. 35 mi w of El Centro, Imperial Co., 27 
California. Turner 4779 
CICHORIEAE 
Hieracium cf. crepi- 12 min of Cuernavaca, Morelos. 722 9 
dispermum Fries 
Malocothrix fendleri Gray 25 mi w of Deming, Luna Co., New Mex- i 
ico. Turner 4747 
Pinaropappus roseus Less. 8 mi ne of Tepeaca, Puebla. 634 18 
SUMMARY 


Chromosome counts are reported for 113 taxa (108 species in 56 gen- 
era) of Compositae from Mexico and the southwestern United States. 
Initial reports are included for 62 species, some of which belong to the 
following previously unreported genera: Bartlettia (n= 11), Bebbia 
(n = 9), Chrysanthellum (n = 8), Geraea (n = 18), Milleria (n = 15), 
Clappia (n = 16), Pseudoclappia (n = 18 + 1), and Varilla (n = 18). 

New basic numbers for the multibasic genera Stevia (x = 11, 12, 17), 
Bidens (x = 10, 11, 12, 14), Calea (x = 16, 18, 19), Cosmos (x = 11, 
12), Hvmenoxys (x = 11, 14, 15), and Psilostrophe (x = 16, 17) are 
reported. 

The phylogenetic relationship of the three halophytic genera Clappia, 
Pseudoclappia, and Varilla is suggested as being closer than previously 
indicated by taxonomic dispositions. 


The Plant Research Institute and 
Department of Botany 
The University of Texas, Austin 


LITERATURE CITED 
BLAKE, S.F. 1913. A redisposition of the species heretofore referred to Leptosyne. 
Proc. Am. Acad. 49:355-357. 


Cave, M.S. (Ed.) 1960. Index to plant chromosome numbers for 1960. University 
of North Carolina Press. Chapel Hill. 


Crowe, L.K. 1954. Incompatibility in Cosmos bipinnatus. Heredity 8:1-11. 


DariincTon, C.D. and A. P. Wy ie. 1956. Chromosome atlas of flowering plants. 
Macmillan Co., New York. 


Eviison, W.L. 1961. A systematic study of the genus Bahia (Compositae). 
Ph.D. dissertation, Univ. of Texas, Austin 


FERNALD, M.L. 1950. Gray’s manual of botany. American Book Co., New York. 


GRANT, W.F. 1953. A cytotaxonomic study in the genus Eupatorium. Am. Jour. 
Bot. 40:729-742. 


Gray, A. 1849. Plantae Fendlerianae Novi-Mexicanae. Mem. Am. Acad. 4:85. 


HEIsEer, C.B. 1960. Jn Documented chromosome numbers of plants. Madrono 
15:219-221. 


140 MADRONO [Vol. 17 


HorrMann, O. 1894. Compositae. Jn Engler, K. and A. Prantl. Die Natiirlichen 
Pflanzenfamilien IV(5) :87-391. 

Jackson, R.C. 1959. In Documented chromosome numbers of plants. Madrofio 
15:52. 

Jounston, M.C. and B. L. TurNER. 1962. Chromosome numbers of Dyssodia (Com- 
positae—-Tagetinae) and phyletic interpretations. Rhodora 64:2-15. 

Raven, P.H. and D.W. Kyuos. 1961. Chromosome numbers in Compositae. II. 
Helenieae. Am. Jour. Bot. 48:842-850. 

, O. T. Sotsric, D.W. Kyuos, and R. Snow. 1960. Chromosome numbers 
in Compositae. I. Astereae. Am. Jour. Bot. 47:124-132. 

Rypberc, P. A. 1923. A new genus of Senecioid composites. Jour. Wash. Acad. 
13:287-289. 

SHinnERS, L.H. 1950. Notes of Texas Compositae—IV. Field Lab. 18:156-159. 

SMALL, J. K. 1933. Manual of the southeastern flora. New York. 

SPEECE, B.M. and J.T. BAtpwin, Jr. 1952. Chromosomes of Hymenoxys. Amer. 
Jour. Bot. 39:685-688. 

StEepBins, G. L. Jr., J. A. JeENKins, and M. S. Watters. 1953. Chromosomes and 
phylogeny in the Compositae, tribe Chichorieae. Univ. Calif. Publ. Bot. 26: 
401-430. 

TurNER, B. L. 1963. Cytophyletic analysis of Hymenoxys odorata: a recapitulation. 
Madrono 17:77-79. 

, J. H. Beaman, and H. F. L. Rock. 1961a. Chromosome numbers in 
Compositae. V. Mexican and Guatemalan species. Rhodora 63:121-129. 

-———— and W. L. ELLison. 1960. Chromosome numbers in the Compositae. 

I. Texas species. Texas Acad. Sci. 12:146-151. 

, W. L. Extitson, and R. M. Kinc. 1961b. Chromosome numbers in the 
Compositae. IV. North American species, with phyletic interpretations. Am. 
Jour. Bot. 48:216-223. 

and M. C. JOHNSTON. 1961. Chromosome numbers in the Compositae. 
III. Certain Mexican species. Brittonia 13:64-69. 

, M. Pow_E tt, and R. M. Kinc. 1962. Chromosome numbers in the Com- 

positae. VI. Additional Mexican species. Rhodora 64:251-271. 


REVIEWS 


California Spring Wildflowers. 122 pp., 96 color photographs, 173 line drawings, 
2 maps. 1961. California Desert Wildflowers. 122 pp., 96 color photographs, 172 line 
drawings, 2 maps. 1962. California Mountain Wildflowers. 122 pp., 96 color photo- 
graphs, 180 line drawings, 2 maps. 1963. All by Puirtip A. Munz. University of Cali- 
fornia Press, Berkeley and Los Angeles. Paper, $2.95 each; cloth, $4.95 each. 

Death Valley Wildflowers. By Roxana S. Ferris. 141 pp., 144 species illustrated 
by line drawings. Death Valley Natural History Association, Death Valley, Calif. 
1962. $1.75. 

Ornamental Shrubs of California. By LEonID Enart. 214 pp., 181 figs. Ward Ritchie 
Press, Los Angeles. 1962. $5.95. 


Staff members of the several herbaria in California devote much time attempting 
to meet the public’s interest in California plants, both native and horticultural. 
Dominant among the queries received are those on plant identification. Not only 
does the layman want to know the names and characteristics of plants which interest 
him, frequently he wants to find these things out for himself. Yet there have been 
few nontechnical books available for the interested amateur to aid him in the iden- 
tification of plants which are native to California, and there are fewer still to help 
him identify plants which have been introduced for horticultural use. 

The republication in 1955, by the California Academy of Sciences, of Parsons’ 
“The Wildflowers of California,” first published in 1897, was a boon to those ama- 


1963 | REVIEWS 141 


teurs interested in native plants. Now there are available to help meet the need three 
handbooks by Philip A. Munz, author of the comprehensive and technical “A Cali- 
fornia Flora” published in 1959, and one by Roxana S. Ferris, author of the fourth 
and final volume, published in 1960, of Abrams’ monumental “Tllustrated Flora of 
the Pacific States.” 


Considering first Munz’ three handbooks, they are regional in character and 
together are intended to include the more commonly met plants of the state of Cali- 
fornia. The first, “California Spring Wildflowers,” includes spring-blooming plants 
found between the Sierra foothills and the sea, the second, “California Desert Wild- 
flowers,” includes plants of the desert areas east of the Sierra Nevada crest and south- 
ward through the Mojave and Colorado deserts and the Imperial Valley, while the 
third, “California Mountain Wildflowers,” covers the area above the Yellow Pine belt 
of the Sierra Nevada and other mountainous areas of California. 


Almost every plant discussed in these handbooks is illustrated by line drawing 
or by color photograph. Although the line drawings in the spring and desert wild- 
flower books are the work of a number of artists, they are all of high calibre. Those 
of the mountain wildflower book are mainly from the skilled pen of the very capable 
botanical artists, Jeanne R. Janish. The color photographs vary considerably, from 
many of outstanding quality to a few, unfortunately, which lack true color rendition 
or sharpness of image. However, the great majority of both types of illustration 
catch the ‘“‘feel” of the plant depicted, and thus fulfill their intended use of being one 
of the principal means of identfiying the plant in question. The texts include neither 
keys nor detailed descriptions, but do describe briefly for each plant the more obvious 
morphological characters as well as giving its geographical range and interesting facts 
about it. In all three books, text and illustrations for the plants illustrated by line 
drawings are grouped according to flower color, i.e., “flowers blue to violet,” etc. 
Text and illustrations for the plants with color photographs occur as a 32-page insert 
which is necessarily separated from the line-drawing section of each book by the 
demands of the printing and binding processes. 


The combination of flower color groupings for the plants treated in the line- 
drawing section along with the mechanical necessity of handling separately the plants 
illustrated with color photographs produces a situation in which closely related 
species may sometimes be found in three or more different sections of each book. 
Cross references are not always included in the spring wildflower book, and one must 
consult the index to find that there are, for instance, six different lupines in three 
sections of this book. Cross references appear more consistently in the desert and 
mountain wildflower books. 


In each of the three books there are two indices, a complete index serving both 
line-drawing and color photograph sections of the text, and a separate one for the 
plants illustrated only by color photographs. The latter organizes the plants so illus- 
trated into the same flower-color groupings that are used in the line-drawing section 
of each text. 

Wildflowers are usually considered as native annual or perennial herbs, and the 
titles of these handbooks imply such a limitation of subject matter. Together, how- 
ever, the books contain a number of common woody plants apt to pique the interest 
of the traveler, a few aliens, a few grasses, some rarities, and in addition the desert 
and mountain books include some ferns and one gymnosperm. Since there is such a 
vast number of California plants to choose from among wildflowers proper, it seems 
unfortunate, in some ways, that the number of these was reduced in favor of other 
plants. 

The problems are always manifold in finding a satisfactory way to make a non- 
technical botanical presentation of a large area within a very limited space, yet at the 
same time to give a meaningful text with “meat” sufficient for more than just a pretty 
book. Despite limitations these handbooks fill a long-felt need and contain much 
information of interest. 


142 MADRONO [Vol. 17 


Roxana Ferris’ handbook on “Death Valley Wildflowers” describes and illustrates 
approximately 150 plants of Death Valley National Monument. Although the nearly 
two million acres of the Monument are highly diversified in topography, climate, 
and soil, and support perhaps a thousand kinds of natural plant life, Mrs. Ferris has 
selected carefully those species which are particularly apt to pique the curiosity of 
the interested visitor. In addition to the more common herbaceous annuals and per- 
ennials, there are included several grasses, a number of highly characteristic shrubs, 
five trees which are outstanding in the vegetation at higher elevations of the Monu- 
ment, and three non-flowering plants (a stiped puffball and two ferns). 


The plants are arranged first by habitat-groups (following Munz’ classification 
of plant communities in “A California Flora”), second by growth-form (herb, shrub, 
tree), and third by flower-color, the clue-words appearing in large type at the bottom 
of each plant “story” or description. Identification is achieved by combined use of 
these clue-words, the description, and the illustration. Each species receives usually 
a full page divided between description and illustration. Almost no technical terms 
are used in the descriptions, and the latter are livened by references to Indian usage, 
animal interrelationships, historical data, etc. 


Not least of the attractions of this handbook are the splendid line drawings made 
for each species by Jeanne R. Janish, after the style she developed in her illustra- 
tions for three other popular handbooks of the Southwest, the “Flowers of the 
Southwest ...” series published by the Southwestern Monuments Association. Space 
is not stinted, and Mrs. Janish’s illustrations are often permitted one-half to three- 
fourths of the page. This latitude allows for a breadth of illustrative expression not 
possible under more cramped circumstances. Her tiny habit- and habitat-sketches 
and her humorous mannikin yardsticks express vividly, with a few strokes of the 
pen, that which words could express only cumbersomely, while her detailed drawings 
for each species are as botanically accurate as they are pleasing. 


For the thousands who visit Death Valley National Monument each season, as well 
as to the National Park Service itself, “Death Valley Wildflowers” should be a very 
welcome addition to the growing list of popular literature available on the natural 
history of the Southwest. 


Enari’s “Ornamental Shrubs of California” is intended for a wide audience, but 
it is reviewed here mainly from the standpoint of its use to the horticulturally-minded 
layman. Within its 214 pages it describes 277 ornamental shrubs, both native and 
introduced, which are grown in the gardens, parks, and nurseries of California. Al- 
though it uses botanical terms, keys, and technical descriptions, the botanical treat- 
ment is handled in such a manner that it should be of value not only to students and 
teachers of botany and research workers in taxonomy, but also to nurserymen, gar- 
deners, landscape architects, and in addition, homeowners and amateur botanists 
with horticultural interests. 


A preface is followed by a seven-page glossary of technical terms and six pages 
of outline drawings illustrating terms found in the glossary. Next occurs a modified 
dichotomous key (keying directly to species), preceded by a one-page “trial run” 
to illustrate its use. In the body of the text, families are arranged according to the 
usual Engler and Prantl system, and genera and species are arranged alphabetically 
within each family. There are no descriptions for families or genera, but these are 
unnecessary since the key takes the user directly to species. The species descriptions 
are drawn up carefully and logically and should be understandable to anyone who 
studies carefully the introductory material of the book. Outline leaf drawings accom- 
pany all of the species treated. Inclusion of floral patterns would have been helpful, 
especially in cases where leaf outlines do not illustrate distinctive features. Both 
common and scientific names are used, and there is a separate index for each. There 
should be much demand for this book.—HeELen K. SHarsMITH, University of Cali- 
fornia, Berkeley. 


1963 ] NOTES AND NEWS 143 


Meet Flora Mexicana. By M. WALTER PESMAN. 280 pp., fold-out vegetation map, 
illustrated. Dale S. King, Six Shooter Canyon, Globe, Arizona. 1962. $4.00 and $5.00 
paper-back ; $6.00 cloth. 


Mr. Pesman has written a delightful and original book about the roadside flora 
of Mexico. Meet Flora Mexicana is the first nontechnical work to be published on the 
area covered. In the words of the author, ‘We realized that many other tourists 
would want information about the very plants that attracted our attention, mainly 
those along the highways,” and so the present volume was born. The role of the 
author’s enthusiastic wife in the project is aptly expressed by an appreciative hus- 
band in the accolade “Blessings on a wife who is a front-seat driver!” On four trips 
to Mexico, Mrs. Elizabeth Pesman was chauffeur and an inspiration to her husband 
in various ways. 


Two-hundred and seventy original line drawings, all made on the spot by the 
author, illustrate the work. Included are some of the showiest and most unusual 
plants to be found as one travels by car along the principal roadways of the country, 
particularly in the winter months. Mexico is divided into 11 floral zones, i.e. desert, 
mesquite and grassland, thorn forest, tropical evergreen forest, and so forth. Once 
a person finds his geographical location on the colored fold-out map, an unknown 
plant is determined by comparison with the illustrations and the accompanying brief 
descriptions. Each plant is provided with a Spanish and an English vernacular name, 
as well as a botanical name. A bibliography of some references to the literature on 
Mexican botany ought to please many arm-chair users of the work, and a printed 
rule in inches and centimeters will please users who wish to use the book in the field. 


Although written for the layman, professional botanists and others ought to 
profit by this book. Some might criticize it for incompleteness, but this book is not 
intended to be a comprehensive technical flora. As the title suggests, readers are merely 
introduced to the Mexican flora through the eyes of the author; he hopes “it will 
do something to make you feel more at home among the many, many interesting 
plants” of Mexico. Perhaps the present work will encourage others to do better, since 
the author hopes to whet appetites and stimulate curiosities. Laurels to M. Walter 
Pesman for being the first to write a popular handbook about the plants of our 
good neighbor south of the border. The publishers are to be highly commended for 
the pleasing format, the very attractive cover, and the relatively modest cost of the 
book.—FrReEpeErick G. MEveEr, U.S. Department of Agriculture, Beltsville, Maryland. 


NOTES AND NEWS 


Notre oN DAMAGE TO THE HOOKER OAK. This well-known specimen of Quercus 
lobata Nee., located in Chico, California’s Bidwell Park, was severely damaged dur- 
ing a storm which battered the West Coast on October 12 and 13, 1962. Approxi- 
mately half of the tree, named by Annie E. K. Bidwell in honor of Sir Joseph Dalton 
Hooker, was broken off 25-40 feet above the ground on the southeast side, ruining 
its symmetry. The City of Chico is spending $5,000 this winter to shore up and prune 
the remaining branches and to prune the roots. Long-range plans are being prepared 
by local officials for restoration of the tree through continued seasonal pruning, but 
its future appearance is problematical. A plaque placed at the site in 1953 by the 
Native Daughters of the Golden West indicates the estimated age of the tree to be 
1,000 years, and the press has frequently reported its age as 1,100 years. However, 
ring counts of the largest branch broken off during the storm suggest that the age 
may be no more than 400 years. This coincides with a recent estimate by California 
Extension Forester Emeritus Woodbridge Metcalfe. Dimensions of the Hooker Oak 
given in 1953 are as follows: height, 96 ft.; circumference of tree 8 ft. from ground, 
29 ft.; spread of north and south branches, 153 ft.; circumference of outside 
branches, 481 ft.; lineal measurement—largest south branch, 111 ft.; diameter of 


144 MADRONO [Vol. 17 


trunk 8 ft. from the ground, 9 ft. [cf. H. A. Dutton’s note in Madrofio 1(5) :97-98. 
1922]. A few other less famous California specimens of Quercus lobata are as large 
or larger than the Hooker Oak. Mr. Metcalfe kindly made his file of data on such 
trees available for study, and the following information on three of them was culled 
by Dr. H. L. Mason: 

The Henley Oak on Wolf Hop Company Ranch, Round Valley, Mendocino 
County: DBH 8.65 feet, height 145 feet, first fork at 20 feet (1937 measurements). 

The Bower’s Ranch Oak near East Biggs south of Oroville. DBH 11’ 8” in August 
1955. 

Tree one-half mile south of Guinda School, Yolo County: DBH 103”, height 
100 feet—KiNGsLEY R. STERN, Department of Biological Sciences, Chico State Col- 
lege, Chico, California. 


EtuHet Bartey Hiccins.—Born in Vassalboro, Maine, August 10, 1866; died in 
San Diego, California, March 9, 1963. 

Mrs. Higgins, as a little girl, learned about the wildflowers which grew in the 
pastures of her grandfather’s farm, and continued that interest while a student at the 
Maine Wesleyan Seminary and Female College. In 1961 she returned as a special 
honored alumna for the 75th anniversary of her graduation at Kents Hill School, 
Kents Hill, Maine, which is the descendent of the college. Coming to California as 
a young woman, Mrs. Higgins opened a photographic studio in the village called 
Hollywood. Exploring the Hollywood hills and San Fernando Valley in her horse- 
drawn buggy, she collected wild flowers to photograph, building up a collection of 
300 plant portraits. It was in order to properly identify these that she took up the 
study of botany. Mrs. Higgins was a resident of San Diego after 1915, when she and 
her inventor husband opened a foundry and machine shop there. Botany remained 
an avocation for many years, but in 1931 Mrs. Higgins published a popular book, 
“Our Native Cacti,’ and in 1934 joined the staff of the Botany Department of the 
San Diego Natural History Museum. She was always interested in the work of early 
botanical collectors and had just completed a manuscript of the history of the botani- 
cal exploration of Baja California at the time of her death. She also had a strong 
interest in Mexico—its culture, history, and art, as well as its flora. In 1952 she 
realized an ambition of many years, and collected in the Cape region of Baja Cali- 
fornia, thus at 86 years of age starting an active program of travel in Mexico, which 
included visits to Mexico City, Oaxaca, and many other areas.—GeEorcE E. Linpsay, 
San Diego Natural History Museum. 


CALIFORNIA BOTANICAL Society.—Lest it be said that proper attention is not paid 
to the passage of time and the occasions thus generated, it should be noted that the 
year 1963 marks the 50th anniversary of the founding of the California Botanical 
Society. The Society through its activities and its journal, Madrofno, has contributed 
to the development of botany not only in California, but also outside of California. 
It is very difficult to single out individual and specific contributions, but one can 
point with pride to the papers of the first two editors of Madronfio, Willis Linn Jepson 
and Herbert L. Mason. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume I, 1916-1929 . . . . =. $5.00 
Volume II,,1930-1934,. . . 2. 2) 35:00 
Volume ITI, 1935-1936 aS eee Gre 0,0) 
Volume IV, 1937-1938. . . . . 5.00 
Volume V, 1939-1940 . . . . . 5.00 
Volume VI, 1941-1942 Re hoes coon ne? <) 00) 
Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 eres. eee: 0,0) 
Volume X, 1949-1950 . . . . . = 7.00 
Volume XI, 1951-1952 50 aes, en ae ORO 
Volume XII, 1953-1954 . . . . = 8.00 
Volume XIII, 1955-1956 . . . . = 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 ol Ret ae ot OLOO 
Volume XVI, 1961-1962 . . . . 12.00 


Single numbers: 


VolumesTandII . .; «2 ©) 37%. 50 
Volumes II through XIV... . 1.00 
Volumé 20) 005 Ah ais G2 ae 1.25 
Volume’ XVEU 0. “Aiea ee 1.50 


The subscription price of MApRONo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


VOLUME 17, NUMBER 5 JANUARY, 1964 
Contents 

PAGE 
PLANT COLLECTION IN NEPAL, D. D. Bhatt 145 


CYTOLOGICAL STUDIES IN THE GENUS Fiscus. III. 
CHROMOSOME NUMBERS IN SIXTY-T wo SPECIES, /ra 
J. Condit 153 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF 
THE GENUS MIMULUS (SCROPHULARIACEAE). VI. 
NEw NuMBERS IN M. GuTTATUS, M. TIGRINUS, AND 
M. cLaBratus, VM. M. Mia, B. B. Mukherjee, and 
R. K. Vickery, Jr. 156 


EXTENDED DORMANCY OF CHAPARRAL SHRUBS DURING 
SEVERE DroucHt, R. A. Harvey and H. A. Mooney 161 


CYTOLOGICAL OBSERVATIONS ON SOME GENERA OF THE 
AGAVACEAE, Marion S. Cave | 163 


REVIEWS: George Neville Jones, Flora of Illinois 
(Grady L. Webster); Samuel J. Pusateri, Flora of 
Our Sierran National Parks (Wallace R. Ernst) 170 


Notes AND News: A NEw LOCALITY FoR ASPLENIUM 
i 
VESPERTINIUM, Barbara Joe 172 


A WEST AMERICAN JOURNAL OF BOTANY 


HIBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


EDGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENsSoN, Pomona College, Claremont, California 
HERBERT F. COPELAND, Sacramento College, Sacramento, California 
JoHN F. Davipson, University of Nebraska, Lincoln 
WALLACE R. ErNsT, Smithsonian Institution, Washington, D.C. 
Mivprep E. Martutas, University of California, Los Angeles 
ROBERT ORNDUFF, University of California, Berkeley 
Marton OwnBEyY, Washington State University, Pullman 
REED C. RoLiins, Gray Harbarium, Harvard University 
IrA L. WiccINns, Stanford University, Stanford, California 


Editor—JoHN H. THoMAS 
Dudley Herbarium, Stanford University, Stanford, California — 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: G. Ledyard Stebbins, Department of Genetics, University of California, 
Davis. First Vice-President: Annetta Carter, Department of Botany, University of 
California, Berkeley. Second Vice-President: Carl W. Sharsmith, Department of 
Biological Sciences, San Jose State College. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding 
Secretary, Margaret Bergseng, Department of Botany, University of California, 
Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State 
College. 


1964 | BHATT: NEPAL 145 


PLANT COLLECTION IN NEPAL 
D. D. BHATT 


‘“«., And the wildest dreams of Kew are the 
facts of Kathmandu .. .”—Kipling. 


Nepal, lying in the lap of lofty Himalayas, is a naturalist’s paradise. 
The physical features, climate, and geographical position all seem to 
have contributed to make this country ideal for a plant collector. Although 
small in size, Nepal contains a variety of plant and animal life, from lush 
tropical to alpine types such as are seldom met in any one country. It is 
here that the Indian, Southeast Asiatic, and Sino-Japanese floristic regions 
of the world merge with each other, making it an interesting place to 
study plant distribution on the two sides of the Himalayan Mountains. 

In the late eighteenth and early nineteenth century, British botanists 
were actively engaged in the study of the flora of the Indian subcontinent. 
The pioneering work of Wallich, Hooker, and Roxburgh resulted in the 
production of such excellent floral studies as Tentamen Florae Napalensis 
(Wallich, 1820-24), Flora of British India (Hooker, 1875-97), and 
Flora Indica (Roxburgh, 1832). Due to its geographical location and par- 
ticularly to the isolationist policy of the Rana rulers, Nepal at first re- 
mained inaccessible to the British botanists and was a “land of mystery” 
in the eyes of western observers. However, after the 1814-1816 war with 
the British, the Nepalese rulers agreed to let some British botanists peep 
through a window (Kathmandu) into the country. How difficult it was 
for westerners to visit Nepal is obvious from the following remarks of 
Hooker (1855), “... Dr. Campbell gained the friendship of Jung Baha- 
door, the most remarkable proof of which is the acceding to his request, 
and granting me leave to visit the eastern parts of his dominions; no 
European that I am aware of, having been allowed, either before or since, 
to travel anywhere except to and from the plains of India and valley of 
Kathmandu, in which the capital city and British residency are situated.” 

The situation changed suddenly in 1950 when the autocratic rule of 
the Rana family came to an end as a result of popular uprising, and the 
country was opened to westerners for the first time. This opportunity was 
seized by mountaineers throughout the world and several expeditions 
were sent to Nepal to scale the renowned Himalayan peaks. The terrain 
of the country makes it very difficult for an individual collector or a small 
party to travel alone, and except for the British Museum parties the 
botanists visiting Nepal during the last decade have all gone as members 
of mountaineering parties. 


PHYSIOGRAPHY AND PLANT DISTRIBUTION 


Until a decade back very little was known about rainfall, temperature, 
or humidity in parts of the country outside Kathmandu, except that 


Maprono, Vol. 17, No. 6, pp. 145-172. January 31, 1964. 


146 MADRONO [Vol. 17 


Hooker (1855) did keep an accurate record of these features during his 
explorations in eastern Nepal. Some scattered accounts are now avail- 
able (Karan, 1960; Kawakita, 1956) which throw some light on these 
matters. 

Nepal is usually divided into five climatic regions, each characterized 
by its own topography, climate, and vegetation: subtropical, tropical, 
temperate, subalpine, and alpine. However, Hagen (1960) proposed 
seven natural divisions occurring in the following order: 1, the Terai; 
2, the Siwalik hills; 3, the Mahabharat Lekh (Lekh-mountain) ; 4, the 
Nepal midlands; 5, the Himalayas; 6, the inner Himalayas; 7, the 
Tibetan marginal mountains. The five climatic regions are grouped into 
the following three categories: 1, subtropical and tropical; 2, temperate; 
and 3, subalpine and alpine (fig. 1). 


Ne pt ed \ 


Fic 1. Climatic zones of Nepal: subtropical and tropical (stippled), temperate, 
and subalpine (hatched). 


SUBTROPICAL AND TROPICAL REGION. These regions consist of warm 
and humid lands and siwaliks (hills) known as Terai, Bhabbar, and 
Churiya respectively. The altitude varies from less than 500-5000 ft. 
Terai is part of the alluvial (cis-gangetic) plain of northern India and 
has a rainfall of 40-80 in (Karan, 1960). The rain occurs mostly during 
the summer months and the vegetation is more luxuriant in the eastern 
part of the country than in the western. Terai covers an area of 8000 sq mi 
and is 10-20 mi in width. Shorea robusta (sal or sakhu) is the most 
dominant species. Other timber species found in Terai and adjoining 
parts of Nepal are: Terminalia tomentosa, T. belerica, Anogeissus lati- 
folia, Dalbergia sisso, D. latifolia, Adina cordifolia, Bombax malabari- 
cum, Acacia catechu, and Cedrela toona. 

Bhabhar is a narrow strip of land between siwalik and Terai extend- 
ing over 3500 sq mi. The soil contains mainly gravel, boulders, and sand 
brought down by the streams from the siwalik and Mahabbarat ranges. 
New forests in this area contain Acacia catechu (khiar), Dalbergia sisso 
(sisso), and grasses, many being species of Saccharum. In old forests 
khiar and sisso are replaced by Bombax malabarica (semal), Albizzia 


1964 | BHATT: NEPAL 147 


lebbek (siris), Hymenodictyon excelsum (lati karma), and Anogeissus 
cordi (karma). 

The siwaliks known in Nepal as Churiya pahad (hills) extend from 
Afghanistan to Assam. The soil is gravelly, very porous, and contains 
little organic mtater. A number of duns (valleys) lie between the sil- 
waliks and the Mahabharat ranges. Forests are common in the valley 
floors and are composed largely of Acacia catechu, Lagerstroemia parvi- 
flora, Careya arborea, Adina cordifolia, and Michelia champaca (champ). 
The latter is found only in the eastern part of the country. 

TEMPERATE REGION. This area consists principally of the Mahabharat 
ranges and has a cool, humid climate. The elevation varies from 5000- 
10000 ft. Rainfall and temperature vary from east to west, and are gen- 
erally higher in the east than in the west. The “‘heart of the country,” as 
Hagen (1960, p. 69) calls these Nepal midlands, lies between Maha- 
bharat and the high Himalayas. Hagen divides it into nine natural 
regions, each including a wide valley drained by one of the major branches 
of the great rivers which form the main drainage system of Nepal. They 
run mainly from north to south in the following west to east sequence: 
Mahakali, Seti, Karnali, Bheri, Kali Gandaki, Buri Gandaki, Trisuli, 
Soonkosi, Arun, and Tamur. The vegetation consists of pure or mixed 
forests of pine and oak. At lower altitudes, Pinus wallichiana (Himalayan 
blue pine), P. roxburghii, Quercus lemellosa, QO. dilatata, Q. incana, Acer 
campbellu, Castanopsis indicus, and Pyrus pashia are the more important 
tree species. At middle altitudes, 6000-10000 ft, Quercus semecarpifolia, 
Q. incana, Juglans regia, Alnus nepalensis, Salix tetrasperma, Magnolia 
campbellu, Cedrus deodora, Picea morinda, P. smithiana, Abies webbiana, 
Pinus excelsa, P. chylla, Rhododendron arboreum, R. barbatum, Betula 
utilis, Thalictrum rotundifolium, Berberis aristata, B. nepalensis, Gaul- 
theria fragrantissima, Pieris ovalifolia, Clematis buchananiana, Spiraea 
vaccinifolia, Rubus paniculatus, Hypericum patulum, and Crategus cren- 
ulata are found. Epiphytic orchids, ferns, and angiosperms (Peperomia 
reflexa and Hoya longifolia) abound in these forests. Species of the tropi- 
cal and subtropical region interdigitate with those of the temperate region 
at 6000-7000 ft elevation. The upper limit of the temperate forest is 
12000 ft. 

SUBALPINE AND ALPINE REGIONS. These regions are composed mainly 
of the outer and inner Himalayan ranges. The tree line in the central 
Himalaya lies between 12000 and 13000 ft, extending to 15000 ft in the 
east. Snowline in the eastern Himalaya is at about 17000 ft (Swan, 1961). 
Rhododendrons have been reported from the subalpine and alpine regions 
of Nepal: R. cowanitanum (10500 ft), R. campanulatum (11000-13000 
ft), R. lepidatum, and R. anthopogon (12000-14000 ft). Other woody 
species at elevations of 10000-14000 ft are: Quercus semecarpifolia, Ju- 
niperus squamata, Tsuga dumosa, Larix griffith, Abies webbiana, Coton- 
easter sp., and Ephedra gerardiniana, the latter being the dominant 
species between 12500 and 16000 ft. Herbaceous plants found at these 


148 MADRONO [Vol. 17 


heights are various species of Pedicularis, Aconitum, Thalictrum, Poten- 
tilla, Gentiana, Meconopsis, Primula, Saussurea, Lonicera, Arenaria, and 
Saxifraga. The upper limit of vegetation in the Himalaya has been re- 
ported to be 20000 ft. Some of the species occurring between 15000 and 
20000 ft are: Rhododendron setosum, Juniperus squamata, Ephedra 
gerardiana, Gentiana vesusta, Primula sikkimensis, Meconopsis horridula, 
and several lichens. In western Nepal (Jumla-Humla) near Mohala 
Bhanjyang (Bhanjyang pass, 19500 ft), Lagotis glauca, Potentilla saun- 
dersiana var. caes pitosa, Pedicularis sp., and Arenara sp. have been found 
(Polunin, 1960). Recently Swan (1961) has reported Stellaria decumbens 
at 20130 ft, a new record in the Nepal Himalaya. Saussurea gossypiphora 
D. Don, which forms clubs of soft white wool from six inches to a foot 
high (first described as S. gossypina by Wallich in 1831), was brought 
back to Kathmandu by pilgrims from Gosainthan. It has been reported 
up to an altitude of 17000 feet (Hooker, 1855). 


PLANT COLLECTORS IN NEPAL 


The history of plant collection in Nepal can be divided roughly into 
two phases, from 1802 to 1950, and from 1950 to the present. During 
the first period, Nepal was a “forbidden land” to foreigners, and hence 
plant explorations were carried out in very small regions of the country. 
Sir Brian Hodgson, British Resident in Kathmandu from 1822 to 1843, 
was the first person to bring to the notice of western scientists the in- 
credible variety of plants and animals of Nepal. He published 127 papers 
which deal with birds, mammals, and reptiles of Nepal. As he was 
obliged to stay in Kathmandu by an order of the Nepalese Court, he 
employed Shikaris or professional hunters to collect for him. His primary 
interests were ornithology and herpetology, and he did little plant collect- 
ing himself. However, he encouraged his fellow countrymen to look into 
the plant life of Nepal. Buchanan Hamilton, later Sir Francis Hamilton, 
onetime Superintendent of the Honourable East India Company’s Botanic 
Gardens at Calcutta, was perhaps the first botanist to visit Nepal. He 
stayed in Kathmandu for about a year (1802—1803) and collected plants 
en route to the valley from the border town of Raxaul (India) and in the 
valley itself. Nathaniel Wallich, who succeeded Hamilton as Superin- 
tendent of the Gardens at Calcutta, was granted permission by the 
Nepalese Government to visit Kathmandu in 1820. He stayed in the 
capital city from December 20, 1820 to November 7, 1821, and “visited 
only one place outside Nepal Valley, viz. Nuwakot, but persuaded pil- 
grims to bring back curiosities to him when they went to Gosainthan”’ 
(Burkill, 1904). The collections of Hamilton and Wallich were catalogued 
by David Don (1825). 

After prolonged negotiations, J. D. Hooker got permission to under- 
take a botanical expedition in eastern Nepal in 1848. He followed the 
Tamir (Tamur) and Arun valleys for a considerable distance and reached 
as far north as Walunchung. He wanted to cross into Tibet, but the por- 


1964] BHATT: NEPAL 149 


ters refused to go any farther north. He, therefore, crossed into Sikkim 
which he is credited with having ‘‘discovered”’ for the British. Hooker’s 
Himalayan Journals (1855) are truly classics in the botanical literature 
of the world. He reported on the general features, topography, vegetation, 
and climate of a region unknown to the western world at that time. His 
description of the local inhabitants is candid and humorous. 

Dr. J. Scully in 1876 and J. F. Duthie in 1880 to 1884 are reported 
(Kitamura, 1955) to have collected plants in parts of western Nepal that 
adjoin Kumaon division of India. 

In 1904, 83 years after Wallich, Burkill (1904), another British botan- 
ist, visited Kathmandu, Nuwakot, and other points in the valley where 
Wallich had collected. It is obvious from reading accounts by Burkill and 
other visitors of that period that the foreigners were forbidden to wander 
around by themselves, and the Nepalese Government rarely gave them 
permission to visit parts of the country other than Kathmandu. 

From 1904 to 1927, no significant activities in plant exploration were 
carried out in Nepal. However, between 1927 and 1937, Captain Lall 
Dhwoj, a Nepali, and Prof. K. N. Sharma, an Indian, collected exten- 
sively for the British Museum in central and western Nepal. The Nepalese 
Government at this time also took interest in the native plants, especially 
those with medicinal importance. A botanical farm was established in 
Shivpuri at 8943 ft altitude about six miles northeast of Kathmandu, to 
produce on a commercial scale plants such as Aconitum, Digitalis, and 
Lobelia. Sharma, who headed the Botanical Deaprtment of the Govern- 
ment of Nepal, claims to have collected plants from such inaccessible 
areas as Rasuwa Garhi and Humla. As a result of the efforts of Dhwoj 
and Sharma, many new species, among them Meconopsis regia and 
M. dhwojii, were discovered. Some of these now have a permanent place 
in the private gardens of the world. Landon’s book contains a list of 
plants from Nepal which was compiled with the help of the Director of 
the Royal Botanical Gardens at Kew (Landon, 1928). 

Oleg Polunin (1950, 1952) was a member of Tilman’s expedition to 
Nepal Himalaya. He surveyed the flora of Langtang Himal and Rasuwa 
Garhi region (1952), and found Gentiana nubigena, Saussurea gossipi- 
phora, primulas and rhododendrons at an elevation of 16000 ft. 

As already mentioned, the revolution of 1950-1951 brought about a 
significant change in the outlook of the Government and people of Nepal 
toward foreigners. In the post-revolution period, westerners were given 
greater freedom to explore the country, and the many mountaineering 
expeditions that have come to Nepal in the last decade gave opportunities 
to a number of botanists to explore plant life in the Himalayas. Col. D. 
Lowndes (1954), a member of Tilman’s second expedition to Nepal 
in 1950, collected seeds and plants in the vicinity of the Marsyandi 
River, Managbhot, and the Jargeng Khola. All of these areas are in 
central Nepal, justly famous for such mountains as Annapurna, Macha- 
pucche and Dhaulagiri. Some of the high altitude plants collected by 


150 MADRONO [Vol. 17 


Col. Lowndes were: Pedicularis (12000 ft), Primula (10000-14000 ft), 
Lonicera (11500-13000 ft), Ephedra (10000-14000 ft) and Delphinium 
(16000 ft). Zimmerman (Vautier, 1959), who accompanied the Swiss 
expeditions to the peaks at Everest (29082 ft) and Cho Oyu (26750 ft) 
in 1952 and 1954 respectively, collected plants en route to Namche Bazar 
from Kathmandu, and also in some of the eastern districts of Nepal. 

In the spring of 1952 the British Museum in conjunction with the 
Royal Horticultural Society sent a team of three botanists, L. H. J. 
Williams, Oleg Polunin, and W. R. Sykes, to collect in western Nepal. 
They explored an area of about 1000 sq mi in the districts of Jumla, 
Humla, Jajarkote, and Sallyan, and collected a total of 5000 herbarium 
specimens, 250 live plants, and 150 seed specimens in their eight-month 
expedition, collecting mostly in areas lying between 7500 and 20000 ft 
altitude (Williams, 1952; 1953). Williams led another expedition to 
West Nepal in 1954, the party consisting of W. R. Sykes and J. D. A. 
Stainton, both botanists, K. Hyatt, a zoologist, and J. Auinlan, an 
entomologist. Dr. V. Puri of Meerut College, India, was also with this 
expedition for about two months. They collected plants south of Dhaula- 
giri (26795 ft), in the Annapurna Range (20000-26041 ft), and in the 
Kaligandaki Valley. A number of new plants were described from the 
collections made in 1952 and 1954 by these British Museum parties 
(Sykes, 1956). 

The Fauna and Flora Research Society of Japan sent two scientific 
and mountaineering expeditions to Central Nepal in 1952 and 1953. 
Sasuke Nakao, a botanist, accompanied the Japanese expedition to 
Manaslu both years, collecting plants in the vicinity of Ganesh Himal, 
Siringi Himal, Manaslu, and also en route from Kathmandu to Pokhara. 
The latter town has served as a base camp for many climbing parties in 
Central Nepal. Members of these two Japanese expeditions collected a 
total of 958 species of phanerogams and 210 species of crytogams, of which 
300 were new to Nepal (Hirano, 1955; Horikawa, 1955; Kawakita, 
1956). Because the Japanese explorers surveyed only a limited area in 
Central Nepal, some of the identification and remarks on distribution 
may not be very accurate. 

In recent years some Indian botanists have visited Nepal. S. K. 
Mukerji, Keeper of the Indian Botanic Garden at Calcutta, collected 
along the Nepal-Sikkim border in 1949. Banerji (1952; 1952a; 1953; 
1958) made six trips to Nepal between 1948 and 1956, but he confined 
his collections to along the main road connecting the capital with all 
the district headquarters lying east of Kathmandu. He listed 169 species 
of flowering plants belonging to 123 genera and 51 species, of which 
16 are new records for the country. D. D. Awasthi (1960) reported upon 
the lichen flora of Nepal. His collections are mainly from Biratnagar and 
Kosi Valley. 

J. E. M. Arnold (Davidson, 1955), a member of the Oxford University 
Expedition to West Nepal, collected plants for the British Museum in 


1964 | BHATT: NEPAL rod 


that region. Swan (1961) made two trips to the Nepal Himalaya, first 
with the American Himalayan Expedition to Makalu (27790 ft) in 
1954, and again in 1960 with Sir Edmund Hillary’s Yeti hunting expedi- 
tion. He collected plants and animals in the neighborhood ‘of Barun 
Glacier and certain other unnamed peaks in that region,’ and found 
evidence of life at the extreme altitudes of 19000 to 22000 ft. 


FLORISTIC REGIONS OF NEPAL 


Hooker (1904) divided the Himalaya into eastern and western phyto- 
geographic regions. He considered the eastern region to extend from 
Kumaon to Mishmi hills in upper Assam, and the western region to 
extend from Chitral to Kumaon division. However, subsequent studies 
have shown that “Nepal is a meeting place of the eastern and western 
Himalayan flora” (Stearn, 1960). The flora of humid mountain areas 
shows a close correspondence with that of western China, Formosa, and 
Japan (Kitamura, 1955), while the plants in the high mountain arid 
zone are the same as those found in Tibet. There seems to have been a 
migration of plants across the Himalayan mountains through gaps in the 
eastern and northwestern end of the chain. The southeastern elements 
of the Nepalese flora are known to extend as far west as Kali Gandki, 
while boreal species (Poa annua and Capsella sp.) have been collected in 
Walungchung. Thus the dividing line between the eastern and north- 
western floristic elements cannot run along Baghmati River, as Banerji 
(1952a) seemed to think, because the flora on the two sides of Kali Gan- 
daki is known to be different (Williams, 1953). It is therefore reason- 
able to assume that the western limit of the southeastern flora could as 
well be this river. 

The collections from Nepal are stored in the herbaria of the British 
Museum, Royal Botanic Gardens at Kew, Indian Botanic Garden, Lin- 
nean Society of London, Missouri Botanic Garden, and Harvard Univer- 
sity. Collections are also known to occur in the museums at Paris and 
Geneva. The author wishes to express his sincere thanks to Lincoln 
Constance of the University of California, Berkeley, for his help in 
securing a fellowship that made this survey possible during the author’s 
stay at the above University. 

Tri-Chandra College, Kathmandu, Nepal 


BIBLIOGRAPHY 


ABRENDT, L. W. A. 1956. A new dwarf barberry from Nepal. Jour. Roy. Hort. Soc. 
81:135 

ASAHINA, Y. 1955. Lichens. Jn Fauna and Flora of Nepal Himalaya. Kyoto Univer- 
sity, Japan. 

AwastTHt, D. D. 1960. Contributoins to the lichen flora of India and Nepal. I. Genus 
Physcia (Ach.) Vain. Jour. Indian Bot. Soc. 39:1-21; I]. The Genus Anaptychia 
Korb. Ibid. 39:415-442. 

BANERJI, M. L. 1952. Some noteworthy plants from east Nepal. Jour. Indian Bot. 
Soc. 31:152-153. 

. 1952a. Observations on the distribution of gymnosperms in eastern Nepal. 
Jour. Bombay Nat. Hist. Soc. 51:156-159. 


152 MADRONO [Vol. 17 


. 1953. Plants from east Nepal. Parts I-III. Jour. Bombay Nat. Hist. Soc. 
51:407—423, 543-560, 773-788. 
. 1958. Botanical exploration in east Nepal. Jour. Bombay Nat. Hist. Soc. 
55:243-268. 
BurkKILL, I. H. 1904. Notes from a journey to Nepal. Rec. Bot. Soc. India 4:59-140. 
Davipson, I. 1955. Oxford University Expedition to West Nepal (1954). Bull. Oxford 
Univ. Exploration Club. 
Don, D. 1825. Prodromus florae nepalensis. London. 
Hacen, T. 1960. Nepal—the kingdom in the Himalayas. Oxford Book & Stationery 
Co. Calcutta: 
Herxtots, G. A. C. 1962. Some plants from Chandragiri ridge. Part I. Jour. Roy. 
Hort. Soc. 87:18-23. 
Hirano, M. 1955. Fresh water algae. Jn Fauna and Flora of Nepal Himalaya. Kyoto 
University, Japan. 
Hooker, J. D. 1855. Himilayan Journals. 2 Vols. John Murray, London. 
. 1904. A sketch of the flora of British India. London. 
, and others. 1875-1897. The flora of British India. 7 Vols. London. 
HorriKkawa, Y. 1955. Mosses. Jn Flora and Fauna of Nepal Himalaya. Kyoto Univer- 
sity, Japan. 
Karan, P. P. 1960. Nepal—a cultural and physical geography. University of Kentucky 
Press, Lexington. 
Kawakira, J. 1956. Vegetatoin. Jn Land and crops of Nepal Himalaya. Kyoto Uni- 
versity, Japan. 
KITAMURA, S. 1955. Flowering plants and ferns. Jn Fauna and Flora of Nepal Hima- 
laya. Kyoto University, Japan. 
Lanpown, P. 1928. Nepal. Constable, London. 
LownpkEs, D. G. 1954. News from Nepal. Quart. Bull. Alpine Gard. Soc. 22:215-229, 
313-322. 
Luptow, F. 1955. New Primula from Nepal. Jour. Roy. Hort. Soc. 80:428—-429. 
LyaLL, H. G. 1958. A pink Meconopsis from Nepal. Jour. Roy. Hort. Soc. 83:122. 
PoLuntn, N. 1960. Introduction to plant geography. McGraw-Hill, New York. 
PotunIn, O. 1950. An expedition to Nepal. Jour. Roy. Hort. Soc. 75:304-315. 
. 1952. A natural history of the Langtang valley. Appendix. Jn Tilman’s 
Nepal Himalaya. The University Press, Cambridge. 
Rosse, E. 1954. Report to the Government of Nepal on Forestry. United Nations 
Food and Agri. Organ. Rep. No. 209. Rome. 
RoxBurGH, W. 1832. Flora Indica. 3 Vols. Serampore. 
SmiTH, H. 1958. Saxifraga of the Himalaya. I. Section Kabchia. Bull. Brit. Mus. 
Hort. Soc. 79:357-358. 
SmitH, W.W. and H. R. FLETCHER. 1954. A new Primula from Nepal. Jour. Roy. 
Hort. Soc. 79:357-358. 
Soest, J. L. van. 1960. New species of Taraxacum from the Himalayan region. Bull. 
Brit. Mus. Bot. 2:263-273. 
STEARN, W. T. 1960. Allium and Milula in the central and eastern Himalaya. Bull. 
Brit. Mus. Bot. 2:161-191. 
Swan, L. W. 1961. Ecology of the high Himalayas. Sci. Am. 205 (4) :68—-78. 
Sykes, W. R. 1956. 1954 expedition to Nepal. Part I. Jour. Roy. Hort. Soc. 80:538- 
544; Part II. Ibid. 81:6-14. 
VauTIER, S. 1959. Resultats des Expeditions Scientifiques Genevoises au Nepal en 
1952 et 1954. (Partie Botanique). Candollea 17:41-221. 
Wesster, G. L. 1961. The altitudinal limits of vascular plants. Ecology 42:587-590. 
Watticu, N. 1824-1826. Tentamen florae napalensis illustratae. London. 
. 1831. Plantae asiaticae rariores. Vol. 2. London. 
Wuiuiams, L. H. J. 1953. The 1952 botanical expedition to Nepal. British naturalists 
in Nepal. Times (London), January 24. 
. 1953. The 1952 expedition to western Nepal. Jour. Roy. Hort. Soc. 78:323- 
S375 


1964] CONDIT: FICUS 153 


CYTOLOGICAL STUDIES IN THE GENUS FICUS. III. 
CHROMOSOME NUMBERS IN SIXTY-TWO SPECIES 


Ira J. CONDIT 


Ficus is a genus the woody plants of which are widely distributed in 
tropical and subtropical countries. It is a fantastic as well as a confusing 
member of the plant kingdom. It is fantastic because of the extreme diver- 
sity of plant forms exhibited, i.e., size, habit of growth, leaves, flower 
characters, and also by reason of the symbiotic relationship of certain 
insects which inhabit the receptacles. It is a confusing group because 
of the many perplexing problems in nomenclature and because of the 
difficulties encountered in identification of specimens either collected in 
their native habitat or found growing under cultivation. Interest in the 
genus has resulted in the description of many species, variously estimated 
from 600 to 1500. In Index Kewensis, 1887 specific names are listed for 
Ficus. 

Cytologically, however, this large genus is not well known (Condit, 
1928; 1933; Krause, 1930; 1931; Mangenot and Mangenot, 1958; 
Sugiura, 1936). It is believed that the species reported earlier (Condit, 
1928; 1933) were accurately determined with the following exceptions: 
F.. glomerata Roxb. should be F. racemosa L.; F. indica L. growing in the 
Lyon Arboretum, Honolulu, is regarded as a distinct species and not con- 
specific with F. bengalensis L.; the authenticity of plants grown in Cali- 
fornia under the name F’. asperrima Rozb. is questionable and should be 
called F. gibbosa Blume; and F. nigens? should be F. ingens Miq. The 
somatic chromosome number was found to be 26 in 37 species. In seven 
species the gametic number was 13. In some instances the somatic number 
was reported to be 24, 25, 27, or 28 and the gametic number 14. Neither 
the species, nor the unisexual plants with long-styled pistillate flowers or 
the bisexual caprifigs with short-styled pistillate and staminate flowers, 
seem to be distinguished by peculiarities in chromosome morphology. 

This paper presents chromosome numbers of 62 species of Ficus plus 
counts of a few species unidentified or previously reported under other 
specific names (table 1). Nomenclature is based mainly on the recent 
papers of Corner (1960; 1961; 196la) and DeWolf (1960). 

Root tips from potted plants grown mostly in a greenhouse at the 
Citrus Research Center, University of California, Riverside (CRC), fur- 
nisher most of the material for study. Plant introduction numbers (PI) 
of the United States Department of Agriculture and of the Hawaiian 
Sugar Planters’ Association (HSPA) are given when applicable. Root 
tips were fixed in Karpechenka’s solution, embedded in paraffin, sectioned 
and stained in Haidenhain’s iron-haematoxylin, as reported in the earlier 
papers. 

Practically all cultivated species of Ficus have been introduced with- 
out the specific insect which normally inhabits the receptacles. I have 
made no studies nor have I found any report as to the extent to which the 


154 MADRONO [Vol. 17 


TABLE 1. CHROMOSOME COUNTS IN FIcUS 
Diploid group, 2n=26 

Ficus acanthocarpa Lev. & Van., HSPA 1686, China. F. afzelii G. Don, from 
Missouri Botanical Garden, a Belgian Congo species. F. amplissiam Smith, PI 
93398, Poona, India as F. tsiela Roxb. F. auran‘iaca Grif. var. parvifolia Corn., 
PI 134991, Philippines. F. aurea Nutt., CRC, Florida. F. auriculata Lour., PI 77952, 
India. F. avi-avi Bojer, CRC, Madagascar. F. awkeotsang Makino (F. pumila L.), 
CRC, Taiwan. F. bussei Mildbr. & Burr., PI 62806, Tanganyika, Africa. F. cabusana 
Standl. & Steyerm., CRC, from a California nursery, identified at F. F. camarinensis 
Merr., HSPA 1819, 1829, 1849, Luzon, Philippines. F. capensis Thunb., PI 73935, 
Gambia, Africa. F. citrifolia Mill., CRC, Paraguay, as F. eximia Schott var. glabra 
Hassl. F. cocculifolia Baker var. sakalavarum (Baker) H. Perr., CRC, Madagascar. 
F. columnaris Muell. & Moore (F. macrophylla Desf.), PI 141765, Sydney, Australia. 
F. costaricana Miq., PI 262188, El Salvador. F. congesta Roxb., HSPA 1545, 1775, 
Philippines, as F. saiterthwaitez Elmer. F. coronata Spin., CRC, Queensland, Aus- 
tralia, as F. scabra Forst. F. cotinifolia H. B. K., PI 159445, Alamos, Mexico. F. doli- 
aria Mart., CRC, Campinas, Brazil. F. geniculata Kurz., CRC, Buitenzorg, Java. 
F. gnaphalocarpa Miq., CRC, Senegal, West Africa. F. goldmanii Standl., CRC, 
Alamos, Mexico. F. hillii Bailey (F. retusa L. var. nitida Thunb.), Australia. F. his- 
pida L., PI 80081, Darjeeling, India. F. znsipida Willd., PI 74426, Summit, Canal 
Zone. F. iteophylla Miq., PI 137932, Nigeria, Africa. F. lapathifolia Miq., CRC, 
Chiapas, Mexico. F. krishnae C. DC., PI 123211, a bud sport of F. bengalensis L. 
F. mallotocarpa Warb., HSPA 5639, Nairobi, Africa. F. mammilifera Warb., CRC, 
Jamaica. F. minahassae Miq., CRC, Laguna, Philippines. F. monckii Hassl., CRC, 
Buenos Aires, Argentina. F. montana Burm., PI 101330, a Malaysian species com- 
mon in nurseries. fF. nekbudu Warb., Los Angeles State and County Arboretum, 
Arcadia, an east African species common in cultivation. F. nota Merr., PI 134993, 
Philippines. F. nympheaefolia Mill., PI 161328, Caracas, Venezuela. F. obtusifolia 
H.B.K., PI 161324, Mazatlan, Mexico. F. perforata L., CRC, Jamaica, as F. wilsonii 
Warb. F. pertusa L., PI 92350, Chiapas, Mexico. F. petiolaris H.B.K., PI 161331, 
Alamos, Mexico. F. pilosa Blume, CRC, Bogor, Indonesia. F. preuszz Warb., PI 
262356, Netherlands, native in the Kamerun, Africa. F. procera Reinw. var. crassi- 
ramea King, PI 94297, Buitenzorg, Java. F. radulina Wats., PI 159446, Alamos, 
Mexico. F. ribes Blume, HSPA 1464, Philippines; HSPA 3275, Java. F. rigo Bailey, 
PI 94210, British New Guinea. F. rumphii Blume, CRC, Allahabad, India. F. solda- 
nella Warb., CRC, Pretoria, South Africa. F. stricta Miq., PI 268135, Laguna, Philip- 
pines. F. subcordata Blume, CRC, Buitenzorg, Java, as F. garciniaefolia Miq, F. 
thonningit Blume., CRC, Nairobi, East Africa. F. tanctoria Forst., PI 78577, Guam. 
F. umbellata Vahl., PI 75751, Gold Coast, Africa. F. urbaniana Warb., PI 161335, 
Caracas, Venezuela. F. urceolaris Hiern., PI 76424, Uganda, Africa. F. variegata 
Blume, PI 122987, Botanic Gardens, Straits Settlements. F. volkensiz Warb., PI 
78261, Tanganyika, Africa. F. wildemaniana Wildem. & Th. Dur., an African species 
introduced by a Florida nursery from Denmark. F. sp., PI 97571, Grenadine Islands; 
PI 101329, France; PI 103504, India; PI 95089, Singapore. 


Tetraploid group, 2n=52 
Ficus burkei Miq., CRC, Pretoria, South Africa. F. hochstetteri A. Rich., CRC, 
Nairobi, East Africa. F. pretoriae Burtt-Davy, PI 137595, Pretoria, South Africa. 
F. sonderi Miq., CRC, Pretoria, South Africa. F. stuhlmani Warb., PI 161334, Pre- 
toria, South Africa. 


Miscellaneous Group 
Ficus elastica Roxb. ‘Decora,’ 2n=39, a common horticultural variety. F. duseni 
Warb., 2n=26, 52, France, native of tropical Africa. F. macrosyce Pitt. (F. insipida 
Willd.), 2==26, 52, CRC, Caracas, Venezuela. F. -palmeri Wats., 2n=26, 50’?, CRC, 
La Paz, Baja California, Mexico. 


1964] CONDIT: FICUS 155 


staminate flowers of cultivated species develop before the fruit shrivels 
and drops. 

Of the new counts reported, 53 are diploid with a somatic chromosome 
number of 26. One triploid is reported, F. elastica ‘Decora,’ in which 
2n=39. This cultivar of the common india rubber is apparently from a 
seedling selected in Belgium over 30 years ago on account of its broad 
leaves and bright coloration. Since no other similar variety of F. elastica 
has been reported with diploid chromosome groups, there is a possibility 
that the triploid ‘Decora’ is a hybrid. 

Five species are tetraploid with 2n=52. Four of these, F. burkei, F. 
pretoriae, F. sonderi, and F. stuhlmant, are indigenous to South Africa, 
while F. hochstetteri is native in East Africa. The last is closely related 
to F. thonningu which is diploid (table 1). Further cytological studies 
of these two species are warranted. Plants of these five species show no 
superficial characters suggesting a condition of tetraploidy when under 
cultivation in southern California. 

In three species some prepared materials show chromosome counts to 
be 26, others 52. Ficus dusenit, for example, a speceies of tropical Africa, 
was represented in our collection by four potted plants. Chromosome 
counts of root-tip material from these plants were conflicting. In one 
plant, both complements, 26 and 52 were counted. In the other three the 
prevailing number was 52. The occurrence of islands of tetraploid cells 
in root-tips of diploids has been reported in tomato (Lesley, 1925). Pos- 
sibly some species are periclinal chimeras of diploid and tetraploid tis- 
sues. Ficus palmer is represented in our collection by four separate slide 
preparations. Two showed diploid complements and two both diploid and 
tetraploid on the same slide. In F. macrosyce one slide showed a diploid 
complement, another showed both diploit and tetraploid complements. 


Citrus Research Center, University of California, Riverside 


LITERATURE CITED 


Conpit, I. J. 1928. Cytological and morphological studies in the genus Ficus. I. 
Chromosome number and morphology in seven species. Univ. Calif. Publ. Bot. 
1:233-244. 1933. II. Chromosome number and morphlogy in thirty-one species 
Ibid. 17:61-74. 

Corner, E.J.H. 1960. Taxonomic notes on Ficus Linn., Asia and Australasia. 
Gard. Bull. Singapore 17:368-485; 1961. Ibid. 18:1-69, 83-97. 

. 1961a. Evolution. Jn R.E. MacLeod and L.S. Cobley, editors. Contem- 
porary botanical thought. Quadrangle Books, Chicago. 

DeWotr, G. P., Jr. 1960. Ficus. In R. E. Woodson, Jr. and R. W. Schery. Flora 
of Panama. Ann. Missouri Bot. Gard. 47:147-165. 

KRAUSE, O. 1930. Cytologische Studien bei den Urticales. Ber. Deutsch. Bot. Ges. 
48:9-13. 

. 1931. Zytologische Studien bei den Urticales. Planta 13:29-84. 

LestEy, M.M. 1925. Chromosomal chimeras in the tomato. Am. Nat. 59:570-574. 

MANGENOT, S. and G. Mangenot. 1958. Deuxiéme liste de nombres chromosomiques 
nouveaux chez diverses Dicotyledones et Monocotylédones d’Afrique Occiden- 
tale. Bull. Jard. Bot. Bruxelles 28:315-329. 

SuciurA, T. 1936. Studies on chromosomes in higher plants. Cytologia 7:544-595, 


156 MADRONO [Vol. 17 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF THE 
GENUS MIMULUS (SCROPHULARIACEAE). VI. NEW 
NUMBERS IN M. GUTTATUS, M. TIGRINUS, 

AND M. GLABRATUS! 


M. M. M1, B. B. MUKHERJEE, AND R. K. VICKERY, JR. 


The new chromosome numbers found in this investigation significantly 
enlarge the cytogenetic concept of M. guttatus DC. and amplify the 
knowledge of such other species of the section as M. nasutus Greene, 
M. tigrinus Hort. ex Sieb. & Voss, M. luteus L., and M. glabratus 
H.B.K. (figs. 1, 2). The cytological techniques employed were slightly 
modified from those previously used (Mukherjee and Vickery, 1962). 
The fixative was strengthened from 2 to 3 parts glacial acetic acid mixed 
with the usual 1 part absolute ethanol saturated with ferric acetate. The 
length of fixation was reduced from 24 to 4 hours. The slides were made 
permanent by first, dehydrating them in an ethanol vapor chamber for 
3 to 4 days. Next, the cover slips were ringed with diaphane and the 
slides were left in the vapor chamber for an additional 2 to 3 days. Rep- 
resentative chromosome configurations were recorded either photograph- 
ically or by drawings made with the aid of a camera lucida. Vouchers of 
each culture will be deposited in the University of Utah herbarium. 

Among the M. guttatus populations ten, in addition to the 34 pre- 
viously sampled (Vickery, 1955; Mukherjee et al., 1957; Mukherjee and 
Vickery, 1959; 1960), were found to have n=14 chromosomes (table 1). 
Two others had n= 16 and genetic tests now in progress may well indi- 
cate that these populations belong to a new species that should be seg- 
regated from M. guttatus. The final M. guttatus population had n=28 
chromosomes. 

This tetraploid, n= 28 population was studied cytologically and genet- 
ically. In meiosis, its chromsomes formed regular bivalent associations 
with no indications of tri- or tetravalent configurations. Several plants 
of this population were crossed with a highly fertile diploid WM. guttatus 
(5052). The Fi hybrids were vigorous but sterile. Of 31 pollen mother 
cells observed at or near first metaphase of meisosis 23 had 14; and 14;, 
three had 12;, and 18,, and five had 9;; and 24;. This pairing behavior 
and that of the tetraploid plants themselves indicate that if the Verde 
Valley population is an autotetraploid as its close morphological sim- 
ilarity to diploid M. guttatus suggests, and not an allotetraploid, then 
it must have arisen long enough ago to permit the accumulation of suf- 
ficient gene differences in its chromsomes to prevent autosyndesis. Even 
though this tetraploid population will not produce fertile hybrids with 


1 This work was supported by grants from the National Science Foundation and 
the University of Utah Research Fund. Many of the counts here reported are 
included in the dissertations submitted by the senior authors to the University of 
Utah in partial fulfillment of the requirements for the Ph.D. degree. 


1964] MIA, MUKHERJEE, & VICKERY: MIMULUS 157 


6260 6212 
See ae : 
0 10 20 micra 


Fic. 1. Meiotic chromosomes of Mimulus: M. tigrinus, 6260, has n = 30 chromo- 
somes, M. guttatus, 6212 and 6250, has n= 16 and n = 28, respectively. The first 
cell is in first anaphase whereas the latter two are in first metaphase. The dotted 
line delimits the two planes of focus in the cell of 6260. The camera lucida drawings 
were at an original magnification of X 2,720, reduced to approx. x 900 in repro- 
duction. 


diploid populations, we hesitate to suggest that it is a separate species. 
Probably other such tetraploid populations have arisen or will arise here 
and there throughout the extensive range of M. guttatus (Grant, 1924). 
Possibly, as in Galium (Ehrendorfer, 1955) such polyploid populations 
would be able to exchange genes. If such a situation proved to be the 
case, a restricted gene flow would still be potentially possible from the 
diploids to the tetraploids and amongst the latter. This gene flow would 
hold the tetraploid and diploid forms together, at least loosely, in a 
common evolutionary pathway. 

The chromosome counts for M. nasutus, M. tigrinus, and M. luteus 
(table 1) include a new number, n=30, for M. tigrinus. Previous reports 
(Brozek, 1932; Mukherjee and Vickery, 1959; 1960) are for n=32 
populations. The latter number is also characteristic of M. luteus 
(table 1; Mukherjee and Vickery, 1959; 1960) from which the horticul- 
tural species M. tigrinus was derived. However, one M. luteus culture 
(Mukherjee and Vickery, 1960) was found to have n=30 + 0, 1, or 2 
chromosomes. In view of this variable population, the discovery of a 
consistently n=30 M. tigrinus population is not surprising, but inter- 
esting. 

Mimulus glabratus H.B.K. is a highly polymorphic, widespread com- 
plex of related varieties (Grant, 1924). It occurs as scattered popula- 
tions that range from the north woods of Michigan (Fassett, 1939) and 
the Great Basin of the Western States (Pennell, 1947), south through 
Mexico to Guatemala and then from Peru on south to Argentina and 


158 MADRONO [Vol. 17 


the Juan Fernandez Islands (Skottsberg, 1953). The striking diversity 
and scatter of the members of the complex raise a question as to the 
closeness of their relationship. 

The most outstanding result of the investigation was the demonstra- 
tion of aneuploidy at each of the three polyploid levels in the complex, 


5747 6201 6192 


6209 


y” oO 2 micra 


0 107 920 


Fic. 2. Meiotic chromosomes of Mimulus: M. glabratus var. utahensis, 5747 
(n = 14), 6201 (n=15);.M. glabratus var. glabratus,. 6192, 6193. (mn =30), 6197, 
6209 (n = 31); M. glabratus var. parviflorus, 6163, 6184, 6185 (n = 46). The figures 
were selected to illustrate the appearance of the chromosomes in several stages of 
meiosis, diakenesis, 6192; metaphase I, 6209, 6184, and 6185; metaphase II, 5747, 
6201, 6193, and 6163; anaphase II, 6197. The camera lucida drawings were at an 
original magnification of X 2,100, reduced to approx. X 700 in reproduction. 


1964] MIA, MUKHERJEE, & VICKERY: MIMULUS 159 


TABLE 1. CHROMOSOME CoUNTS IN MIMULUS, SECTION SIMIOLUS 


M. glabra‘us var. glabratus.n = 30: e of Morelia along route 15, Michoacan, Mexico, 
8300 {t., Wiens 2519 (6192) ; Hidalgo, along route 15, Michoacan, Mexico, 8300 
ft, Wiens 2520 (6193); El Salto, km 1104 on Durango-E] Salto rd, Durango, 
Mexico, 8500 ft, Wiens 2639 (6210).n = 31: Santa Rosa, along route 15, Michoa- 
can, Mexico, ca 8500 ft, Wzens 2521 (6194); n of Tehuacan along route 150, 
Puebla, Mexico, 7200 ft, Wiens 2552 (6197) ; Ciudad Mendoza, along route 150, 
Vera Cruz, Mexico, 4800 ft, Wzens 2555 (6198); Cofre de Perote, Vera Cruz, 
Mexico, 9500 ft, Wiens 2575 (6199); n of Puebla along route 119, Tlaxcala, 
Mexico, 6700 ft, Wzens 2588 (6200); El Salto, km 1050 on Durango-El Salto 
rd, Durango, Mexico, 8000 ft, Wzens 2635 (6209). 

M. glabratus var. parviflorus. n = 46: Auetrihue, Argentina, Diem, in 1959 (6162) ; 
Lumaco-Puente del Cina, Malleco, Chile, 660 ft, Kunkel, Nov. 29, 1958 (6163) ; 
Tafi del Valle, La Quebradita, Tucuman, Argentina, 7300 ft, de la Sota, Feb. 7, 
1959 (6184); e of La Divisiona, in the Cordillera Azul, Loreto, Peru, 6600 ft, 
Mathias 5151 (6185). 

M. glabratus var. utahensis. n = 14: Telephone Canyon, Pilot Cone, Mineral Co., 
Nevada. 5500 ft, Figg-Hoblyn, July 4, 1950 (5747).n = 15: Geneva Steel Plant, 
Utah Co., Utah, 4490 ft, Lindsay, Apr. 4, 1959 (6156) ; Rio Tierra Quemada and 
route 57, Guanajuato, Mexico, 5700 ft, Wzens 2598 (6201); Saltillo, route 57, 
Coahuila, Mexico, 5500 ft, Wzens 2509 (6203). 

M. guttatus.n = 14: Lewiston grade, Nez Perce Co., Idaho, 840 ft, Preece, Murdoch, 
& Rumely 2167 (5262); Botanic Garden, Wageningen, Netherlands, Venema 
1949 (5306) ; Mutica, Sonora, Mexico, Gentry 2194 (5321); Mono Lake, Mono 
Co., California, 6440 ft, Vickery 200 (5397); Holberg’s, Lake Co., California 
3000 ft, Vickery 2043 (6138); near Cache Creek, Lake Co., California, 1300 
ft, Vickery 2044 (6139); West Thumb, Yellowstone Park, Wyoming, 7000 ft, 
Mia, July 25, 1959 (6186) ; Cache Creek, Lake Co, California, 1250 ft, Campbell, 
June 18, 1948 (6288), seeds from 131652 (TFX); Atlantic City, Wind River 
Mtns., Fremont Co., Wyoming, 7680 ft, Edmunds, Aug. 18, 1960 (6303) ; Lobdel 
Lake, Sweetwater Mtns., Mono Co., California, 9400 ft, Beaman 957 (5257). 
n= 16: km 1155, Durango-Mazatlan rd, Durango, Mexico, 8500 it, Wiens 
2643 (6212); w of crest of Durango-Mazatlan rd, Durango, Mexico, 8300 ft, 
Vickery 2616 (6273). n= 28: Verde Valley, Yavapi Co., Arizona, 3010 ft, 
Vickery 2593 (6250). 

M. luteus. n = 32: Auetrihue, Aregntina, Diem, in 1959 (6161). 

M. nasutus.n = 14: Alabama Hills, Inyo Co., California, 500 ft, Dedecker, May 30, 
1956 (6060). 

M. tigrinus. n = 30: Cultivated in gardens, Kathmandu, Nepal, 4500 ft, Brydon, 
May, 1960 (6260). 


as follows: M. glabratus var. utahensis Penn., n = 14 and n = 15; 
M. glabratus var. glabratus, n = 30 and n = 31; and M. glabratus var. 
parviflorus (Lindl.) Grant, n = 45 and n = 46 (table 1). For M. glab- 
ratus var. utahensis, the additional determinations verified the previous 
reports (Vickery, 1955; Mukherjee, e¢ a/., 1957; Mukherjee and Vickery, 
1960). The counts now available suggest a generally central and west- 
ern Great Basin distribution for the n = 14 form and an eastern Great 
Basin and Mexican Plateau distribution for the n = 15 form. At first, 
culture 5852 from the Wendover, Utah population was thought to be 
M. guttatus (Vickery, 1955), but further study has shown it to belong 
to M. glabratus var. utahensis. It is the most eastern n = 14 population 
found thus far. 


160 MADRONO [Vol. 17 


The new counts show WM. glabratus var. glabratus to be a tetraploid 
as is M. glabratus var. fremontu (Benth.) Grant (Mukherjee and Vick- 
ery, 1960). However, morphologically var. glabratus is far closer to var. 
utahensis than it is to var. fremontu. The few n = 30 populations of 
var. glabratus found to date occur only in Michoacan and Durango, 
Mexico whereas the n= 31 populations are more common and more 
widespread (table 1). 

The new counts for M. glabratus var. parviflorus plus the previous 
n — 45 report (Mukherjee and Vickery, 1959) reveal that this variety 
aslo has two chromosome forms. Morphologically and cytologically M. 
pilosiusculus H.B.K. (Mukherjee and Vickery, 1959) belongs to this 
taxon. In contrast, the Tafi del Valle population (6184) is very distinct 
from the other n = 46 or n = 45 populations. It is erect and the others 
are prostrate. 

Cytologically, the WM. glabratus complex presents a most interesting 
picture of a polyploid series with aneuploidy at each level. The polyploid 
levels increase from north to south in a striking manner. However, while 
these counts clearly indicate the complexity of the group, an improved 
taxonomic treatment of it must await a cytological study of the other 
varieties (Fassett, 1939; Skottsberg, 1951, p. 784) and a genetic in- 
vestigation of the whole complex. 


Pakistan Atomic Energy Commission, Ramma, Dacca, Pakistan 
Genetics Department, McGill University, Montreal, Canada 
Department of Genetics and Cytology, University of Utah, Salt Lake City 


LITERATURE CITED 


Brozexk, A. 1932. Mendelian analysis of the “red-orange-yellow” groups of flower 
colours in Mimulus cardinalis Hort. Preslia 11:1—10. 

EHRENDORFER, F. 1955. Hybridogene Merkmals-Introgression Zwischen Galium ru- 
brum L. s. str. und G. pumilum Murr. s. str. (Zur. Phylogenie der Gattung 
Galium. IV.) Osterr. Bot. Zietschr. 102:195-234. 

FasseEtT, N. C. 1939. Notes from the herbarium of the University of Wisconsin. XVIII. 
Rhodora 41:524—525. 

GranT, A. L. 1924. A monograph of the genus Mimulus. Ann. Missouri Bot. Gard. 
11:99-388. 

MUKHERJEE, B.B. and R.K. Vickery, Jr. 1959. Chromosome counts in the section 
Simiolus of the genus Mimulus (Scrophulariaceae). III. Madrofio 15:57-62. 

. 1960. Chromosome counts in the section Simiolus of the genus Mimulus 
(Scrophulariaceae). IV. Madronio 15:239-245. 

. 1962. Chromosome counts in the section Simiolus of the genus Mimulus 
(Scrophulariaceae). V. The chromosomal homologies of M. guttatus and its 
allied species and varieties. Madrono 16:141-155. 

MUKHERJEE, B. B., D. Wiens, and R. K. VicKERY, JR. 1957. Chromosome counts 
in the section Simiolus of the genus Mimulus (Scrophulariaceae). II. Madrono 
14:128-131. 

PENNELL, F. W. 1947. Some hitherto undescribed Scrophulariaceae of the Pacific 
States. Proc. Acad. Phila. 99:155-199. 

SKOTTSBERG, C. 1920-1953. The Natural History of Juan Fernandez and Easter 
Islands Vol. 2. Botany. Almqvist & Wiksells, Uppsala. 

VickeErY, R. K., Jr. 1955. Chromosome counts in the section Simiolus of the genus 
Mimulus (Scrophulariaceae). Madrono 13:107-110. 


1964 | HARVEY & MOONEY: CHAPARRAL SHRUBS 161 


EXTENDED DORMANCY OF CHAPARRAL SHRUBS 
DURING SEVERE DROUGHT 


R. A. HARVEY AND H. A. Mooney 


Chaparral shrub species display a diversity of rooting types ranging 
from shallow fibrous systems to those which are deep penetrating (Hell- 
mers et. al., 1955). It might be assumed that these differences result in 
differential growth periods in response to seasonal changes in the moisture 
levels in the soil profile. The objective of this study was to establish and 
quantify this probable relationship. Toward this end a small topographic 
unit of apparent climatic and edaphic homogeneity was selected that 
included shrubs of several species representing different rooting types. 
This area is located in the Mill Creek drainage of the San Bernardino 
Mountains at an elevation of 3000 ft. Three species were chosen for 
detailed growth measurements. These were Heteromeles arbutifolia 
(Lindl.) M. Rom. (toyon), a woody shrub with a deep penetrating root 
system, Cercocarpus betuloides Nutt. ex T. & G. (birchleaf cercocarpus), 
a woody shrub with a shallow, wide spreading root system, and Salvia 
apiana Jeps. (white sage), a subshrub with a shallow, fibrous root system. 

At monthly intervals commencing in November, 1960, ten samples 
were taken from five shrubs of each species in order to record increases 
in leaf number, area, and weight, as well as increases in terminal shoot 
length and weight. Procedural details are available elsewhere (Harvey, 
1962). These measurements were continued through May, 1961. Soil 
moisture samples also were taken at the same time intervals at six inch 
increments from the surface to a depth of 36 inches. On one occasion 
additional samples were taken to a depth of 120 inches. Precipitation 
data for the area were obtained from the Mill Creek Ranger Station 
which is located one mile from the study area at a slightly lower elevation. 

From July, 1960, to July, 1961, a total of 5.47 in. of precipitation was 
recorded at the Mill Creek Ranger Station. All of this rain fell during 
the period of investigation, but almost half of the total came during 
November. This is far below normal and may be contrasted with the 
previous season’s total of 15.03 in., and with the 20.33 in. received fol- 
lowing the study in 1961-1962. 

Soil moisture was at low percentages at depths below six in. during 
the entire study period. Following the November rain the percentage 
moisture in the soil at a 12 in. depth was 8.5 per cent. From this amount 
the percentage steadily decreased to 4.6 per cent in May. The 15 atmos- 
phere tension value for this soil and depth is 5.1 per cent. Thus, during 
the entire period of study the 12 in. deep soil mosture was at tensions 
near or greater than 15 atmospheres. Deeper than 12 in. the soil drought 
was even more pronounced. 

The terminal branches of Salvia apiana grew only slightly in length, 
mostly less than an inch. Beginning in January a great number of small 


162 MADRONO [Vol. 17 


leaves were produced; however, none of these expanded appreciably. By 
May, because of leaf drop in April, leaf area and number were of the 
same magnitude as in November, and die-back of terminal shoots was 
quite evident. 


Cercocarpus betuloides had no gain in either stem growth or leaf pro- 
duction. Similarly, there was no stem elongation on Heteromeles arbuti- 
folia, Although on the latter species several leaves were produced on each 
terminal branch during December and January, as many were lost during 
March through May. There was no seasonal production of flowers and 
fruit on either species. 

Terminal stem length measurements only were made on shrubs of 
Garrya veatchi Kell., Rhamnus californica Esch., Prunus ilicifolia 
(Nutt.) Walp., and Quercus dumosa Nutt. from February to June. No 
stem growth or flower production occurred on these species during this 
period. 

In summary, shrub growth during the study period as measured by 
the given criteria was essentially non-existent. The one exception was the 
slight growth displayed by the shallow, fibrous-rooted Salvia plants. 

There were no apparent subsequent adverse effects of the prolonged 
drought on these shrubs. The following year, in the fall of 1962, a cur- 
sory examination was made of the three primary shrub species studied. 
In contrast to the negligible growth of Salvia during the drought year, 
terminal branches of this species averaged 46 inches growth during the 
1961-1962 season (flowering branches). Vigorous growth was also evi- 
dent on Heteromeles and Cercocarpus. On all three species, many of the 
terminal buds present during the drought year were dried and had been 
replaced by lateral bud growth. 

Scant evidence is provided to fulfill the original objective of estab- 
lishing a relationship between the rooting type of chaparral shrubs and 
the period of growth activity. Only Salvia apiana, a shallow, fibrous 
rooted species, displayed measurable growth. Presumably, this shrub 
utilized what small amounts of precipitation penetrated the upper levels 
of the soil profile. A sufficient amount of soil moisture was apparently 
not available for the growth of deeper rooted species. The drought that 
occurred during the period of study was of unusual severity. Precipita- 
tion for the 1960-1961 season in the San Bernardino region was the 
lowest recorded in 92 years, and followed two years of subnormal rainfall. 


Even in a normal year the chaparral environment may be considered 
severe for plant growth. The growth period is largely restricted to early 
spring. In the winter when soil moisture is normally available, low tem- 
peratures limit growth. In the summer, although equitable growing tem- 
peratures prevail, soil moisture is limiting, and due to a reduction in 
physiological activity, the plants become dormant. In some seasons, 
under certain circumstances as described in this report, this dormant 
state may be of protracted length with conditions at no time being suit- 


1964 | CAVE: AGAVACEAE 163 


able for growth. That these plants can survive such a prolonged period of 
adverse moisture conditions is a striking indication of their xerophytic 
adaptiveness. 

Department of Botany, University of California, Los Angeles 


LITERATURE CITED 
Harvey, R. A. 1962. Seasonal growth of chaparral shrubs in relation to root types. 
M.A. thesis. Univ. of California, Los Angeles. 
Hetime_ers, H., J. S Horton, G. JUHREN, and J. O'KEEFE. 1955. Root systems of 
some chaparral plants in southern California. Ecology 36:667-678. 


CYTOLOGICAL OBSERVATIONS ON SOME GENERA 
OF THE AGAVACEAE 


Marton S. CAVE 


Even before Hutchinson (1934) set up the family Agavaceae to in- 
clude tribes of woody xerophytes from both the Liliaceae and the 
Amaryllidaceae, McKelvey and Sax (1933), Whitaker (1934), and Sato 
(1935) pointed out that Yucca and Agave, together with certain of their 
allies, must be related because they all have a basic karyotype of 5 long 
and 25 short chromosomes. This karyotype is too unusual to have been 
developed along two different evolutionary lines. 

Since the new family has been constituted there have been numerous 
comparative studies of the genera within the family evaluating the evi- 
dence for and against the erection of the family (Moran, 1949; Wunder- 
lich, 1950; Cave, 1953). According to many botanists today there is 
strong evidence for a close relationship of Yucca and Agave, but their 
agreement with Hutchinson to place these genera with others such as 
Cordyline, Dracaena, Sansevieria, Phormium, Nolina, Dasylirion, and 
Doryanthes is not so strong. 

One of the lines of cytological evidence useful in taxonomy is the study 
of the karyotype. Granick (1944) has summarized the information of 
that date concerning the chromosome numbers in the Agavaceae. Both 
Yucca and Agave appear to have widespread hybridization within each 
genus, but polyploidy so far has been reported only in the latter. Granick 
discussed polyploidy within Agave and concluded that the karyotypes 
were of little value in determination of individual species, but that there 
was a definite correlation between polyploidy and vegetative develop- 
ment. The polyploids appear also to have a wider distribution than the 
diploids. Her counts were made on root tip materials which offer little 
information as to the possible hybrid nature of the plant examined, 
especially in the polyploids. 

The comparatively long time needed for most of the Agavaceae to 
mature is probably one of the reasons more cytotaxonomic work has not 
been achieved on the family. Recently a number of specimens growing 
at the University of California Botanical Garden, Berkeley, have flow- 


164 MADRONO [Vol. 17 


ered, and thus an opportunity to study their meiotic chromosomes was 
presented. In line with the cytological program of the Garden, chromo- 
some numbers of these plants, as well as those of three additional species 
from material fixed in the wild, have been determined. This paper pre- 
sents information that may be helpful in future cytotaxonomic studies 
of the family. 

In making chromosome counts flower buds were fixed in three parts 
absolute alcohol to one part glacial acetic acid. Anthers were squashed 
in aceto-carmine and slides were made permanent according to the tech- 
nique of Bradley (1948). Immediately following the name of the species 
is the number of the permanent slide and the next number is the Uni- 
versity of California Botanical Garden cultivation number. These are 
followed by the place of collection, collector, and the herbarium in which 
vouchers are deposited. The species of Agave are arranged by subgenera 
according to the classification of Berger (1915). Agave, Furcraea, and 
Beschorneria have a basic karotype of 5 long (L) and 25 short (s) 
chromosomes; Nolina and Dorvanthes do not. 


AGAVE, SUBGENUS MANFREDA 


A. sp. (5805; UCBG 56.624-1; Mexico, origin unknown). n = 30. 
Meiosis in PMC’s was regular with 5L -++ 25s chromosome pairs (fig. 1). 
The plant was growing in the greenhouse, but has now been set outdoors. 

A. sp. (62126; near San Luis Potosi, Mexico, Kimnach 294).n = 30. 
Buds collected and fixed in the wild gave a count of 5L + 25s chromo- 
some pairs in regular meiotic behavior (fig. 2). Identification and her- 
barium voucher must await flowering of the specimen at present in the 
Huntington Botanical Garden. 


AGAVE, SUBGENUS LITTAEA 


A, filifera Salm-Dyck. (6106; UCBG 60.328-1; Mexico, origin un- 
known; UC). n = 30. This is an ordinary diploid with 5L + 25s chromo- 
some pairs. No irregularities were noted in meiosis (fig. 3). This number 
agrees with that of Horowitz (McKelvey and Sax, 1933). 

A. lechuguilla Torr. (6266; UCBG 57.475—1; Texas, Helotes, O. Sokol; 
UC). n = 55-60. Granick counted the chromosomes in root tips of three 
specimens of this species and lists 20L + 100s in each. In our plant II M 
in PMC’s gave counts of 10L + 45-50s (fig. 4). It is possible that in 
some instances some of the small chromosomes were missed. Meiosis 
was regular, with only rare lagging of small chromosomes at I A, and 
numerous capsules were produced along the raceme-like inflorescence. 
A few mature seeds were formed. 

A, victoria-reginae T. Moore. (6274; UCBG 57.494—1; Mexico, origin 
unknown; UC). n= 30. This species has 5L + 25s pairs of chromo- 
somes. Meiosis was regular (fig. 5). No seed was set, however. 

A. celsii Hook. (62119; UCBG 49.2087-1; Mexico, Huntington Botan- 
ical Garden #20.128; UC). n = 30. This is also a diploid with 5L + 25s 


1964 | CAVE: AGAVACEAE 165 


pairs of chromosomes. Meiosis was regular (fig. 6). Capsule formation 


was heavy, but seed set was negligible. 
A. toumeyana Trel. (UCBG 62.254—1; Arizona, origin unknown; UC). 


This species is probably a polyploid, judging from the number of long 


.“ g3 ~ ’ 

. 8 @4 Drags Pol a 
ae “< Wy yPan 

ea 4 => = \ ov 


° = pee Yh ate 
& r @ g* & oe ne <%,° Py a é 
:* @ oo ~~ “r ta ce " ed 


*v 
qe: ot 


a7 6, tee OP 


e Fas 4 
@ " e =~ - ekg -. om 
Sa Pie, a 
ET an 
< a °° ome 
e Y od 8 e sy of : 


Fics. 1-9. Chromosomes of Agave species: 1, Agave sp. (Manfreda), I T, polar 
view, n = 30; 2, Agave sp. (Manfreda), I T, side view, n = 30; 3, Agave filifera, I M, 
n= 30; 4, A. lechuguilla, II] M, n=58; 5, A. victoria-reginae, I M, n= 30; 
6, A. celsti, diak., n = 30; 7, A. deserti, diak., n = 59; 8, A. salmiana, diak.; 9, A. 
asperrima, microspore mitosis, n = 87. All X 833. 


166 MADRONO [Vol. 17 


arms of chromosomes in II M. However, meiosis was so disturbed that 
no count was possible. Like other irregular meioses the development of 
PMC was not synchronous, but all stages of meiosis were found together 
in one portion of an anther. 


AGAVE, SUBGENUS EUAGAVE 


A. deserti Engelm. (6289; UCBG 52.1911-1; Baja California, Mexico, 
Hutchison 710; UC) n. = 59. This specimen had 10L + 49s chromo- 
some pairs (fig. 7). Meiosis was regular with 10 pairs of large chromo- 
somes and only little lagging at I A. No multivalent formation was noted. 
Many capsules developed to a large size, but only a very few mature 
black seeds with embryos were present. Seeds without embryos were the 
same size, but remained white. 

A. salmiana Otto. (62103; UCBG 61.1518-1; Mexico, origin unknown; 
UC, US, MO). This species was reported by Vignoli as quoted by Granick 
(1944) to be tetraploid. Meiosis in our plant was so disturbed that a 
count was impossible at this stage. Development of PMC was not syn- 
chronous, but each cell was likely to be at a different stage of meiosis. 
There were only 5 units that could be considered as associations of large 
chromosomes, but the number of small chromosomes was considerably 
greater than 25 (fig. 8). Whether these small units represented pairs 
or univalents was impossible to say. At I A many bridges and fragments 
or lagging small chromosomes were present. At the tetrad stage micro- 
nuclei were seen which persisted even after the young pollen grains were 
formed. In pollen grain mitoses 8—9 large chromosomes could often be 
made out, suggesting that the large units at I M were multivalents. The 
pollen grains varied in size and the exine might be said to be malformed, 
in that the pore was enormous, sometimes being about a fourth of the 
surface of the grain. The walls were thick and sculptured and the grains 
were often held together in pairs. At maturity some showed two male 
gametes, while others had only one, or even none, judging from the lack 
of stainability. 

A. asperrima Jacobi. (5909; UCBG 49.2095-1; Mexico, Huntington 
Botanical Garden #20.192; UC). n = 74-93. This plant had developed 
beyond the stage of meiosis in PMC before the present investigation was 
started. However, divisions in many pollen grains were studied. The 
pollen did not vary greatly in size and few empty grains were observed. 
The exine, though sculptured, was not as thick as in A. salmiana. In 27 
grains in which metaphase plates were counted, there were 16 large 
chromosomes, one of which was slightly smaller than the rest. In 22 of 
these plates the small chromosomes could be counted, and ranged in 
number from 58 to 77 (fig. 9). Counts of small chromosomes cannot 
be as accurate as those of large. The error should cause underestimation 
of the number, since it is fairly easy to miss some of the small chromo- 
somes. A. asperrima is therefore a hexaploid with one large chromosome 
extra. It would seem that the small chromosomes may segregate unevenly 


1964] CAVE: AGAVACEAE 167 


at meiosis without affecting the viability of the pollen to develop at least 
through the microspore division. Doughty (1936) reported a variable 
number of small chromosomes within each of the high polyploid species 


p° 0 J 22 
wa b, " 


'e “Ng Geo & 
‘Pps e se A > 
eg Nat 
Pp 
cS Pe 
0 
ni 4 ¢! 
rf e a 
,@ @ | 
e <5 & ) I 3 zt 
ae 
ae all 9 
8 ae 3° e % 


® per £4 
sos” t nth 
ey’ oo ites 
16 


Fics. 10-17. Chromosomes of Agave, Furcraea, Beschorneria, Nolina, and Dory- 
anthes: 10, Agave vexans, II T, n= 87; 11, Furcraea andina, diak., n = 30; 12, 
Beschorneria yuccoides, diak., n = 30; 13, Nolina parviflora, diak., n = 19; 14, N. 
bigelovii, diak.,n = 19; 15, N. bigelovii, I M, n= 19; 16, N. beldingii, diak., n = 19; 
17, Doryanthes palmeri, I M, n= 24. All X 833, except 14 and 16 which are X 770. 


168 MADRONO [Vol. 17 


studied by him. Sharma and Bhattacharyya (1962) report variation in 
somatic cells of a number of species in the number of small chromosomes 
present, and suggest that this irregular behavior may be an aid to 
speciation in the genus, particularly since various types of vegetative 
reproduction are common. Unfortunately nothing is known as to whether 
the mature pollen of A. asperrima is viable, or what sort of seed set 
occurred on this plant. 

A. vexans Trelease. (62132; UCBG 49.2089-1; Mexico, Huntington 
Botanical Garden #20.157B; UC, US, HEID). n = 87. This species is 
also a hexaploid. Meisosis was regular with respect to the large chromo- 
somes. Multivalents were rare in both large and small chromosome pairs. 
At I A occasional lagging small chromosomes could be seen, but tetrads 
were normal and there were no micronuclei. At II T (fig. 10) there were 
15 large chromosomes and 72 small. 

It is of interest that even though meiosis is regular and pollen is 
apparently good in many agaves, seed set may be negligible or entirely 
lacking under botanical garden conditions, at least, e.g., when there is 
only one plant of a species in bloom at one time. Widespread vegetative 
reproduction in the genus may favor the retention of self-incompatibility 
factors. 


FURCRAEA 


F. andina Trelease. (62104; UCBG 61.1490-1; South America, origin 
unknown; UC). n= 30. There were 5L + 25s pairs of chromosomes 
(fig. 11). No irregularities were observed. Pollen grains remained in 
tetrads. Seed was not set, but bulbils were produced. 


BESCHORNERIA 


B. yuccoides C. Koch. (6222; UCBG 57.384-S1; Chiapas, Mexico, 
MacDougall 377; UC, US, K). n = 30. The 5L + 25s pairs of chromo- 
somes were microscopically indistinguishable from those of Furcraea 
andina (fig. 12). The pollen grains were also held together in tetrads. 
Seed set was good, but no bulbils were formed as in the latter genus. The 
count agrees with Koeperich’s findings of 1930, although she reported 
12L + 48s chromosomes in somatic cells. One of the largest of the short 
chromosome pairs must have been considered as long chromosomes. 

Except in the genus Agave no polyploidy has been reported in any 
of the Agavaceae with the characteristic karyotype of 5L + 25s chro- 
mosomes. 


NOLINA 


N. beldingii Brandegee. (5505; Sierra de la Victoria, Baja California, 
Mexico, Carter & Ferris 3331; DS, UC). n= 19. The plants were all 
male and showed 19 pairs of chromosomes at diakinesis (fig. 16). 

N. bigelovii (Torr.) Wats. (5504; In-ko-pah Gorge, Imperial County, 
California; Pray s.n., in 1955); (6283; near Los Angeles Bay, Baja Cali- 
fornia, Mexico, Moran 9732; SD). n= 19. Nineteen pairs of similar- 


1964] CAVE: AGAVACEAE 169 


sized chromosomes were present in meiosis of microsporangiate flowers 
in both collections (figs. 14, 15). 

N. parviflora Hemsl. (6297; UCBG 51.1242-1; Mexico, origin un- 
known; UC). n= 19. This male plant has been growing in the Garden 
since 1951, having been moved from the main campus. At meiosis there 
were 19 pairs of chromosomes showing no great differences in size 

(fig. 13). 

Previous counts in the genus Nolina have varied. McKelvey and Sax 
(1933) counted about 38 chromosomes in somatic cells of an unnamed 
species, and Whitaker (1934) showed 36 chromosomes in the root tips 
of NV. recurvata. Lewis (1959) counted 20 pairs in NV. parryi, although 
Lenz (1950) pictured only 19 pairs in this species. Satd (1942) showed 
36 somatic chromosomes in NV. microcarpa. Gioelli’s count of 10 haploid 
chromosomes in JV. longifolia, as reported by Granick (1944) is remark- 
able in deviating distinctly from those reported for the rest of the genus. 


DORYANTHES 


D. palmeri W. Hill. (62151; UCBG 32.3895-1; Australia, origin un- 
known; UC). n= 24. This plant has been growing in the Garden since 
1932, and has been transplanted three times. An inflorescence appeared 
in the fall of 1962, and by January 1, 1963 the flowers were in the process 
of meiosis in the PMC. There were 24 pairs of chromosomes segregating 
regularly at I A. There was no great difference in size among the pairs, 
except that one was considerably larger than the rest (fig. 17). Sato 
(1938) depicts 4 large chromosomes in somatic cells. Although delimita- 
tion of microspores was “simultaneous” as in Phormium (the only other 
genus of the Agavaceae with this type of delimitation), the mature pollen 
grains were monosulcate like those in Yucca and Agave, and distinctly 
different from the trichotomosulcate grain of the former. 

In the genus Doryanthes three different somatic numbers have been 
reported: in D. excelsa, Newman (1929) found 44; in D. palmeri, Whit- 
aker (1934) counted 36; in both these species and also in D. guilfovylei, 
Sat6 (1938) counted 48. 

Department of Botany, University of California, Berkeley 


LITERATURE CITED 


BerceER, A. 1915. Die Agaven. Gustav Fischer, Jena. 

Braviey, M. V. 1948. A method for making aceto-carmine squashes permanent with- 
out removal of the cover slip. Stain Tech. 23:41—44. 

Cave, M. S. 1953. Cytology and embryology in the delimitation of genera. Chron. 
Bot. 14:140-153. 

. 1956. In Documented chromosome numbers of plants. Madrono 13:205- 

206. 

Dovucuty, L. R. 1936. Chromosome behavior in relation to genetics of Agave. Jour. 
Genet. 33:197-205. 

Granick, E. B. 1944. A karyosystematic study of the genus Agave. Am. Jour. Bot. 
31:283-298. 

Hutcumson, J. 1934. The Families of Flowering Plants. Vol. II. Monocotyledons. 
Macmillan, London. 


170 MADRONO [Vol. 17 


Koepericu, J. 1930. Etude comparative du noyau, des chromosomes et de leurs rela- 
tions avec le cytoplasme. La Cellule 39:307-398. 

Lenz, L. W. 1950. Chromosome numbers of some western American plants. I. Aliso 
2:317-318. 

Lewis, H. 1959. In Documented chromosome numbers of plants. Madrofo 15:49-52. 

McKetvey, S. D. and K. SAx. 1933. Taxonomic and cytological relationships of 
Yucca and Agave. Jour. Arnold Arb. 14:76-81. 

Moran, R. 1949. The Agavaceae. Desert Plant Life 21:64—69. 

Newman, F. V. 1929. The life history of Doryanthes excelsa. II. Proc. Linn. Soc. 
New S. Wales 54:411-435. 

Sato, D. 1935. Analysis of karyotypes in Yucca, Agave and related genera with special 
reference to the phylogenetic significance. Jap. Jour. Genet. 11:272-278. 

. 1938. Karyotype alteration and phylogeny IV. Cytologia. 9:203-242. 
. 1942. Karyotype alteration and phylogeny in Liliaceae and allied families. 

Jap. Jour. Bot. 12:57-161. 

SHaRMA, A. K. and U. C. BHATTACHARYYA. 1962. A cytological study of the factors 
influencing evolution in Agave. La Cellule 62:259-279. 

WHITAKER, T. W. 1934. Chromosome constitution in certain monocotyledons. Jour. 
Arnold Arb. 15:135-143. 

WUNDERLICH, R. 1950. Die Agavaceae Hutchinsons im Lichte ihrer Embryologie, 
ihres Gyn6ozeum, Staubblatt, und Blattbaues. Osterr. Bot. Zeitschr. 97:437-502. 


REVIEWS 


Flora of Illinois. By GEORGE NEVILLE JONES. 3rd ed. American Midland Naturalist 
Monograph 7. vi + 401 pp. Notre Dame, Indiana. 1963. $7.50. 


Jones’ Flora of Illinois will be familiar to most Midwestern botanists, as it has 
been for a number of years one of the only up-to-date identification manuals of a 
Midwestern state. Since Deam’s Flora of Indiana has been out of print for some 
years (apparently not to be reprinted in the near future) and the floras of Wisconsin 
and Michigan are still in preparation, there ought to be considerable local demand 
for the Flora of Illinois. 

The general format of the book follows that of the second edition (1950), with 
one major exception. The usual sequence of families, based on the system of Engler 
and Prantl, has been abandoned in favor of a modification of that of Hutchinson; 
a conspectus of this new classification is presented toward the back of the book 
(pp. 369-373). By and large, Jones appears to have followed the arrangement in 
such Hutchinsonian works as Clapham, Tutin, and Warburg’s Flora of the British 
Isles, but he has introduced some innovations of his own. Unfortunately, some of 
these modifications are highly questionable, at least if any significance at all is to 
be placed on the linear arrangement of families. Some of the more debatable assign- 
ments in the conspectus include: 1. The Violaceae and Cistaceae are placed in the 
Papaverales, apparently because of their parietal placentation. As far as I can de- 
termine, this is the first time that anyone has ever circumscribed the Papaverales 
in such a manner. 2. The inclusion of the Lauraceae in the same order with the 
Lythraceae, 60 families away from the Ranales, is baffling. This assignment is not 
likely to encourage Hutchinson to claim Jones as a disciple! 3. The Callitrichaceae 
are placed in the Myrtales between the Haloragaceae and Hippuridaceae. These three 
families do grow in wet places and tend to have reduced flowers, but there is no good 
evidence that the Callitrichaceae are any closer to the Hippuridaceae than they are 
to the Euphorbiaceae (where they were misplaced in the Engler system). 4. The 
Cactaceae are placed between the Passiflorales and Loasales, despite the fact that 
much biochemical and anatomical evidence demonstrates their affinity to the Centro- 
spermae (Chenopodiales of Jones). 5. The Aristolochiales are placed after the 
Myrtales rather than the Ranales. 6. Jones unaccountably rejects one of Hutchin- 


1964] REVIEWS 171 


son’s better innovations and separates Yucca and Agave into different orders; this 
disregards cytological and anatomical evidence that they are fairly closely related. 

Also debatable, but admittedly a matter of opinion, is the consistency of treat- 
ment in delimitation of groups. If a “foolish consistency is the hobgoblin of little 
minds,” still one expects a reasonable uniformity of circumscription. Thus Jones on 
the one hand accepts the dissolution of the Nymphaeaceae into three families (as 
Li has unconvincingly proposed) and maintains Menyanthaceae distinct from Gen- 
tianaceae, but on the other hand includes the Monotropaceae and Pirolaceae within 
the Ericaceae. The Saxifragaceae are split up 4 la Hutchinson into a number of 
families, but the Taxaceae are included in the same order with the Pinaceae. In 
general, Jones has tended to split supraspecific taxa rather finely; but here he at 
least has a lot of distinguished company. 

The author implies in the introduction that the taxonomic treatments have been 
brought up to date by consulting the latest monographs and revisions. It is surpris- 
ing, therefore, to find no reference to Haucke’s important work on Equisetum subg. 
Hippochaete. The authorities cited for the two species of Poinsettia are incorrect. 
The bitterweed is listed as Helenium tenuifolium Nutt., although the correct name 
was shown in 1957 to be H. amarum (Raf.) Rock. However, it does appear that on 
the whole the nomenclature of the book is reliable. 

Lest it appear that I am being too captious, it ought to be pointed out that the 
objections listed above do not seriously detract from the utility of the book. The 
author (p. 2) specifically states that “the main objective of this work is... to 
afford a ready means of identification of the approximately 2400 species of flowering 
plants and fernworts growing without cultivation in Illinois.” It is only fair, there- 
fore, to judge the book primarily by this standard. The physical lay-out of the 
volume is attractive, and the contents are logically arranged. The difficulty to the 
user may come as a consequence of the telegraphic style. As in previous editions, there 
are no separate descriptions of the taxa; the information is all presented in synoptic 
form in the leads of the keys. This works rather well when a species is being keyed 
down within a genus, since each species usually has a reasonably detailed diagnosis 
in the key. However, the characteristics of genera and families often have to be 
reconstructed from successive lines in a key, and this may give trouble to those (i.e., 
most of us!) who have short memories. For a common plant such as Cichorium 
intybus, for example, there is no description on the page where it is listed; and even 
in the key there is no indication of the characteristics of leaves (except that they are 
probably alternate), general habit, or involucre. In practice, such brevity will not 
seriously inconvenience the professional taxonomist or experienced amateur, who 
will in fact find identifications expedited by the uncluttered format. However, many 
beginning students in plant taxonomy may run into serious problems and need some 
supplementary reference with full descriptions. Perhaps part of the difficulty is that 
the book is mistitled; it should not be called Flora of Illinois but perhaps rather 
“Keys for Identification of the Flora of Illinois.” 

Judged as a manual of keys for ready identification, Jones’ “flora” stands up well 
indeed. Perhaps, having seen that a little modernity can be a dangerous thing, he 
will in the next edition abandon the capriciously restyled sequence of families in 
the present book and either return to the hackneyed but convenient Engler and 
Prantl system or else take the trouble to carefully modify some new svstem to the 
needs of the Illinoian flora—-Grapy L. WesBsTER, Purdue University, Lafayette, 
Indiana. 


Flora of Our Sierran National Parks. By SAMUEL J. PUSATERI. 170 pp., 177 line 
drawings, 26 black & white photos, 11 plates (66 color photos). Carl & Irving Printers, 
Tulare, California, 1963. Available from the author at Red Bud Acres, Three Rivers, 
California. $3.75, paper-back ; $4.75 cloth. 

The aim of Pusateri in presenting this book on the plants of a portion of the 
Sierra Nevada of California is “to bridge the gap which exists between the many 


172 MADRONO [Vol. 17 


publications covering the more common species of the region and the advanced texts 
which are difficult to use for the person without considerable training in botany.” 
The area covered is Sequoia and Kings Canyon National Parks as well as Yosemite 
National Park. 

Ferns, grasses, sedges, rushes, and many other plants are omitted. The inclusion 
of taxa from beyond the limits of the parks is confusing as ranges of distribution 
are ignored. Mention of some species (Adenostoma sparsifolium, Chamaesaracha 
nana, Eschscholzia elegans, Oxytheca perfoliata, Pedicularis densiflora, Penstemon 
centranthifolius) which must be excessively rare or nonexistent in the region covered, 
should have been documented with herbarium vouchers. Anisocoma acaulis, normally 
expected east of the Sierra and in Kern Valley to the south, is said to occur along 
the western approaches to Sequoia and Kings Canyon National Parks. 

The sequence of families follows no known order and many plants (Aesculus, 
Cornus nuttalli, Dicentra, Eriogonum, Eschscholzia, Iris, Kelloggia, Mahonia, 
Platystemon) cannot be identified because of ambiguities in the keys. Some other 
species are keyed but not described or described but not keyed. Pusateri has relied 
upon Jepson’s A Manual of the Flowering Plants of California (1925) “as the final 
authority for most of the scientific names... .” 

The black and white photos scattered through the text are excellent. The species 
shown in 21 of the 66 color photos also are represented by line drawings even 
though common plants (corn lily, chinquapin, poison oak, mountain misery, cow 
parsnip, arrowleaf groundsel (arrowhead butterweed), common bracken, staghorn 
lichen) are not illustrated. About 38 per cent of the species included are illustrated 
and these illustrations probably will be helpful to those who have no knowledge 
of plants since almost nothing else is available of local coverage for the Sequoia and 
Kings region.—WALLace R. Ernst, Smithsonian Institution, Washington, D.C. 


NOTES AND NEWS 


A New Locality FoR ASPLENIUM VESPERTINUM.—The fern Asplenium ves perti- 
num Maxon has been reported for the San Gabriel Mountains, the San Bernardino 
Mountains, and from San Rafael, Baja California. Its appearance in the more north- 
erly Santa Monica Mountains may indicate that it is more widespread than formerly 
suspected. A very small single colony was found on a rock outcropping at Sherwood 
Lake, Ventura County, California, in March, 1963 under an overhanging sandstone 
rock ledge with a northern exposure (Joe s.n., Oct. 20, 1963, DS, LA). The ferns, 
which were but a few inches tall, were growing in moist rather silty soil and pro- 
tected in front wtih clumps of Dryopteris arguta. Adiantum jordani, Dryopteris 
arguta, and Pityrogramma triangularis were abundant in the vicinity. Less common 
were Cheilanthes californica and Polypodium californicum.—BaRBARA JOE, University 
of California, Los Angeles, and Los Angeles City College. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volumes], 1916-1929) wy Vine, ee $5.00 
Volume IT, 1930-1934 . . . . . 5.00 
Volume ITI, 1935-1936 ee me A, 0) 
Volume IV, 1937-1938 .°.0. 7. 3 5 95200 
Volume V, 1939-1940 . . . . . ~ 5.00 
Volume VI, 1941-1942 he cn Cee SeOO 
Volume VII, 1943-1944 Fee os 5.08 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 5. Wake Ger yee ORO 
Volume X, 1949-1950) 3 3). re 00 
Volume XI, 1951-1952 Seah Ga) OL OW 
Volume XII, 1953-1954 . . . . 8.00 
Volume XIII, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 a a O08 
Volume XVI, 1961-1962 . . . ~~ . 12.00 


Single numbers: 


Volumes:l and Il... 725.) ae eee 50 
Volumes II through XIV. . .. 1.00 
Volume XV <6) a7 Ss) Ai ee 1.25 
VolumieXVio8 . 3 hese eee 1.50 


The subscription price of MaproNo is $6.00 per year. 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 membership in the California Botanical Societv on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


MADRONO 


VOLUME 17, NUMBER 6 APRIL, 1964 


Contents 


PAGE 


CYTOTAXONOMY AND DISTRIBUTIONAL ECOLOGY OF 
WESTERN NorTH AMERICAN VIOLETS, 
Jens Clausen 173 


NOMENCLATURAL PROBLEMS IN THE ACACIA CORNI- 
GERA ComPLEX, Velva E. Rudd 198 


NOTES ON THE LEAF EPIDERMIS AND CHROMOSOME 
NUMBER OF SWALLENIA (GRAMINEAE), 
Dennis Anderson 201 


NOTES AND NEws: NEw DISTRIBUTION RECORD FOR 
HELEOCHLOA ALOPECUROIDES IN OREGON, 
William H. Baker 197 


SAXIFRAGA ESCHSCHOLTZII STERNB., Kenion L. 
Chambers; NEw PUBLICATIONS 203 


A WEST AMERICAN JOURNAL OF BOTANY 


\ 


UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS | 


EpcAR ANDERSON, Missouri Botanical Garden, St. Louis 
LymMAN Benson, Pomona College, Claremont, California 
HERBERT F, CopELAND, Sacramento College, Sacramento, California 
Joun F. Davinson, University of Nebraska, Lincoln 
WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. 
Mitprep E. Martuias, University of California, Los Angeles 
ROBERT ORNDUFF, University of California, Berkeley 
Marion OwnBeEy, Washington State University, Pullman 
REED C. RoLiins, Gray Harbarium, Harvard University 
IrA L. Wiccrns, Stanford University, Stanford, California 


Editor—JoHN H. THomAs 
Dudley Herbarium, Stanford University, Stanford, California 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: G. Ledyard Stebbins, Department of Genetics, University of California, 
Davis. First Vice-President: Annetta Carter, Department of Botany, University of 
California, Berkeley. Second Vice-President: Carl W. Sharsmith, Department of 
Biological Sciences, San Jose State College. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding 
Secretary, Margaret Bergseng, Department of Botany, University of California, 
Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State 
College. 


1964] CLAUSEN: VIOLA Wf) 


CYTOTAXONOMY AND DISTRIBUTIONAL ECOLOGY OF 
WESTERN NORTH AMERICAN VIOLETS 


JENS CLAUSEN 


During the years from 1932 to 1943 the chromosome numbers of vio- 
lets from many natural habitats throughout western North America were 
determined. In some species determinations were made on plants from 
many populations. The present paper is a report on these studies. They 
were conducted in cooperation with my friend, the late Milo S. Baker of 
Santa Rosa Junior College, who made many trips through the western 
states and fixed plants in the wild when they were in early blooming 
stage. He also brought plants back to his garden at Kenwood, Sonoma 
County, California, and made other fixations there. Through many yeers 
this garden contained a remarkable collection of Viola species from out- 
of-the-way places which he treated with tender care. 

Other western violets were collected and bud materials fixed by myself 
and David D. Keck on collecting trips throughout the western states, 
especially California. In the early years of the study some of these col- 
lections were grown in the greenhouses at the Department of Plant Biol- 
ogy of the Carnegie Institution of Washington. It soon became evident, 
however, that most of the wild species of the Chamaemelanium section 
of Viola were ill-adapted for cultural practices and were difficult to keep 
alive in the experiment garden at Stanford. Even in Baker’s more favor- 
able garden at Kenwood the Chamaemelanium violets were unable to sur- 
vive for a longer period than two to four years. For these reasons the 
Viola project was primarily limited to the approaches of cytotaxonomy 
and distributional ecology. 

Baker and I cooperated on the major taxonomic classification which 
was greatly aided by the chromosome numbers, by careful field studies 
and by study of specimens from about 14 herbaria throughout the coun- 
try. The specimens were pooled for the study and annotated through 
1936-1938. A complete list of these specimens was prepared and sup- 
plemented by impressions of leaves and flowers of critical specimens. The 
list and impressions were of great help in the study of the distributional 
ecology and morphology of the taxa and is deposited as documentation 
at the herbarium of the University of California, Berkeley. 

It was agreed that Baker should write the taxonomic descriptions pro- 
viding the nomenclature. The taxonomic papers (Baker, 1935-1960) 
were to be followed by a paper on the chromosome situation and its im- 
plications for classification. 

At the 1935 meeting of the American Association for the Advancement 
of Science at St. Louis, Missouri, I gave a paper on the chromosomal 
distributional situation in the Purpureae and the Nuttallianae of the 
Chamaemelanium section. Eventually Baker (1949d) provided a tabular 


Maprono, Vol. 17, No. 6, pp. 173-204. June 11, 1964. 


174 MADRONO [Vol. 17 


list of the chromosome numbers of the various taxa, mentioning that the 
cytological work had been done by me. The proper bibliographical cita- 
tion of these chromosome numbers is therefore Baker (1949). 

In 1951 most of the chromosome numbers of the species were listed 
and diagrammed (Clausen, 1951), but the numbers were never docu- 
mented by reference to the herbarium sheet or the locality where the 
plant was taken. 

The purpose of the present paper is to provide the documentation and 
to integrate the field evidence with the cytological information and with 
the findings on the close relatives of our western violets from other parts 
of the world investigated by authors such as Gershoy and Miyaji. Ger- 
shoy in 1936 provided me with information on the sources of some of 
the plants he used in the determination of the chromosome numbers, and 
this information is also included in the listing (table 1). Unless other- 
wise stated the chromosome determinations are based on fixations made 
of bud materials from the plants growing in their natural habitats. When 
proper meiotic stages were unavailable mitotic divisions in the floral 
regions were successfully used for the counting of chromosomes. 


CLASSIFICATION AND EVOLUTIONARY PATTERNS 


In the chromosome list (table 1) the taxa are arranged alphabetically 
under sections, subsections (cenospecies), species, and subspecies. In the 
text the arrangement is ecologic-geographical, combined with cytological 
evidence in order to provide a background for an evolutionary under- 
standing. 

The genus Viola has many natural hybrids, mostly sterile. Some of 
these are discussed later. A study of these revealed some of the evolution- 
ary patterns. The violets have utilized the methods of polyploidy in 
their processes of speciation. It was therefore no accident that the first 
example of polyploidy within a genus (n — 6, 12, 18, 24, 36) was in an 
early paper on chromosome numbers in violets (Miyaji, 1913). Written 
in Japanese and generally unknown to the rest of the scientific commu- 
nity, it was later translated to German (Miyaji, 1927). 


SECTION CHAMAEMELANIUM 


Within western North America the Chamaemelanium violets contain 
several evolutionally ascending species complexes (Clausen, 1951). These 
species complexes (cenospecies) are here considered subsections and 
are listed as Pedunculatae, Purpureae, Nuttallianae, Chrysanthae, Nudit- 
caules, Canadenses, and Biflorae, the latter a less closely knit taxon. 
Species belonging to such distinct subsections can occur together within 
one and the same habitat; in contrast, the taxa within a single subsec- 
tion occupy geographically or ecologically separate regions. The taxa 
within the subsection are therefore either ecotypic subspecies of one 
species, or they are ecospecies within a species complex. Experimental 


1964] CLAUSEN: VIOLA 1/5 


evidence to determine between these two possibilities is not available ex- 
cept through natural hybrids. 

At a nontropical latitude it is unusual to find so many distinct species 
complexes existing within a geographic region, as in Chamaemelanium. 
The section is probably old, and the complexity mirrors evolutionary 
histories within a region that during millions of years has been geologi- 
cally unstable. Obviously, many links with the past have been lost, al- 
though some still exist. The bridges between complexes are provided by 
species of recent discovery. 

CHANGES IN CLASSIFICATION. The classification used in this paper 
differs slightly from Baker’s. One of the changes (Baker, 1953) is that 
V. aurea ssp. aurea and ssp. mohavensis are here considered to be sub- 
species of V. purpurea. This is in accord with Baker’s earlier concept and 
avoids the unnatural arrangement whereby V. aurea occurs as an island 
within the geographical territory of V. purpurea. 

In contrast, V. linguifolia is retained as a species instead of being con- 
sidered a subspecies of V. praemorsa. The morphological characters are 
not highly distinct in this case, although both the leaves and flowers of 
the two are different. There is reason to believe that the hexaploid V. 
linguifolia does not occur in the states of the Pacific slope. Forms from 
these more westerly regions previously thought to be V. linguifolia have 
consistently proved to be octoploid and belong to V. praemorsa ssp. 
oregona and ssp. major. 

Viola praemorsa ssp. praemorsa, n = 18, from the coastal prairies, and 
24 paired V. praemorsa ssp. major and ssp. oregona are retained as one 
species in accordance with Baker’s classification. However, I do not feel 
that there is justification for maintaining V. praemorsa ssp. arida because 
ssp. major itself comes from the same transmontane arid region. 

Another uncertainty exists in V. bakeri ssp. shastensis Baker. This 
taxon was referred to V. bakeri because it was thought to be octoploid, 
n — 24. Baker (1960) cited several specimens he thought belonged to 
the new taxon. A closer study of the type specimen, Baker 13045, from 
Postpile Camp in the Yolla Bolly Mountains, Tehama Co., and of Baker 
13139 from Scott Mountain, Trinity Co., on which the chromosome num- 
ber was determined, indicates that they belong to different species. The 
type specimen has an elongated taproot and broadly lanceolate leaves 
with entire ciliate margins, scattered shaggy leaf pubescence, and pubes- 
cent capsules, the latter two characters separating it from the octoploid 
V. bakert but placing it with the diploid relict V. tomentosa. The plant 
fron. Scott Mountain has an abbreviated truncate rootstock and crenu- 
late leaf margins, placing it with the octoploid V. praemorsa ssp. oregona 
(table 1). The taxonomic status of the rare taxon, V. bakeri ssp. shas- 
tensis, and its chromosome number are therefore unknown. Most likely 
the taxon represents a disjunct population of the rare V. tomentosa. 

SUBSECTION PEDUNCULATAE. The most coastal and southern of the 
North American Chamaemelanium species is Viola pedunculata, n= 6 


176 MADRONO [Vol. 17 


(table 1). This is the johnny-jump-up or wild pansy of California, which 
is remarkably adjusted to the California Coast Range climate of a mild, 
humid winter followed by a dry summer. It is a geophyte, having a deep- 
seated digitate roostock or bulb. During the winter and spring multi- 
tudes of underground stolons develop from the bulb, terminating into 
patches of flowering stems (Baker, 1949d). The stems dry up at the 
onset of summer, the plant becoming summer-dormant. This species occu- 
pies primarily the Coast Range regions of the southern half of California, 
including some offshore islands, and the northern part of Baja California. 
Viola pedunculata ssp. tenuifolia (table 1) occurs in scattered colonies 
in the inner Coast Ranges and the foothills of the Sierra Nevada. 


Viola charlestonensis (Baker, 1949c) is a narrow endemic, high alti- 
tude counterpart limited to the Charleston Mountains of southeast Ne- 
vada. It is also diploid (table 1) and resembles V. pedunculata in the 
rounded outline of the leaves, its deeply buried rootstock and its large 
seeds. 


The scattered populations of the Pedunculatae suggest that this type 
of a violet previously had a wider distribution. Possible Mexican rela- 
tives are V. flagelliformis Hemsley and V. galenaensis Baker (Baker, 
1947), the latter at 9000-12000 ft in the state of Nuevo Leon. 

SUBSECTION PURPUREAE, AN EXAMPLE OF ECOLOGIC-GENETIC SPECI- 
ATION. Viola purpurea and its close relatives occupy most of the area 
from the Pacific to the Rocky Mountains, attaining an altitude of 11000 
ft, but do not enter the prairie states. Viola quercetorum is cohabitant 
with V. pedunculata but is separated from it through its higher number 
of chromosomes, n = 12. 


Leaf shapes of the various taxa within the Purpureae are pictured in 
Baker (1949d). Gross silhouettes of herbarium specimens of the succes- 
sion of taxa are on a profile of western North America in Clausen (1951). 
West of the summits of the Sierra Nevada and the Cascades the taxa of 
this subsection have well developed aerial stems from root crowns in the 
surface of the ground; east of the summits the stem nodes are abbre- 
viated, and the taxa native to the dry desert areas are geophytes having 
their renewal buds buried below the ground level. 

One diploid species, V. purpurea, n = 6, extends from the 3000 ft level 
on the west side of the Sierra Nevada eastward to the Rocky Mountains 
and has developed about 10 distinct subspecies which replace each other 
through a series of climatic zones and edaphically distinct habitats. 


1. Viola purpurea ssp. purpurea occupies the transition zone of Cali- 
fornia from about 3000-6000 ft, being associated with Pinus ponderosa, 
Libocedrus decurrens, and Quercus kelloggii. It is also associated with 
these species in the high mountains of southern California and in the 
northern California Coast Ranges (table 1). It has a tap root, rounded 
basal leaves with a truncate base, and aerial stems 10-15 cm long devel- 
oping from the ground level. The type was collected by Kellogg near the 


1964] CLAUSEN: VIOLA Lid 


present day Placerville but was presumably destroyed in the San Fran- 
cisco fire of 1906. 

2. In the Sierra Nevada the change from ssp. purpurea to ssp. meso- 
phyta coincides more or less with the replacement of Pinus ponderosa by 
P. jeffrevi and of Quercus kelloggii by Q. vaccinifolia. Viola purpurea ssp. 
mesophyta continues, however, throughout the territory of Pinus mur- 
rayana, following it to the summit of the passes at 10000 ft (table 1). 
It differs from ssp. purpurea by having shorter, more slender stems and 
distinctly narrower deltoid pubescent leaves. 

3. The former two subspecies have finely serrate leaf margins, but 
north of the Lake Tahoe region and still on the west side of the moun- 
tains ssp. mesophyta is gradually replaced by ssp. integifolia, which dif- 
fers from mesophyta primarily by having nearly entire leaf margins 
(table 1). 

4. Viola purpurea ssp. xerophyta occupies gravelly terraces on the east 
side of the Sierra Nevada at or above 10000 ft (table 1). These tiny 
plants have stems buried in the gravel, renewal buds below the surface, 
and narrow, highly pubescent leaves. This subspecies is clearly related 
to the neighboring ssp. mesophyta and transitions are common near the 
passes. Viola purpurea ssp. xerophyta extends southward from the Lake 
Tahoe region to the San Bernardino Mountains of southern California. 

5. Viola purpurea ssp. geophyta (table 1) is common in volcanic ash 
deposits at 4000-6000 ft east of the Cascades of central Oregon to Lassen 
Co., California, reappearing in the volcanic ash deposits of Mono Co. 
east of the Sierra Nevada. Its stems are deeply buried in ash and sand, 
the leaves are gray-puberulent and sinuate-toothed. 

6. An interesting transitional ssp. dimorpha occupies a triangle in the 
northeastern corner of California (table 1) between ssp. integrifolia and 
ssp. geophyta. Its spring leaves are mesic, resembling the leaves of the 
Great Basin ssp. atriplicifolia, but unlike other subspecies from east of 
the Cascades and Sierras ssp. dimorpha develops aerial stems in the 
summer. 

7. The canescent V. purpurea ssp. aurea inhabits even drier zones than 
those inhabited by ssp. geophyta. It occurs locally east of the central 
Sierra Nevada in Washoe to Ormsby counties, Nevada, at about 4000 ft 
(table 1). Its stems and renewal buds are deeply buried in dry gravel. 


8. Viola purpurea ssp. mohavensis (table 1) is morphologically related 
to ssp. aurea. Its herbage is gray, varying from smooth to short-puberu- 
lent, and like aurea its leaves are sinuate-toothed. It occupies dry sites 
in the higher parts of the Mohave desert and montane areas in southern 
California, 4000-7000 ft altitude. The lower parts of its stems are buried, 
but it develops about 10 cm long aerial stems. 

Two subspecies occupy montane to alpine altitudes on the ridges of 
the Great Basin. Both are dwarfish and have green herbage, varying from 
smooth to slightly pubescent. Their renewal buds appear at the surface of 


178 MADRONO [Vol. 17 


the ground and aerial stems are either absent or very short. They are: 

9. Viola purpurea ssp. atriplicifolia has leaves of rounded outline with 
deeply sinuate leaf margins. It occurs from 6000-10000 ft altitude at 
widely scattered localities from eastern Washington, Oregon and Cali- 
fornia to Idaho and Wyoming. 

10. Viola purpurea ssp. venosa is a dwarf alpine with subcordate and 
subentire leaves growing from eastern Washington, Oregon, and western 
Nevada to Montana, Utah, and Colorado. 

Field and herbarium studies indicate that moderate recombinations 
of characters occur at the points where these taxa meet, suggesting some 
gene interchange. Such observations provide evidence that these taxa 
are subspecies adjusted to life in their particular environments rather 
than distinct species. Nevertheless, even though local crossings happen, 
the trends in subspecific replacements are fairly sharp, and the extremes 
are highly distinct. It is reassuring to notice how the characteristic leaf 
shape of ssp. atriplicifolia has been faithfully transmitted between points 
as remote as Yellowstone Park (type locality) and valleys on the east 
side of the Sierra Nevada. 

An interesting example of the contiguous existence of the subspecies 
of V. purpurea from diverse areas is the Slate Creek Valley in the Harvey 
Monroe Hall Natural Area surrounding the Timberline transplant sta- 
tation of the Carnegie Institution of Washington. Within this east-facing 
hanging valley at 10000-11000 ft altitude, ssp. mesophyta occupies south 
slopes in the Pinus murrayana forest, ssp. xerophyta gravelly east-facing 
terraces at 10400 ft on Mount Conness, and the morphologically highly 
distinct ssp. atviplicifolia was found in grassy areas in the bottom of the 
valley at about 10000 ft. The three subspecies have therefore preserved 
their morphologic and ecologic identities over great distances. 

Two tetraploid species, n = 12, are included in the Pur pureae complex. 
Both of the tetraploids occupy environmental regions outside of the 
regions inhabited by the subspecies of the diploid V. purpurea. Geo- 
graphically, therefore, these tetraploids are arranged as true members 
of the subsection Purpureae. Both tetraploids probably are amphiploids 
that combine diploid taxa belonging to separate subsections. The reasons 
for this assumption are to be discussed later in the paper. 

Viola quercetorum, n = 12 (table 1) is a species of the oak savannas 
of the California Coast Ranges and the Sierra Nevada foothills, extend- 
ing into the southern Oregon Coast Ranges (Baker, 1949a). In many 
characters it resembles the diploid V. purpurea ssp. purpurea of the Sier- 
ra Nevada transition zone and ssp. mohkavensis of the arid transition 
zone at montane altitudes in southern California. In other respects, such 
as its semi-woody, much-branched rootstock, its long-pedicelled subcor- 
date leaves, long-peduncled large flowers and generally large size, V. quer- 
cetorum reminds of the equally diploid V. pedunculata with which it is 
often cohabitant. Viola quercetorum follows Quercus douglasu and Pinus 
sabiniana, but above the 3000 ft level where these trees are being re- 


1964] CLAUSEN: VIOLA 179 


placed by Quercus kelloggi and Pinus ponderosa, V. quercetorum is also 
replaced by the diploid V. purpurea ssp. purpurea. 

Viola utahensis, n = 12 (table 1), is the other tetraploid of the Pur- 
pureae. It is a fairly narrowly endemic species in moister sagebrush lands 
of central Utah (Baker, 1949b). Occasionally V. utahensis is found to- 
gether with the putative parents. The Utah violet combines subsections 
Purpureae with Nuttallianae, whereas V. quercetorum possibly is a Pur- 
pureae-Pedunculatae combination. 

SUBSECTION NUTTALLIANAE, DIFFERENTIATED THROUGH POLY- 
PLOIDY. In contrast with the Purpureae the Nuttallianae constitute a 
polyploid species complex. The chromosome number of its species range 
from n= 6 to 24 at intervals of six pairs of chromosomes. The species of 
the Nuttallianae cover western North America from the coastal plains 
of the Pacific in Washington to the western prairie states east of the 
Rocky Mountains, although they are absent in the Coast Range regions 
from central California southward. 

With one exception, the species of the Nuttallianae differ from the 
Purpureae by having glabrous seed capsules compared with appressed 
puberulent capsules in the Purpureae. The Nuttallianae generally lack 
elongated aerial stems, and their basal leaves (Baker, 1957; Clausen, 
1951) are much larger and of a different shape than those of the Pur- 
pureae, Since the members of the two subsections mostly cover the same 
area they often cohabitate. 

The Nuttallianae subsection has two basic diploid species, each with 
its morphologically distinct series of polyploid derivatives. The diploids 
are geographically separated: one, in the central Sierra Nevada has tap- 
root and entire leaves with ciliate margins, and the other, in the inter- 
mountain region, has abbreviated, truncate rootstock and crenulate, non- 
ciliate leaf margins. 

1. Viola tomentosa, n -= 6 (table 1), is the Sierra Nevada diploid, a 
recently discovered rare endemic at 5000-6000 ft from Eldorado to Plu- 
mas counties (Baker, 1949b). It is a highly distinctive violet with a 
taproot similar to that of V. purpurea and small, lanceolate leaves 
(Baker, 1949d). It differs from others of the Nuttallianae by having 
densely tomentose leaves, woolly pubescent capsules and fairly well 
developed stems in the fruiting stage (Baker, 1957). It combines, there- 
fore, characters of the Nuttallianae with those of the Purpureae and 
hybridizes with the equally diploid cohabitant V. purpurea, although the 
hybrids are sterile (see later). 

2. Viola bakeri, n= 24 (table 1), is the only polyploid species that is 
morphologically and geographically related to the diploid V. tomentosa. 
Before its chromosome number was known, V. tomentosa was even sus- 
pected of being a subspecies of V. bakeri. The octoploid V. bakeri has 
a long taproot and lanceolate leaves with entire, ciliate margins, but it 
has glabrate capsules and lacks the tomentose pubescence and the elon- 
gated summer stems that characterize V. tomentosa (Baker, 1957). Viola 


180 MADRONO [Vol. 17 


bakert is a montane species of forested areas, ranging at altitudes of 
5000-6000 ft from southern Washington through the Cascade Moun- 
tains of Oregon to the central Sierra Nevada where it surrounds V. 
tomentosa. 

No hexaploid species is known that with V. tomentosa could produce 
the octoploid V. bakeri, unless the extremely rare V. bakeri ssp. shastensis 
(Baker, 1960) should prove to be the missing hexaploid. Morphologi- 
cally, however, ssp. shastensis classifies as a geographically well sepa- 
rated, less tomentose population of V. tomentosa. It could also be a relict 
hybrid derivative of V. tomentosa with V. bakeri. 

3. Viola vallicola, n = 6 (table 1), is the inter-mountain diploid that 
presumably is one progenitor of a couple of polyploid derivations. It has 
a truncated vertical rootstock; thin, smooth ovate leaves with truncate 
base and crenulate, non-ciliate leaf margins (Baker, 1957). It is a species 
growing in grassy and open wooded areas in the mountains, among sage- 
brush with some moisture, and in grassy prairies. It has the widest dis- 
tribution of any species within the Nuwttallianae, ranging from eastern 
Washington, Oregon and Nevada to Alberta, the Black Hills of South 
Dakota, Wyoming and Colorado, altitudes 4500-7500 ft. 

4. Viola nuttallu, n = 12, is the only tetraploid species among the 
Nuttallianae, a prairie species that remains east of the continental divide. 
It is a geophyte, having a deep seated truncated rootstock with the re- 
newal buds below the surface, and lanceolate, puberulent leaves with 
crenulate margins. The morphology contradicts that V. nuttallii could be 
an autoploid of V. vallicola as suggested by Baker (1957). Chromosomes 
have been counted only on plants from Canada and Colorado (table 1), 
and undiscovered diploid V. nuttalli could exist somewhere within the 
large area of the species, extending from Arizona, eastern Colorado, 
Wyoming, Montana to Alberta, North and South Dakota, Nebraska, and 
Kansas. 

5. Viola linguifolia, n = 18 (table 1), is a montane to subalpine for- 
est violet of the Great Basin Ranges, 2500-10000 ft altitudes. It is a 
hexaploid member of the V. vallicola derivatives, possessing the key 
characteristics of the group; short, erect, truncate rootstock and crenu- 
late-margined, non-ciliate, erect elongated-ovate leaves; the floral ped- 
uncles do not surpass the leaves (Baker, 1957). The leaf shape suggests 
a compromise between the leaves of V. vallicola and V. nuttallu, but V. 
linguifolia is larger than either and occupies a moister, higher altitude 
zone than either. Viola linguifolia extends from eastern Washington, 
Oregon and western Nevada to Montana, Wyoming and Colorado. 

6. Another closely related hexaploid of this group, V. praemorsa ssp. 
praemorsa, n = 18 (table 1), occupies the coastal prairies of Washing- 
ton, Oregon, and northern Califonia at low altitudes. It differs from V. 
linguifolia in having divaricate leaves with cordate bases, and the flow- 
ers on the equally divaricate peduncles surpass the leaves (Baker, 1957). 
It is densely pubescent with long hairs. Douglas’s name, V. praemorsa, 


1964] CLAUSEN: VIOLA 181 


refers to the truncated rootstock that is so characteristic of the polyploid 
violets derived from V. vallicola and V. nuttallu. _.. 

A gap of at least about 150 mi separates the coastal V. praemorsa from 
the nearest recorded interior and essentially montane V. linguifolia; this 
gap is now occupied by the Cascade Mountains. It is tempting to suggest 
that before the rise of the mountains these two hexaploids were con- 
nected across the then low plains. 

The mountains and valleys of the Pacific states that now le between 
the coastal prairies and the inter-mountain region provide habitats for 
three octoploid Nuttallianae taxa, n = 24, that probably are conglom- 
erates of several origins. One of these, the tap-rooted V. bakeri, n = 24, 
was discussed in connection with the Sierra Nevada diploid, V. tomen- 
tosa, n = 6. Two other octoploids, relating to the inter-mountain V. val- 
licola diploid and to the hexaploids on both sides of the Cascades-Sierra 
Nevada, have the praemorsa type truncated rootstock and the crenulate, 
non-ciliate leaf margins and will be discussed below. 

7. Viola praemorsa ssp. major, n = 24 (table 1), is the largest and 
most robust of the taxa within the Nuttallianae. It differs from ssp. prae- 
morsa by its leaves equaling or surpassing the flowers, by generally erect 
leaves and peduncles and larger flowers (Baker, 1957). These charac- 
ters are fairly well correlated with the chromosome number and particu- 
larly with the geographic distribution. This taxon was first described by 
Hooker as a variety of V. nuttallit on the basis of Douglas’ specimens 
taken “under solitary pines on the dry, sandy soil of the Columbia,” pre- 
sumably somewhere near the present Hood River, Bingen, and the Dalles. 

Viola praemorsa ssp. major occurs from Washington and central Cali- 
fornia in a broad band through the Cascades and the Sierra Nevada with 
extensions to neighboring mountains and intervening valleys at altitudes 
from about 600-6000 ft. The border between V. praemorsa ssp. major 
and V. linguifolia follows closely the dividing line between Washington- 
Oregon (in ssp. major territory) and Idaho (in V. dinguifolia territory). 
There is a westward bulge of ssp. major into the northern California 
Coast Ranges and the Siskiyou Mountains of California and Oregon, 
where it meets ssp. praemorsa. 

8. Viola praemorsa ssp. oregona, n = 24, has broadly lanceolate basal 
leaves, resembling V. dinguifolia. It occupies a small area at mid-altitude 
in southeastern Oregon and northeastern California (Baker, 1957). 

The taxonomist has good reason to become confused by the octoploid 
“muddle” of V. praemorsa ssp. major, ssp. oregona, and V. bakeri from 
the same general territory. These taxa are less distinct than the diploids, 
and it appears that an octoploid complex species is in the process of 
emerging through genetic interchange between the elements. The indi- 
viduals within each population are highly variable, and the populations 
differ from each other. Such differences go unnoticed until the herbarium 
sheets from each local population have been gathered from the various 
herbaria throughout the country. 


182 MADRONO [Vol. 17 


From an over-all point of view it appears that V. praemorsa ssp. major 
is morphologically related to hexaploid ssp. praemorsa on the coast, 
whereas ssp. oregona is suggestive of the hexaploid V. lingutfolia to the 
east, but no present-day diploids are known that with the hexaploids 
could synthesize the octoploid subspecies. Hexaploid and tetraploid pro- 
genitors of V. bakeri are also missing. The present chromosome survey 
is very limited, providing only a skeleton to which substance can be 
added. 

SUBSECTION CHRYSANTHAE. The yellow-flowered violets of the Chry- 
santhae have rosettes of deeply dissected leaves, deep seated rootstocks, 
and long-peduncled flowers on short aerial stems. They constitute a 
strictly polyploid complex through the states of the Pacific slope. With 
the species of both Purpureae and the Nuttallianae they produce sterile, 
natural hybrids. 

Viola sheltonti, n = 6 (table 1), is the only known diploid species 
within this subsection. It has palmatisect leaves of cordate outline and 
hybridizes with V. tomentosa of the Nuttallianae. 

Viola beckwithi T. & G. and V. trinervata Howell have leaves of simi- 
lar outline to those of V. sheltoni, although they are palmatifid rather 
than palmatisect. They are native to the northern Great Basin, but their 
chromosome numbers are not known. 

Viola douglasii and V. hallii have bipinnatifid leaves of ovate outline. 
Viola douglasii is native to the inner Coast Ranges and foothills of the 
Sierra Nevada from Kern Co. to Oregon. It is tetraploid, n = 12, in its 
southern half, and octoploid, n = 24, north of the San Francisco Bay 
(table 1). It hybridizes with species of the Purpureae and the Nuttal- 
lianae. 

Viola hallu, n = 36 (table 1; Gershoy, 1932) is a 12-ploid species of 
the northern Coast Ranges. 

The chromosome numbers of n = 18 and n = 30 are still missing in 
completing the polyploid series within this subsection, but the two Great 
Basin species above might provide them. 

SUBSECTION NUDICAULES. This subsection has erect succulent stems, 
usually poorly developed rosette leaves, and yellow flowers. The Nudt- 
caules extend to eastern North America and to east Asia and are inhabi- 
tants of fairly humid woods and forests. The diploid V. lobata, n — 6, 
and the only tetraploid, V. glabella, n = 12, are western species. A di- 
ploid close relative of the latter is the Japanese V. brevistipulata, n = 6 
(V. glabella in Miyaji, 1913). Other diploid relatives grow in eastern 
North America and in Japan (table 1). 

SUBSECTION CANADENSES. This subsection constitutes a complex of 
species that also have prominent and erect but leafy stems but have 
whitish to purple, rather than yellow flowers. The Canadenses have spe- 
cies both in western and eastern North America, including two diploid 
species, V. ocellata, and V. scopulorum, n= 6, and two tetraploids V. 
canadensis, and V. rugulosa, n = 12 (table 1). Bold and Gershoy (1934) 


1964] CLAUSEN: VIOLA 183 


found that the latter two intercrossed easily and their hybrids appeared 
fully fertile. The chromosome numbers of two western species of this 
complex V. cuneata and V. flettt are not known. 

SUBSECTION BIFLORAF. In a previous paper Clausen (1929) stated 
that it would be logical to consider the yellow-flowered V. diflora, n = 6, 
a Chamaemelanium violet. In certain respects this species is among the 
least differentiated species of the section and of the genus as well. In 1929 
the term Biflorae was applied only to V. biflora and V. sarmentosa (syn- 
onym: V. sempervirens). 

A better understanding of the Bzflorae is now possible. They consti- 
tute a loosely knit group of species that previously were scattered among 
the sections Dischidium, Chamaemelanium, and Plagiostigma. The Viola 
sections were based exclusively on the characters of the pistil and stig- 
ma. These make sensible distinctions between such sections as Melanium, 
Plagiostigma, and especially Rostellatae (Clausen, 1929) that were best 
known to European botanists. In contrast, within the Chamaemelanium 
section the species-to-species variability in pistil-stigma characters is 
greater than the section-to-section variabliity of the other violets (Clau- 
sen, 1929). 

As here interpreted, the Bzflorae follow the distinctive chromosomal 
pattern of the Chamaemelanium violets, have yellow flowers, and a cen- 
tral corm from which emerge either the flower-bearing but weaker hori- 
zontal stolons or rhizomes, or short ascending runners. The Bzflorae are 
inhabitants of humid woods, forests and mountains. 

The best known western species of the Biflorae is Viola sempervirens. 
Within the redwood forests of central California it is a tetraploid, n = 12 
(Clausen, 1929), but it is octoploid, n = 24, at lower altitudes in the 
forests of Oregon (Gershoy, 1934). The closely related V. orbiculata, 
n — 12 (Gershoy, 1932), is a species of higher altitudes in humid moun- 
tain forests of the northern Cascades and beyond; it has ascending, 
short runners. 

Another member of the Biflorae is V. rotundifolia, n = 6 (Gershoy, 
1928). This is a yellow-flowered violet of cool, rich woods in eastern 
North America. It was previously classified with the section Plagios- 
tigma on account of the shape of its pistil and stigma (Clausen, 1929), 
disregarding its flower color and chromosome number. Viola rotundifolia 
is reminiscent of the tetraploid western V. orbiculata by its gross mor- 
phology and by its short, rambling summer stems (Baird, 1942). 

The name-giving species of the complex is V. biflora, n = 6 (table 1). 
Because of its primitive pistil and split stigma it was previously referred 
to the monotypic section Dischidium, but this character is also found in 
other genera of the family. The pistil of V. sempervirens represents a 
transition from that of V. diflora (Clausen, 1929). Viola biflora has a 
short, thick rootstock from which rambling, deciduous flowering stems 
emerge; these contrast with the evergreen, trailing, flower-bearing stems 
of V. sempervirens from a more southern latitude. 


184 MADRONO [Vol. 17 


Viola biflora is a montane and high latitude circumpolar species that 
reaches western North America in Alaska and also occurs in isolated 
locations within the Pikes Peak region of Colorado. Gershoy (1934) 
produced a weak hybrid between V. biflora, n = 6, and V. rugulosa, 
n — 12, an indication that a degree of genetic relationship exists be- 
tween V. biflora and a Chamaemelanium violet. 

In Japan (table 1) the diploid V. dizflora remains below 1500 m alti- 
tude. Above 2100 m it is replaced by a closely related octoploid, V. 
crassa, n = 24 (Miyaji, 1929). 

Chromosomally and otherwise V. biflora appears to belong to a primi- 
tive group within the Chamaemelanium section, a group that has highly 
divergent pistil and stigma characters. Such a situation suggests that the 
Biflorae are central among the violets of the northern hemisphere and 
therefore are related to several sections of the genus Viola and probably 
to other genera as well. 

NATURAL INTER-SUBSECTIONAL HYBRIDS OF CHAMAEMELANIUM. 1. 
Purpureae — Nuttallianae, V. purpurea, n= 6, V. tomentosa, n= 6. 
Two hybrid localities were discovered by Baker within the territory of 
V. tomentosa. One was in Nevada Co. near Excelsior Point at 5000 ft in 
the yellow pine and incense cedar zone (Baker 8729) where the parental 
V. purpurea ssp. purpurea and V. tomentosa were found nearby. The 
other was at Little Grass Valley, Plumas Co. (Baker 9968), also there 
between parents, but in this case the purpurea parent belonged to ssp. 
integrifolia (Baker 9970) and the V. tomentosa parent was Baker 9969. 
In leaf form the hybrids resemble the V. purpurea parents but they pos- 
sess Shaggy, scattered pubescence reminding of the V. tomentosa parents. 
Both hybrids were fixed for chromosome investigations but were too old 
for meiosis. The pollen size varied greatly in both, suggesting unequal 
distribution of the parental chromosomes and accordingly general lack 
of homology. The seed capsules contained empty seeds. Obviously, V. 
purpurea and V. tomentosa are only remotely related, although the exist- 
ence of the normally developing hybrids indicates a measure of genetic 
relationship between the parental species. 

2. Chrysanthae « Purpureae; V. douglasu, n = 12, &* V. quercetor- 
um,n — 12. Throughout its tetraploid area V. douglasi regularly crosses 
with the equally tetraploid V. quercetorum, whenever they occur in the 
same habitat. Hybrids have been found in both Kern Co. and Santa 
Clara Co. and are completely sterile. In growth form the hybrids resem- 
ble the V. douglasii parent: they are geophytes, having their renewal 
buds buried below the ground surface. The hybrids are easy to recog- 
nize, however, as their leaves are pinnatifid instead of pinnatisect. Hy- 
brids of this kind were found in the foothills west of Glenville, Kern Co. 
(Keck & Clausen, 3188), and in the village of Glenville (Keck & Clau- 
sen, 3194). In Santa Clara Co.-on the east side of the Mount Hamilton 
Range at Arroyo Bayo, similarly growing with Quercus douglas and 
Pinus sabiniana, another hybrid population was discovered between the 


1964] CLAUSEN: VIOLA 185 


parental species (Keck & Clausen, 4546). In the Kern Co. hybrids the 
24 chromosomes were arranged in six pairs plus 12 single chromosomes, 
but the meiosis was more irregular in the Santa Clara Co. hybrid 
(table 1). It is not known whether the six pairs derive from conjugation 
between douglasti and quercetorum chromosomes or from autosyndesis 
among the chromosomes of one of the parent species. 

A hybrid V. douglasii «& V. purpurea was reported by Baird (1936). 
This was before V. quercetorum was published. The hybrid was collected 
with the parent species on April 18, 1935, in Walker Basin, between 
Caliente and Bodfish, Kern Co., on sunny slopes. The altitude of the 
Basin is approximately 3500 ft, and Mrs. Baird agreed that the altitude 
she quoted, 6000 ft, was too high. The Keck and Clausen hybrids were 
collected on April 13 the same year around Woody and Kernville, only 
about 30 mi farther north. Mrs. Baird’s careful description suggests that 
her V. purpurea was the tetraploid V. quercetorum, although her draw- 
ing is somewhat suggestive of the diploid V. purpurea ssp. mohavensis. 
Lacking the chromosome numbers of the Baird specimens the parentage 
of this hybrid cannot be decided with certainty. No seed developed on 
the garden transplant of this hybrid. 

Within the octoploid territory of V. douglasii a cytologically puzzling 
V. douglasi plant was found that could be a hybrid derivative of V. 
douglasu < V. quercetorum. This plant (Baker 8645) from Indian Val- 
ley, Plumas Co., has 29 pairs and one single chromosome, 2n = 59 (cited 
as V. douglasu in table 1). This plant occurred with V. quercetorum, 
n= 12 (Baker 8644, table 1). Baker suspected hybrids in this popula- 
tion and transplanted the V. douglasii plant to Kenwood, where the 
buds were fixed. It is possible that an unreduced V. douglasii ovule, 
2n — 48, pollinated by a reduced V. quercetorum pollen, n = 12, could 
result in a subdecaploid hybrid derivative, 2n = 59. The douglasii 
chromosomes would be expected to pair among themselves, providing 
about 24 pairs and possibly 6 pairs of qguercetorum chromosomes, equal- 

ling about 30 pairs. In such a hybrid the proportion of douglasii : quer- 
~ cetorum chromosomes would be 80 : 20 per cent, strongly in favor of 
douglasti. In comparison, the proportion is 50 : 50 per cent in the tetra- 
ploid hybrids (above), and in the octoploid hybrids discussed below. 
Morphologically, the 50 : 50 hybrids are already overwhelmingly to- 
wards V. douglasii. The strong hereditary influence of the species of the 
subsection Chrysanthae widens the possibilities concerning the ancestry 
of the species of that subsection. 

3. Chryvsanthae «K Nuttallianae; V. douglasti, n = 24, « V. praemorsa 
ssp. major, n — 24. Applegate discovered this hybrid (Applegate 8319, 
DS) in a narrow zone between the parental species on an exposed rocky 
slope among junipers and sagebrush in 1933 (table 1). During the first 
meiotic metaphase of the hybrid 48 unpaired chromosomes were ob- 
served, suggesting general absence of homology between the parental 
chromosomes. The hybrid was completely sterile. Morphologically, this 


186 MADRONO [Vol. 17 


octoploid Oregon hybrid is indistinguishable from the tetraploid Cali- 
fornia hybrids of V. douglasti *& V. quercetorum. Both kinds of hybrid 
have pinnatifid rather than pinnatisect leaves and would pass as belong- 
ing to the subsection Chrysanthae. In both kinds of hybrid the propor- 
tion of douglasii to other chromosomes is comparable. This fact, and the 
penetrating influence of V. douglasit makes the similarity understand- 
able. 

4. V. sheltonu, n = 6, * V. tomentosa, n = 6. The hybrid (near 
Weaver Lake, Sierra Co., Baker & Smith 8726) between these two di- 
ploid species grew with V. tomentosa in open coniferous forests at 5400 
ft altitude. It had somewhat tomentose pinnatifid leaves, suggesting its 
origin, but was not fixed for cytological investigation. 

PossIBLE NATURAL AMPHIPLOIDS. The Chamaemelanium section has 
evolved an unusual superstructure of polyploid species, ranging from 
diploid to 12-ploid. A survey of the arrangement of the species within 
the various polyploid levels was previously given (Clausen, 1951). Many 
amphiploids are expected in such a group, but the identification of the 
parents of natural amphiploids is generally hazardous. 

The safest principle in identifying amphiploids is to use the geographic- 
ecologic distribution of the amphiploid and its putative parents sup- 
ported by their gross morphological characters and chromosome num- 
bers. The morphology of the inter-subsectional hybrids listed above sug- 
gests that Vzola hybrids can be deceptive. In general, the parents of 
successful natural amphiploids are fairly remotely related and belong to 
distinct cenospecies, that is, in the Chamaemelanium section they may 
belong to distinct subsections. 

1. Viola quercetorum, n = 12: Suggested parent species, V. pedun- 
culata, n = 6, and V. purpurea, n = 6. Characters of V. quercetorum 
that remind of V. pedunculata are long peduncles, large petals, broadly 
subcordate lower leaves, branched rootstock, widely spreading habit, and 
especially its climatic region. Its morphological similarity to V. purpurea 
is obvious. Viola quercetorum occupies an ecological zone that became 
available after the rise of the Coast Ranges and after the development 
of the oak woodlands between the coast and the coniferous forests of the 
lower Sierra Nevada. Before that time V. pedunculata must have found 
congenial habitats over a considerably larger territory than now because 
the coastal influence reached much farther inland. Stebbins et al. (1963) 
claims that V. purpurea ssp. mohavensis and ssp. purpurea were the par- 
ents, and that V. pedunculata was not a parent. They base the conclu- 
sion upon chromatographic evidence but are aware that their two puta- 
tive parents are natives of cold winter climates, whereas V. quercetorum 
occupies a climatically milder zone. Since V. pedunculata probably was 
contiguous to V. purpurea ssp. purpurea in Pliocene central California 
and to relatives of ssp. mohavensis in Pleistocene southern California 
it is possible that V. quercetorum may have arisen more than once, from 
crossings between V. pedunculata and the two subspecies of V. purpurea. 


1964] CLAUSEN: VIOLA 187 


The high morphological variability within V. quercetorum and especial- 
ly the difference between southern and central California plants of the 
species lend credence to this possibility. 

I wholeheartedly agree with the plea by Stebbins et al. (1963) con- 
cerning the need for an extensive cytological investigation of the V. pur- 
purea relatives, especially in southern California. I disagree, however, 
with the interpretation of the distribution of subspecies presented in the 
map, especially the exclusion of ssp. purpurea from southern California 
and the extension of V. quercetorum to an altitude of 7000 ft. The last 
three V. quercetorum plants listed (Stebbins et al., 1963; table 1) belong 
to V. purpurea ssp. mohavensis or are introgressive hybrids between ssp. 
mohavensis and ssp. purpurea. 

2. Viola utahensis, n = 12: suggested parent species, V. vallicola, n = 
6, and V. purpurea ssp. venosa, n = 6. The narrowly endemic sagebrush 
species, V. utahensis (Baker, 1949b), is ecologically and morphologically 
intermediate between V. vallicola at 4000-6000 ft in open sagebrush 
flats, and V. purpurea ssp. venosa of moister, forested habitats, 5000- 
10000 ft, combining diploid species of subsect. Nwttallianae and Pur- 
pureae. Its herbage and seed capsules are minutely puberulent as in 
Pur pureae. The amphiploid can occasionally be found together with both 
putative parent species and is probably of recent origin. 

3. Viola linguifolia, n = 18: suggested parent species V. vallicola, 
n = 6, and V. nuttalli#i, n = 12. Viola linguifolia has broadly lanceolate 
leaves with ovate base, V. vallicola ovate leaves with truncate base, and 
V. nuttallu lanceolate leaves, tapering at both ends. V. nuttalli is dis- 
tinctly a prairie species, V. vallicola a species of sagebrush flats and open 
woods in the intermountain region, extending to the prairie region, where- 
as V. linguifolia is a species of montane regions, 5000-8000 ft. The two 
putative parents occasionally occur together, as for instance near Hulett, 
Crook Co., NE Wyoming at 4500 ft, where Ownbey 543-544 is V. nut- 
tallii from loose soil and steep hillsides, and Ownbey 553 is V. vallicola 
from grassy parks and divides. 


SECTION MELANIUM 


The homogeneity in the series of chromosome numbers of the Chamae- 
melanium section contrasts with the irregularity in the Melanium sec- 
tion, or the field pansies (Clausen, 1931). One group of its species ap- 
pears to follow a modified 6-series, n = 7, 8, 11, 12, 13, 17, 18, and 24, 
and a different group a 10-series, n = 10, 20, and 30. Both in chrom- 
osome numbers and in their pistil character the 6—series pansies resemble 
certain species of Chamaemelanium, such as V. pedunculata. 

The Melanium violets are almost exclusively European and west 
Asiatic, but one species, V. rafinesquiu, Greene, n = 17, is a native North 
American species. It is a weedy annual predominantly of the southeast- 
ern states but reaches western North America in Colorado. The history 


188 MADRONO [Vol. 17 


and relationships of this species are discussed in a separate paper 
(Clausen, Channell, and Nur, 1964). 


SECTION PLAGIOSTIGMA 


Most of the species of the Plagiostigma section have chromosome num- 
bers in multiples of 12 rather than in multiples of 6, but two of its groups 
deviate by having aneuploid numbers. Chromosomally the Plagiostigma 
violets could be considered to have been derived from an original 6-series. 
None have yellow petals. Examples of pistils and stigmas were shown in 
Clausen (1929). 

SUBSECTION STOLONOSAE (PALUSTRES). The relatives of V. palustris 
are a circumpolar group of species of moist meadow habitats or even shal- 
low bogs. The tops of their pistils are flattened or even expanded and 
provide a platform with a conical stigmatic papilla along the margin of 
the platform, a shape also found in some members of the Chamaemela- 
nium section. The flowers are white or lavender. 

The Stolonosae of North America comprise many diploids, n = 12 
table 1), and include V. mccabeiana, a local British Columbia species 
which Baker (1940) and Baird (1942) related to V. nephrophylla, 
n= 27, of the Boreali-Americanae. Its chromosome number, slender 
rhizomes, and ecology places it now with the Stolonosae. One species, 
V. epipsila, n = 6, is circumboreal, but the chromosome number of its 
western ssp. repens is unknown. A belt of delicate mountain bog species 
comprises V. macloskeyi (west coast and mountains) and V. pallens 
(eastern counterpart) and possibly even V. mccabeiana. Another belt 
of coarser plants at lower altitudes is V. occidentalis (western) V. lan- 
ceolata (eastern), V. vittata (southeastern), and V. primulifolia (east- 
ern half of continent). Gershoy (1932) recorded fertile F1 hybrids of 
V. pallens pollinated by V. lanceolata and V. primulifolia, suggesting 
close genetic relationships between species of both belts. 

Viola palustris, n = 24 (Gershoy, 1928) is a tetraploid, circumpolar 
species. In the wild it hybridizes with other members of the complex 
(Clausen, 1927). The western North American V. palustris ssp. brevipes, 
appears to be such a polyploid, chromosomally irregular hybrid deriva- 
tive (table 1). Since only a couple of samples of the V. palustris complex 
~ have been chromosomally investigated, it can be expected that a more 
extensive study will provide more surprises. 

SUBSECTION VAGINATAE (LANGSDORFFIANAE). This is a small essen- 
tially east Asiatic to western North American group of species. Three 
diploid species, n = 12, are known from Japan in addition to the octo- 
ploid V. langsdor fii (Regel) Fisch., n = 48 (Miyaji, 1929). Viola langs- 
dor fit also occurs in the Aleutian Islands and in Alaska, but its chromo- 
some number within that region is not known. Neither is the number 
known of the closely related V. s¢mulata Baker of moist woodsy areas 
of the Pacific coast from British Columbia to Oregon. 


1964] CLAUSEN: VIOLA 189 


SUBSECTION ADNATAE. The Adnatae, exemplified by V. patrini DC., 
n = 12, are distinguished by having the stipules attached to the pedicels. 
They constitute a huge east Asiatic polyploid complex of at least 24 
species, having chromosome numbers of n = 12, 24, 36 (listed as Plagio- 
stigma and Umbrosae by Miyaii, 1929). Scattered members of the Ad- 
natae occur beyond east Asia and include the western North American 
and circumpolar diploid V. selkirku Pursh, n = 12 (Gershoy, 1928, 
Miyaji, 1929) of the cool mountain forests and the European tetraploid 
V. pinnata L., n = 24, of the Swiss Alps (Clausen, 1927). 

SUBSECTION BLANDAE. The Blandae are chromosomally separated 
from other species of the Plagiostigma section in having a subtetraploid 
number of chromosomes, n = 22 (table 1; Gershoy, 1932). They consist 
of a species pair, V. blanda and V. incognita, eastern species that reach 
as far west as Minnesota and the Dakotas. 

SUBSECTION BOREALI-AMERICANAE. The North American stemless blue 
violets constitute another aneuploid group of the Plagiostigma section, 
consisting of approximately 30 species such as V. cucullata Ait. and V. 
palmata L. All are hypertetraploids, having n = 27 (Gershoy, 1928). 
They have pistils and stigmas similar to those of the Stolonosae but can 
be distinguished from them by their heavy, succulent rootstocks or rhi- 
zomes. Predominantly the Boreali-Americanae are an eastern North 
American group of taxonomically closely knit species that were made 
famous by Ezra Brainerd’s classical studies on Mendelism in wild violets 
during the early years of this century (Brainerd, 1906; 1921; 1924). 

Following Brainerd’s field and garden studies Gershoy (1928; 1932; 
1934) determined the chromosome numbers of the Boreali-Americanae 
and found them generally fairly interfertile. The aneuploid chromosome 
number of the group tends to separate them genetically from the other 
Plagiostigma violets. Before white man came to North America the taxa 
of the Boreali-Americanae occupied edaphically distinct niches within 
the huge forests of the continent. Their genetic barriers were only par- 
tial, however, and when new avenues for migration were opened through 
clearings in the forests they expanded their territories, hybridized, and 
entered an evolutionary cycle of adjustment to man-made environments. 

Viola nephrophylla is the only species of the Boreali-Americanae that 
extends all the way from the Atlantic coast across the continent to the 
Pacific states. It and two close relatives, V. cognata and V. arizonica 
Greene, are the only western North American representatives of the 
group, the latter two by some authors considered varieties of the former. 
Chromosomally, a western form of V. nephrophvlla and V. cognata are 
similar to the eastern taxa of the complex, having n = 27 chromosomes 
(table 1). 

Viola clauseniana conforms in gross morphology with the Boreali- 
Americanae, but it differs by having smooth, beardless petals and by a 
rare chromosome number n = 22 (table 1), one similar to that of the 
Blandae. According to Baker (1938) V. clauseniana occupies a precari- 


190 MADRONO [Vol. 17 


ously small, specialized habitat, not more than about 100 ft in diameter 
in Zion National Park, Utah, growing in a canyon at the base of Weep- 
ing Rock, a spot where sunlight seldom, if ever, comes. Viola nephro- 
phyla lives in the same general region, and V. arizonica was found at a 
higher altitude of 9000 ft. Viola clauseniana exemplifies how among the 
violets a species can become genetically isolated from most of its morpho- 
logically closest relatives. 

Another violet that in gross morphology is loosely reminiscent of the 
Boreali-Americanae is V. pedata L., n = 28 (Gershoy, 1932). It has, 
however, a very different pistil, and genetically it is a highly isolated 
species. In crossings with species of the Boreali-Americanae, Adnatae, 
Canadenses, Rostellatae, and others Gershoy (1932) obtained no seeds 
or only seeds unable to germinate. 


SECTION ROSTELLATAE 


The stemmed blue violets of North America belong to one of the groups 
of the section Rostellatae. These are circumboreal and have chromosome 
numbers in multiples of 10, rather than of 6 or 12 as in the previous sec- 
tions. In the Rostellatae the pistil ends in a straight or curved beak or 
rostellum, and the stigma is at the end of the beak (Clausen, 1929). 
Most of the species of the Rostellatae are Eurasiatic, but one of its sub- 
sections is circumboreal and has representatives in western North Amer- 
ica. The three subsections of the Rostellatae (Clausen, 1931b) are: 
1. Arosulatae Borb., Eurasiatic; simple biaxial growth habit as in the 
Nudicaules and in Canadenses of the section Chamaemelium, the primary 
stems elongate, and the flowers emerge from the leaf axils; n = 10 in 
V. stagnina Kit., n = 20, in V. elatior Fries and V. canina L. (Clausen, 
1926; 1927; 1929); some populations of the latter have accessory chrom- 
osomes (Clausen, 1931la); the Atlantic coastal V. /actea Sm. is subhexa- 
ploid and has n = 29 (Moore, 1959); 2. Scapigerae W. Beckr., Mediter- 
ranean-West Asiatic; biaxial, but nodes of primary stems do not elon- 
gate, and flowers develop from the axils of the rosette as in V. semper- 
virens and in Boreali-Americanae; n = 10 in V. odorata L. and V. hirta 
L.; n = 20 in V. sepincola Jord; 3. Rosulantes Borb., Eurasiatic and 
North American; triaxial, the primary stem develops a leaf rosette only 
and flowers appear from elongated secondary stems that come from the 
axils of the rosette as in the western North American V. adunca; n = 10 
in V. rupestris Schmidt and V. reichenbachiana Jord., n= 20 in V. rivi- 
niana Reich., some populations of the latter having accessory chromo- 
somes (Valentine, 1949). 

SUBSECTION ROSULANTES. Viola adunca, n = 10, occupies habitats 
from the Pacific coast to altitudes of 10000 to 12000 ft in the high Sierra 
Nevada and beyond to the intermountain region and the Rocky Moun- 
tains (Clausen, Keck, and Hiesey, 1940). No exception to n = 10 has 
been found (table 1). The inland and higher altitude forms differ in 


1964] CLAUSEN: VIOLA 191 


morphology from the type of the species which came from the Northwest 
coast. Many segregate names have been proposed for these taxa. Since 
the changes with altitude parallel those of the subspecies and ecotypes of 
Potentilla glandulosa from the same general region, and since the chrom- 
some numbers in about 55 taxa and natural hybrids of western North 
to be one huge species with many subspecies (table 1). 

Viola howelli is a tetraploid and octoploid counterpart of V. adunca 
(table 1). It occurs in the lower Coast Ranges, valleys, and lower Cas- 
cade Mountains of Oregon and Washington. Gershoy (1928) found that 
plants sent by Peck of Willamette University were tetraploid (probably 
n = 20). In contrast, plants from near the type locality of the species, 
at Oregon City near Portland, Oregon, were octoploid, n = ca. 40 (Ger- 
shoy, i932). From this preliminary sampling it is obvious that V. how- 
elliti merits a more thorough chromosomal study throughout its territory. 

The stemmed blue violets of eastern North America are chromosom- 
ally uniform: Gershoy (1928; 1932) reported n = 10 in V. conspersa 
Reichb., V. /abradorica Schwein., V. rostrata Pursh. and V. striata Ait. 
Miyaji (1913; 1929) likewise found n = 10 in 10 Japanese species of 
Rosulantes. The existence of 10 as a basic number in the Rostellatae 
from three continents is therefore well established. 

Attention is called to the fact that the pistils of the two western mem- 
bers of the Rostellatae tend to expand into a head (Clausen, 1929), ap- 
proaching in this character some of the western Chamaemelanium species 
and deviating from Rostellatae in other parts of the world. 


CONCLUSIONS 


This paper documents approximately 115 determinations of chromo- 
some numbers in about 55 taxa and natural hybrids of western North 
American violets (table 1). The text discusses the taxonomic, ecologic, 
geographic, and evolutionary aspects of the chromosome situation of 
these taxa and of their relatives in other parts of the world. 

The Viola genus probably has some 600 species distributed among 
about 14 sections in northern and southern hemispheres. The sections 
of Viola are more like genera among other plants, but it would be taxo- 
nomically unwise to partition the genus Viola. It has evolved a unique 
and distinctive floral apparatus that makes it easy to recognize a violet 
in any part of the world. Four sections are native to the temperate zones 
of the northern half of our earth, and all four are represented within 
our area. 

The Chamaemelanium section is evolutionally distinct from the other 
sections of the genus by strictly following a polyploid series of chrom- 
osome numbers with x = 6 as the basic number. All levels of polyploidy 
from the diploid n = 6 to the duodecaploid, n = 36, are represented 
within the western Chamaemelanium violets. Uniformity in regard to 
basal chromosome number is contrasted with extreme variability in re- 


192 MADRONO [Volei7 


gard to characters of the pistil and stigma, which within other northern 
hemisphere sections are relatively constant. The strict 6—series of chrom- 
osome numbers is mainly limited to the violets of the Chamaemelanium 
section and to a few Melanium species. This particular base number, 
found also in the genus Hybanthus, and the great variability in pistils 
and stigmas are indications that the section is a central and primitive one 
within the violets. 

The Chamaemelanium section has its richest development in the topo- 
graphically and ecologically diversified western North America. In this 
region it has evolved about 33 taxa, belonging to at least seven polyploid 
lines of distinct evolutionary sequences. Some of these lines have repre- 
sentatives in the eastern part of North America and in east Asia as indi- 
cated in table 1, and one species, the diploid V. diflora, n = 6, is cir- 
cumpolar. 

The other 17 taxa from western North America belong to seven sub- 
sections of three other sections. These sections are: 1. Melanium, or the 
field pansies, with one 17-chromosome southeastern species that has 
reached Colorado; 2. Plagiostigma, which follows a 12-series, n = 12, 
24, 36, 48; several strictly polyploid groups are augmented by adjunct 
aneuploid ones, n = 22, 27, 28; and 3. Rostellatae, characterized by a 
10-series, having n = 10, 20, 40 chromosomes. The latter three sections 
are more richly represented in other parts of the world, but the 17 west- 
ern North American taxa provide a fairly representative sampling of 
even these sections. 

One diploid species of Chamaemelanium, V. purpurea, n = 6, anda 
diploid species of the Rostellatae, V. adunca, n = 10, have evolved series 
of ecotypic subspecies that enable them to occupy a nearly complete 
range of habitats within western North America without changing their 
chromosome numbers. 

Spontaneous interspecific hybrids of V. purpurea * V. tomentosa, 
V. douglasit K V. quercetorum, and V. douglasit * V. praemorsa ssp. 
major were studied cytologically. 

The chromosome numbers of the following species have not previously 
been counted: V. scopulorum, n = 12; V. macloskeyi, n = 12; V. mc- 
cabetana, n = 12; V. occidentalis, n = 12; V. palusinis ssp.vOorevepes, 
n = ca. 36-48, irregular, apparently a hybrid; V. cognata, n = ca. 27; 
and V. clausentana, n = ca. 22. 


TABLE 1. SOURCES OF CHROMOSOME NUMBERS IN VIOLAL 


SECTION CHAMAEMELANIUM Ging. 


1 Abbreviations: MSB = M. S. Baker collection numbers, primarily at UC; MSB 
transplant = Baker’s garden at Kenwood, Sonoma Co.; JC = Jens Clausen collec- 
tion numbers; CIW = Carnegie Institution of Washington garden at Stanford. The 
state is not indicated for Califcrnia localities. 


1964] CLAUSEN: VIOLA 193 


SUBSECTION BIFLORAE Clausen 


V. biflora L. n = 6: Botanical garden material (Clausen, 1926; 1927); unknown 
(Gershoy, 1928) ; Japan, below 1500 m (Miyaji, 1929). 

V. crassa Makino. n = 24: Japan, alpine, above 2700 m (Miyaji, 1929). 

V. orbiculata Geyer. n = 24: North America (Gershoy, 1932). 

V. sempervirens Greene. Tetraploid form, southern. n= 12: Santa Cruz Co., 
S of Boulder Creek; garden transplant from central California (Clausen, 1929). 
Octoploid form, northern, n = 24: Oregon, dry woods, collected by Peck (Gershoy, 
1934) ; southern Oregon, collected by Purdy, 1926 (Gershoy, 1934). 

V. rotundifolia Michx. n = 6: Unknown, E North America (Gershoy, 1934). 


SUBSECTION CANADENSES W. Beckr. 
V. canadensis L. n = 12: Unknown (Gershoy, 1928; Bold and Gershoy, 1934). 
V. ocellata T. & G. n= 6: Unknown (Gershoy, 1928); garden transplant from 
central California (Clausen, 1929). 
V.rugulosa Greene. n = 12: Botanical garden source as “V. rydbergit Greene” 
(Clausen, 1926; 1927). 
V. scopulorum Greene. n = 6: MSB V-254 transplant, source unknown. 


SUBSECTION CHRYSANTHAE W. Beckr. 

V. douglasii Steud. Tetraploid form, southern, n= 12: Kern Co., summit of 
Tehachapi Pass (Clausen, 1929); % miS of Tehachapi Pass, MSB, CIW 5576 fixa- 
tion; W of Glenville, Keck & Clausen 3187; Santa Clara Co., Arroyo Bayo, Keck & 
Clausen 4544. Octoploid form, northern, n = 24: Sonoma Co., Melita, JC 732; 
Plumas Co., Indian Valley, MSB 8645 transplant, 2n = 59, possibly derivative of 
V. quercetorum (MSB 8644, n = 12) in same locality; Oregon, Klamath Co., Swan 
Lake Valley, Applegate 9318, 2n = 48. 

V. douglasiz, n= 12, & V. quercetorum, n = 12: Kern Co., Glenville, Keck & 
Clausen 3194-2, 6;; + 12,, 2n = 24; Santa Clara Co., Arroyo Bayo, Keck & Clau- 
sen 4546-1, 2n = 24, metaphase I highly irregular. 

V. douglasii?, n = 24, & V. praemorsa ssp. major, n = 24 (“V. praemorsa ssp. 
arida” according to Baker): Oregon, Klamath Co., Swan Lake Valley, Applegate 
8319 (DS), 2n = 48, no pairing in metaphase I. 

V. hallit Gray. n = 36: Collected by Purdy, April, 1928 (Gershoy, 1932), 2n = 
Cae i 2. 

V. sheltoni Torr. n = 6: Nevada Co., Big Ben Ranger Station, MSB 5310 trans- 
plant. 

SUBSECTION NupIcAuLEs W. Beckr. 

V. alliariifolia Nakai. n = 6: Japan (Miyaji, 1929). 

V. brevistipulata W. Beckr. n= 6: Japan, as V. glabella Nutt. (Miyaji, 1913; 
1927). 

V. eriocarpa Schwein. n = 6: E North America (Gershoy, 1928). 

V. glabella Nutt. n= 12: Garden transplant from central California (Clausen, 
1929). 

V. kiskidai Nakai. n = 6: Japan (Miyaji, 1929). 

V.lobata Benth. n = 6: Unknown source (Gershoy, 1928). 

V. pubescens Ait. n= 6: E North America (Gershoy, 1928); Wisconsin, near 
Madison, in woods (Clausen, 1929). 


SUBSECTION NUTTALLIANAE Baker (W. Beckr. pp.) 

V. bakeri Greene. n = 24 (Baker, 1949c) ssp. bakeri: Tehama Co., Mt. Lassen 
National Park, 6000 ft, MSB 8377; Sierra Co., Webber Lake, open forest, 6770 ft, 
MSB 8388; Nevada Co., W of Big Bend Ranger Station, MSB 5215 transplant; 
Bear Valley—Bowman Lake Rd., MSB 8406; Placer Co., Road Truckee to Tahoe 
Tavern, 6100 ft, MSB 8403. 

V. linguifolia Nutt. n= 18: Idaho, near Moscow, collected by Fisher as “V. 
praemorsa” (Gershoy, 1934); Utah, Cache Co., Brush Canyon, MSB 4465 trans- 
plant. 


194 MADRONO [Vol. 17 


V. nuttalli Pursh. n = 12 (Baker, 1949d): Canada, Alberta, Calgary, McCalla 
7335; Colorado, nursery plant from Andrews Nursery, MSB V-279 transplant. 

V. praemorsa Dougl. ssp. praemorsa. n = 18 (Gershoy, 1934): Washington, S of 
Tacoma, coastal prairie, MSB 9634 transplant; Oregon, Josephine Co., Takilma, 
Purdy nursery stock (Gershoy, 1934) ; Humboldt Co., Yager, MSB 4059 transplant. 

V. praemorsa ssp. major (Hook.) Baker & Clausen. n = 24 (Baker, 1949d): 
Washington, Kittitas Co., E of Goldendale, MSB 7386, region of the type; Idaho, 
Nez Perce Co., near Cul-de-Sac, MSB 7399, type locality of “V. flavovirens Poll.”; 
Idaho Co., Kamiah, N bank of Clearwater River, MSB 7408; Oregon, Klamath Co., 
Swan Lake Valley, with hybrid, Applegate 8317 (DS); Sierra Co., near Webber 
Lake, 6700 ft, MSB 8385, type of “ssp. arida Baker.” 

V. praemorsa ssp. oregona Baker & Clausen. n = 24 (Baker, 1949d): Oregon, 
Klamath Co., McCullom’s Mill on road to Medford, MSB 8292; E side of Klamath 
River, on road to McCullom’s Mill towards California, MSB 8288; Siskiyou Co., 
Yreka, MSB 7354; Trinity Co., Scott Mt., 5800 ft, MSB 13139, mistakenly listed as 
V. bakeri ssp. shastensis, n = ca. 24 (M.A. Nobs). 

V. tomentosa Baker & Clausen. n = 6 (Baker, 1949b): Nevada Co., Cisco-Grass 
Valley Road, Pinus ponderosa forest, 5000 ft, MSB 8699. 

V. vallicola Nels. n = 6 (Baker 1949d): Utah, Cache Co., N outskirts of Provi- 
dence, about 5000 ft, Maguire 16928-8, as “V. praemorsa.” 


SUBSECTION PEDUNCULATAE Clausen 


V. charlestonensis Baker & Clausen. n = 6 (Baker, 1949c): Nevada Charleston 
Mts., MSB transplant; Charelston Mts., Lee’s Canyon, MSB 8673. 

V. pedunculata T. & G. ssp. pedunculata. n = 6 (Clausen, 1929): Solano Co., 
Rockville, 500 ft, Stebbins 6003 (Stebbins et al., 1963) ; San Mateo Co., San Bruno 
Mt., J. & J. M. Webber (Clausen, 1929); Monterey Co., Jolon, 1200 ft, Stebbins 
6019 (Stebbins et al., 1963). 

V. pedunculata ssp. tenuifolia Baker. n= 6 (Baker, 1949d): San Benito Co., 
Pinnacles, M7SB 9267 transplant; San Luis Obispo Co., S of Paso Robles, 500 it, 
Stebbins 6018 (Stebbins et al., 1963). 


SUBSECTICN PURPUREAE Baker 


V. purpurea ssp. atriplicifolia (Greene) Baker & Clausen. n = 6 (Baker, 1949d): 
Oregon, Deschutes Co., Bend, MSB 7414. 

V. purpurea ssp. aurea (Kell.) Baker & Clausen. n = 6 (Baker, 1949d): Nevada, 
near Reno, Hunter’s Creek, among rocks and sagebrush, MSB 8634. 

V. purpurea ssp. dimorpha Baker & Clausen. n = 6 (Baker, 1949d): Tehama Co., 
at entrance to Mt. Lassen National Park, 6000 ft, MSB 8378; Child’s Meadow, E of 
Mineral, 5000 ft, MSB 8379; Plumas Co., S of Humbug Summit, 6000 ft, Keck & 
Clausen 3770; S of Chaparral, 5000 ft, Keck & Clausen 3775. 

V. purpurea ssp. geophyta Baker & Clausen. n-== 6 (Baker, 1949d): Oregon, 
Klamath Co., 30 mi SE of Lapine, volcanic ash in yellow-lodgepole pine forest, 
Keck & Clausen 3707, type; Lassen Co., near Westwood, among Pinus jeffreyi and 
Abies concolor, Keck & Clausen 3766. 

V. purpurea ssp. integrifolia Baker & Clausen. n = 6 (Baker, 1949d): Plumas- 
Butte Co. line, Humbug Summit, 6500 ft, in Abies magnifica forest, Keck & Clausen 
3769, type. 

V. purpurea ssp. mesophyta Baker & Clausen. n = 6 (Baker, 1949d): Mariposa 
Co., Yosemite National Park, Porcupine Flat, 8100 ft, JC 1098; Tuolumne Co., 
Tenaya Lake, 8200 ft, JC 1099; above Tuolumne Meadows, 9000 ft, transition to 
ssp. xerophyta, JC 1100. 

V. purpurea ssp. mohavensis Baker & Clausen. n = 6 (Baker, 1949d): San Benito 
Co., Lockwood Valley, CIW 1815 transplant; San Bernardino Co., Horse Thief Can- 
yon, 328 ft, Clokey 5833, type; 7 mi SE of Hesperia, E side of Mohave River, 3100 
ft, Stebbins 6017 (Stebbins et al., 1963). 


1964] CLAUSEN: VIOLA 195 


V. purpurea ssp. purpurea. n = 6 (Baker 1949d): Trinity Co., 35 mi NW of 
Redding, MSB 7427; Tehama Co., near Payne’s Creek, 1600 ft, MSB 8655; Lake 
Co., 2.5 mi N of Salmina’s Resort, in Pinus ponderosa and Pseudotsuga forest, 2500 
ft, Stebbins 6011; Bartlett Mt., 4100 ft, Stebbins 6008; Loch Lomond Resort, in 
Pinus ponderosa forest, 2800 ft, Stebbins 6022 (Stebbins et al., 1963) ; Tuolumne Co., 
Mather, Hog Ranch, 4600 ft, with Pinus ponderosa, CIW 1015-1 transplant. 

V. purpurea ssp. venosa (Wats.) Baker & Clausen. n= 6 (Gershoy, 1934): 
Washington, Kittitas Co., Blewett Pass, 4100 ft, Keck & Clausen 3550; near Bingen, 
collected by Suksforf (Gershoy, 1934); Utah, Wasatch Co., Timpanogas Highway, 
1 mi E of summit, in aspen grove, MSB 8529. 

V. purpurea ssp. xerophyta Baker & Clausen. n = 6 (Baker, 1949d): Mono Co., 
Mt. Conness, terraces on E slope, 10400 ft, JC 524. 

V. quercetorum Baker & Clausen. n = 12 (Baker, 1949): Oregon, Josephine Co., 
Wimer, MSB 9049 transplant; as “V. purpurea,’ collected by Purdy (Gershoy, 
1934) ; Plumas Co., Indian Valley, MSB 8644; Lake Co., Loch Lomond Resort, 2800 
ft, Stebbins 6021 (Stebbins et al., 1963): S of Kelseyville, MSB 8208; at Lakeview- 
Knoxville Road, MSB 8169, n = 12 + fragment; Napa Co., Howell Mt., Pine Flat, 
MSB 7666 transplant; Howell Mt., near Pacific Union College, 1870 ft, Stebbins 
5986, 6007 (Stebbins et al., 1963) ; Marin Co., Mt. Tamalpais, upper limit of forest, 
JC 1178; Contra Costa Co., Mt. Diablo, at summit, JC 1172, 1173; Santa Clara Co., 
Mt. Hamilton, brick kiln on W side, Keck & Clausen 4516; Mt. Hamilton, east side, 
Arroyo Bayo, Keck & Clausen 4545, hybridized with V. douglasi; Monterey Co., 
9 mi NW of Jolon, 1200 ft, Stebbins 6020 (Stebbins et al., 1963), near V. peduncu- 
lata; San Benito Co., 5.4 mi S of Willow Creek School, CIW 1816 transplant; Kern 
Co., at road from Woody to Kernville, to summit, Keck & Clausen 3185; W of 
Glenville, Keck & Clausen 3186-1, type locality, hybridizing with V. douglasii; 
Tehachapi Pass, MSB 8806. 

V. utahensis Baker & Clausen. n = 12 (Baker, 1949b): Utah, Cache Co., Logan 
Campus, under Artemisia tridentata, Maguire 16022; mouth of Providence Canyon, 
under Artemisia, Maguire 16027-6, type locality; near Providence, under Prunus, 
Maguire 16027-7. 


SECTION PLaGiosTIGMA Godr. (NOMINIUM pp.) 
SUBSECTION Blandae Gershoy 


V. blanda Willd. n = 22 (Gershoy, 1932). 
V. incognita Brain. n = 22 (Gershoy, 1932). 


SUBSECTION BOREALI-AMERICANAE W. Beckr. 


V. clauseniana Baker. n = ca. 22: Utah, Zion National Park, base of Weeping 
Rock area, MSB 8438 transplant. 

V. cognata Greene. n = ca. 27: Colorado, Estes National Park, MSB transplant. 

V. nephrophylla Greene. n = 27 (Gershoy, 1928): Plumas Co., Drakesbad, MSB 
transplant. 

16 related species. n = 27: E and central United States (Gershoy, 1929; 1932). 


SUBSECTION STOLONOSAE Kupffer 


. epipsila Ledeb. n = 12 (Clausen, 1926; 1927): Denmark, Lyngby. 
. lanceolata L. n = 12 (Gershoy, 1928): Unknown locality. 
. mccloskeyi Lloyd. n = 12: Tulare Co., Mineral King, MSB transplant. 
. mocabeiana Baker. n = 12: Canada, British Columbia, Columbia Lake, MSB 
9444 transplant, from type locality. 

V. occidentalis (Gray) Howell. n= 12: Oregon, Josephine Co., Kerby, MSB 
transplant. 

V. pallens (Ging.) Brain. n = 12: (Gershoy, 1928) ; Unknown locality. 


Sh SS 


196 MADRONO [Vol. 17 


V. palustris X V. epipsila. 2n = 36, 12,, + 12,: Denmark, forest meadow (Clau- 
sen, 1927). 

V. palustris ssp. brevipes Baker. Irregular meicsis, 36-38 pairs or singles: Colo- 
rado, Estes Park, MSB 7629 transplant. 

V. primulifolia L.n = 12 (Gershoy, 1928; 1932): E North America. 

V. shihokiana Makino. n = 12 (Miyaji, 1929): Japan. 


SECTION RGSTELLATAE Boiss. (NOMINIUM pp.) 
SUBSECTION ROSULANTES Borb. 


V. adunca Sm. ssp. adunca. n= 10 (Clausen, 1929): British Columbia, Van- 
couver, MSB transplant; Mendocino Co., Albion, bluffs, CIW 1828-1 transplant; 
Marin Co., Mt. Vision (Clausen, 1929); San Mateo Co., San Bruno Mt., collected 
by I. & J. M. Webber (Clausen, 1929) ; Montara, JC 630, CIW 1817 transplant. 

V. adunca ssp. ashtona Baker. n = 10: Colorado, Estes Park, Cub Lake Trail, 
8500 ft, MSB 5430 transplant, type localtiy. 

V. adunca ssp. oxyceras (Wats.) Jeps. n= 10: Oregon, Grant Co., 3 mi W of 
Dixie Pass, Keck & Clausen 3659; Nevada Co., Big Bend Ranger Station, MSB 
transplant; Donner Lake, 7000 ft, MSB transplant; Tuolumne Co., Mather, 4600 
ft, CIW 1829-1 transplant. 

V. adunca ssp. radicosa Baker, (cf. V. bellidifolia Greene). n = 10: Colorado, 
Rocky Mountains National Park, Kewuneeche Valley, 12 mi N of Grant Lake, WSB 
Al 72s LY Pe: 

V. howellii Gray. n = 20, n = 40 (Gershoy, 1928; 1932): tetraploid form, cen- 
tral Oregon, collected by Peck, May 26, 1925 (Gershoy, 1928, ““2n = 42”). Octoploid 
form, Oregon, near Oregon City, collected by Gale, Wildwood Nurseries, Portland, 
type locality (Gershoy, 1932). 


Department of Plant Biology, Carnegie Institution of Washington, 
Stanford, California 


LITERATURE CITED 


Barr, V. B. 1936. A natural violet hybrid. Madrono 3:325-327. 

. 1942. Wild violets of North America. Univ. Calif. Press, Berkeley. 

Baker, M.S. 1935. Studies in western violets. I. Madrofio 3:51-57. 

—. 1936. Studies in western violets. II. New Species and varieties. Madrono 

3:232-239. 

—, 1938. An undescribed species of Viola from Utah. Madrono 4:194-196. 

——-—. 1940. Studies in western violets. III. Madrono 5:218-231. 

. 1947. A new violet from Mexico. Madrono 9:131-137. 

————. 1949a. A new western violet. Leafl. West Bot. 6:101-102. 

————., 1949b. Studies in western violets. IV. Leafl. West. Bot. 5:141-147. 

—_—. 1949c. Studies in western violets. V. Leafl. West. Bot. 5:173-177. 

——-—. 1940d. Studies in western violets. VI. Madrono 10:110-128. 

. 1953. Studies in western violets. VII. Madrono 12:8-106. 

. 1957. Studies in western violets. VIII. The Nuttallianae continued. Brit- 

tonia 9:217-230. 

. 1960. Studies in western violets. IX. Miscellaneous species in the sections 
Nomimium and Chamaemelanium. Madrono 15:199-204. 

Bop, H. C. and A. GersHoy. 1934. Studies in North American violets. IV. Chromo- 
some relations and fertility in diploid and tetraploid species hybrids. Vt. Agr. 
Exp. Sta. Bull. 378. 

BRAINERD, E. 1906. Hybridism in the genus Viola. III. Rhodora 8:49-61. 

. 1921. Violets of North America. Vt. Agr. Exp. Sta. Bull. 224. 

. 1924. Some natural violet hybrids of North America. Vt. Agr. Exp. Sta. 

Bull. 239. 


1964] CLAUSEN: VIOLA 197 


CLAUSEN, J. 1926. Genetical and cytological investigations on Viola tricolor L. and 
V. arvensis Murr. Hereditas 8:1-156. 

. 1927. Chromosome number and the relationship of species in the genus 
Viola. Ann. Bot. 41:677-714. 

. 1929. Chromosome number and relationship of some North American 
species of Viola. Ann Bot. 43:741-764. 

— 193la. Viola canina L., a cytologically irregular species. Hereditas 15:67- 

88. 

. 1931b. The Viola species of Denmark. Bot. Tidsskr. 41:317-335. 

. 1931c. Cyto-genetic and taxonomic investigations in Melanium violets. 
Hereditas 15:219-304. 

. 1951. Stages in the evolution of plant species. Cornell Univ. Press, Ithaca. 

CLAUSEN, J., R. B. CHANNELL, and U. Nur. 1964. Viola rafinesquii, the only Melan- 
ium violet native to North America. Rhodora 66:32-46. 

CLAUSsEN, J., D. D. Keck, and W. M. Hiesey. 1940. Experimental studies on the 
nature of species. I. Effect of varied environments on western North American 
plants. Carnegie Inst. Publ. 520. 

GersHoy, A. 1928. Studies in North American violets. Vt. Agr. Exp. Sta. Bull. 279. 

. 1932. Descriptive notes for Viola exhibit. The Nominium and Chamaeme- 
lanium section. Sixth Intl. Cong. Genetics, Ithaca, N.Y. (A. Gershoy, Vt. Agr. 
Exp. Sta., Burlington, 27 pp.) 

1934. Studies in North American violets. III. Chromosome numbers and 
species characters. Vt. Agr. Exp. Sta. Bull. 367. 

Mriyajl, Y. 1913. Untersuchungen uber die Chromosomenzahlen bie einigen Viola- 
Arten. [In Japanese.] Bot. Mag. Tokyo. 27:443-460. (German abstract, Bot. 
Mag. Tokyo 41:262-268. 1927). 

. 1929. Studien uber die Zahlenverhaltnisse der Chromosomen bei der 
Gattung Viola. Cytologia 1:28-58. 

Moore, D. M. 1959. Population studies on Viola lactea Sm. and its wild hybrids. 
Evolution 13:318-332. 

STEBBINS, G. L., B. L. Harvey, E. L. Cox, J. N. RutcEer, G. JELENcovic, and E. 
YaciL. 1963. Identification of the ancestry of an amphiploid Viola with the aid 
of paper chromatography. Am. Jour. Bot. 50:830-839. 

VALENTINE, D. H. 1949. Vegetative and cytological variation in Viola riviniana. Bot. 
Soc. British Isles, Rep. Conf. Stud. Critical British Groups, pp. 48-53. 


NOTES AND NEWS 


NEw DIstTRIBUTION RECORD FOR HELEOCHLOA ALOPECUROIDES IN OREGON.—Re- 
cently Chambers and Dennis (Madronfio 17:92. 1963) reported on the distribution of 
H. alopecuroides (Pill. & Mitterp.) Host in Oregon. The following record should be 
included as a range extension of 150 air mi across several mountain ranges and 
divides to the Rogue River Valley in western Curry Co. about 35 mi from the Pacific 
Ocean. This is an area which has long been noted for its bizarre and unusual plant 
records (Baker, Leafl. West. Bot. 6:82-84. 1950). The grass was collected on sandy 
shores along the Rogue River, 3 min of Agness, Curry Co. (Baker 16559, Aug. 22, 
1963, ID, OSC). —Wi111AmM H. Baker, Department of Biological Sciences, Univer- 
sity of Idaho, Moscow. 


198 MADRONO [Vol. 17 


NOMENCLATURAL PROBLEMS IN THE ACACIA CORNIGERA 
COMPLEX 


VELVA E. Rupp 


The name Acacia cornigera (L.) Willd., based on Mimosa cornigera L., 
has been applied to two different species by modern authors. The ques- 
tion of the correct usage of the name was raised by Daniel H. Janzen, a 
student of insect ecology at the University of California, Berkeley, who 
is interested in the ants of the genus Pseudomyrmex that inhabit the 
thorns of the Mexican “bull horn” acacias. 

Janzen noted that Standley (1922) placed A. cornigera in synonymy 
with A. spadicigera Schl. & Cham. and recognized A. sphaerocephala 
Schl. & Cham. as a separate species. Britton and Rose (1928) made the 
opposite determination, placing 4. sphaerocephala in synonymy with A. 
cornigera and separating A. spadicigera, the two taxa being designated as 
the segregate genus Tauroceras Britt. & Rose. In at least two recent floras 
(Standley and Steyermark, 1946; Leon and Alain, 1951) the nomencla- 
ture of Britton and Rose was followed as to species, although the generic 
status of Tauroceras was not recognized. 

The identity of A. cornigera and its relationship to A. spadicigera and 
A. sphaeroce phala, as well as the putative synonymy, obviously is in need 
of clarification. 

A part of the confusion is traceable to Linnaeus’ original descriptions 
of Mimosa cornigera (1737, 1753) in which he included in his literature 
citations, references to material from the East Indies, although the type, 
from the garden of George Clifford, was presumably of Mexican origin. 
De Candolle (1825), on the basis of the literature, separated A. cornigera 
into two varieties, americana and indica. 

Schlechtendal and Chamisso (1830), considering A. cornigera to rep- 
resent a mixture of species, rejected the name and published two new 
ones to identify Mexican collections made by Schiede and Deppe, viz. 
A. spadicigera Schl. & Cham. and A. sphaerocephala Schl. & Cham. 

Schenck (1913) followed Schlechtendal and Chamisso in disregarding 
A. cornigera and described three additional species, A. cubensis, based on 
Mexican material introduced into Cuba, A. nicovensis, from Costa Rica, 
and 4. veracruzensis, from Mexico. Safford (1914, 1915) considered A. 
cornigera to be distinct from both A. spadicigera and A. sphaerocephala, 
accepted Schenck’s three species, and added another three names to the 
complex, A. hernandezti, A. furcella, and A. dolichocephala, all based on 
Mexican collections. 

This group of some nine published species is characterized by inflated, 
indehiscent fruits and paired, stipular spines that, in symbiosis with ants, 
may develop into thorns as much as 11 cm long, suggesting miniature 
replicas of the horns of Longhorn cattle. One or more ‘‘boat-shaped”’ 
glands may occur on the axis of the leaf, at least one usually on the 


1964] RUDD: ACACIA 199 


petiole just below the first pair of pinnae. Nectar glands commonly are 
present at the tips of the young leaflets. 


Examination of pertinent herbarium material confirms that two species, 
or groups of species, can be recognized. The members of one group, in- 
cluding A. sphaerocephala, A. veracruzensis, and A. dolichocephala, have 
similar, globose inflorescenses and leaflets with only the midvein, or costa, 
evident. The species of the other group, A. spadicigera, A. cubensis, A. 
nicoyvensis, A. hernandezii, and A. furcella, all exhibit oblong, spicate 
inflorescenses and leaflets with secondary veins clearly visible. 


Standley (1922) and succeeding authors have agreed that the two 
groups of species are reduceable to two species, one of which is referable 
to Acacia cornigera. A third related species, Acacia mavyana Lundell 
(1937), is apparently distinct and is excluded from further discussion in 
this paper. 

The type of Mimosa cornigera L., the basionym of Acacia cornigera, 
is in the Clifford Herbarium (BM). It is a sterile specimen, and the only 
useable comparative character is the venation of the leaflets. Fortunately, 
that is sufficiently distinctive in the two species to permit recognition. 

Through the kindness of W. T. Stearn of the British Museum, I have 
been permitted to examine a few leaflets from that type specimen and 
have found that numerous secondary veins are clearly visible, as in 4. 
spadicigera, et al. Therefore, I believe that Standley’s original interpre- 
tation, placing A. spadicigera in synonymy under A. cornigera, was cor- 
rect, and that A. sphaerocephala is a distinct species. 

Another question raised by Janzen concerns the taxonomic rank of 
Tauroceras. The species discussed above constitute the genus Tauroceras 
Britt. & Rose and also Safford’s “group” Ceratophysae of the genus 
Acacia. It is difficult to categorize this assemblage of species whose chief 
common character is the indehiscence of the pods due to lack of sutures. 
There are other species of Acacia such as those in the segregate genera 
Vacuhellia and Poponax with similar appearing terete or subterete fruits 
in which the sutures are developed and dehiscence may occur. Anatomical 
and biosystematic studies are needed to elucidate the relationships. I do 
not believe that generic, or even subgeneric, distinction is warranted, but 
that the term “Acacia cornigera complex” is preferable at this premature 
stage of our knowledge. 


An exhaustive treatment of the complex is beyond the scope of this 
paper, but the following brief resumé, including citations of specimens 
examined, may be helpful. 


Inflorescences oblong, spicate, the interfloral bracteoles with acuminate, sometimes 
sagittate laminae, the tips often recurved; leaflets with costa and secondary 
Ati nVelt 9 (0) 0 Ware (220 eliaslen 0 Ke (3) 0 Genaeere ata Mann Rey em OR RY: Seer ei: rene eee ars 1. A. cornigera 
Inflorescences globose, capitate, the interfloral bracteoles obtuse; leaflets with costa 
present but secondary venation not evident. 2. A. sphaerocephala 


200 MADRONO [Vol. 17 


1. ACACIA CORNIGERA (L.) Willd. Sp. Pl. 4:1080. 1806. Mimosa cor- 
nigera L. Sp. Pl. 520. 1753. A. cornigera var americana DC. Prodr. 2:460. 
1825. A. spadicigera Schl. & Cham. Linnaea 5:594. 1830. A. cubensis 
Schenck, Repert. Sp. Nov. 12:360. 1913. A. nicoyensis Schenck, Repert. 
Sp. Nov. 12:360. 1913. A. kernandezti Safford, Jour. Wash. Acad. 4:358. 
1914. A. furcella Safford, Jour. Wash. Acad. 4:359. 1914. Tauroceras 
spadicigerum (Schl. & Cham.) Britt. & Rose, N. Am. Fl. 23:85. 1928. 
T. cornigerum (L.) Britt. & Rose, N. Am. Fl. 23:86. 1928, excl. synon. 


Type: Cultivated, presumably from material introduced from Mexico (BM, 
photograph and fragment of type seen). 


Representative specimens seen. MEXICO. Veracruz: near Laguna Verde, Schiede 
& Deppe 685 (US, fragment of type of A. spadicigera ex HAL, photograph of isotype 
ex B); Lake Catemaco, Nelson 427 (US, type of A. furcella); San Francisco, 
Smith 1509 (NY); Alvarado, King 2431 (US); Cuitlahuac, King 2671 (NY, US); 
Veracruz, Miller 89 (NY); Zacuapan, Schenck 836 (US); Purpus 7748 (NY, US); 
Pueblo Viejo, Palmer 448b (NY, US); Cordoba, Fisher 93 (US). San Luis Potosi: 
Rascon, Palmer 669 (NY, US, type of A. hernandezii) ; Las Palmas, Pringle 3691 
(NY, US); Rose & Hough 4870 (US); Tancanhuitz, Nelson 4404 (NY, US). 
Chiapas: Huistla, Purpus 6837 (US); San Bartolomé, Collins & Doyle 112 (US); 
Pichucalco, Collins & Doyle 260 (US). Campeche (as Tabasco): Atasta, Rovirosa 
461 (US). GUATEMALA. Alta Verapaz: near Finca Sepacuite, Cook & Griggs 8 
(US). Izabel: Quirigua, Standley 23836 (US); 24054 (NY, US). Solola: Patulul, 
Kellerman 5915 (US). Suchitepéquez: Rio Bravo, Mell 19 (US); Mazatenango, 
Maxon & Hay 3469 (NY, US); Kellerman 5800 (US). EL SALVADOR. La Liber- 
tad: Ateos, Standley 23360 (US). San Salvador: San Salvador, Standley 19150 (NY, 
US) ; 19343 (US) ; 22464 (NY, US); 22674 (US); 23588 (US) = Calderon 81°(NY, 
US); Renson 89 (NY, US); between San Martin and Laguna de Ilopango, Stand- 
ley 22580 (US). San Vicente: San Vicente, Standley 21290 (US) ; 21687 (NY, US). 
BRITISH HONDURAS. El Cayo: El Cayo, Bartlett 13007 (US). Belize: Belize, 
Lundell 4381 (US). NICARAGUA. Granada, Mell s.n., Jan. 23,1925 (NY.) COSTA 
RICA. Guanacaste: Nicoya, Tonduz (Herb. Pittier No.) 13538 (US, type of A. nico- 
yensis). Alajuela: vicinity of San Ramon, Los Loras, Brenes 22677 (NY). CUBA. 
Retiro, cultivated, Wright 2402 (US, photograph and fragment of isotype.of A. 
cubensis ex GH); Habana, Tulipan, introduced, cultivated, or naturalized, Bro. 
Leon 684 (NY, US, fragment); 3690 (NY, US), LESSER ANTILLES. Guade- 
loupe: Basse Terre, introduced and naturalized, Duss 3226 (NY, US); Bailey & 
Bailey 198 (US); Gosier, Stehlé 499 (NY). Martinique: St. Pierre, Jardin des 
Plantes, introduced frrom Mexico, Duss 1144 (NY, US, fragment); Tivoli, Stehlé 
6685 (US). 

2. ACACIA SPHAEROCEPHALA Schl. & Cham. Linnaea 5:594. 1830. A. 
veracruzensis Schenck, Repert. Sp. Nov. 12:362. 1913. A. dolichocephala 
Safford, Jour. Wash. Acad. 5:355. 1915. 


Type: Actopan, Veracruz, Mexico, Schiede & Deppe 684 (B, photograph and 
fragment of type seen). 

Representative specimens seen. MEXICO. Tamaulipas: Tampico, Palmer 133 
(NY, US); Sta. Rafaela, between Tampico and Tula, Berlandier 2145 (US) ; Ciudad 
Madero, King 3990 (NY, US). San Luis Potosi: Tanquian, Tancanhuitz, Cuevas s.n., 
in 1908 (US). Veracruz: Veracruz, Schenck 916 (US, photograph and fragment 
of type of A. veracruzensis ex Herb. Schenck, B?) ; Greenman 87 (NY, US, isotypes 
of A. dolichocephala) ; Miiller 88 (NY); near Tampico, Palmer 448a (US). 


Smithsonian Institution, Washington, D.C. 


1964] ANDERSON: SWALLENIA 201 


LITERATURE CITED 


Britton, N. L. and J. N. Rose. 1928. North American Flora, Vol. 23. 
DE CANDOLLE, A. P. 1825. Prodomus, Vol. 2. 
LEOn, Bro. and Bro. ALAIN [SAUGET, J. S. and E. E. LiocrEr]. 1951. Flora de Cuba. 
Contr. Ocas. Mus. Hist. Nat. Col. “De La Salle,” Vol. 2. 
LinnaEvs, C. 1737. Hortus Cliffortianus. Amsterdam. 
. 1753. Species Plantarum. Stockholm. 
LUNDELL, C. L. 1937. The Vegetation of Petén. Carnegie Inst. Publ. 478. 
SAFFORD, W.E. 1914. Acacia cornigera and its allies. Jour. Wash. Acad. 14:356-368. 
. 1915. New or imperfectly known species of bull-horn acacias. Jour. Wash. 
Acad. 15:355-360. 
ScHENCK, H. 1913. Acaciae myrmecophilae novae. Repert. Sp. Nov. 12:360-363. 
SCHLECHTENDAL, D. von and A. von CHAmisso. 1830. Plantarum Mexicanum... 
... Linnaea 5:554-625. 
STANDLEY, P. C. 1922. Trees and Shrubs of Mexico. Contr. U.S. Natl. Herb., Vol. 23. 
STANDLEY, P.C. and J. A. STEYERMARK. 1946. Flora of Guatemala. Fieldiana Bot., 
Vol. 24. 
WILLDENOW, C.L. 1806. Species Plantarum, Vol. 4. 


NOTES ON THE LEAF EPIDERMIS AND CHROMOSOME 
NUMBER OF SWALLENIA (GRAMINEAE) 


DENNIS ANDERSON 


During April, 1963, the author explored portions of Death, Saline, and 
Eureka valleys in Inyo County, California. Of special interest in Eureka 
Valley is the large sand dune at the south end and the series of endemics 
growing on and near it (Munz and Roos, 1955). Among the most strik- 
ing of these endemics is the monotypic grass genus Swallenia (formerly 
Ectosperma, Soderstrom and Decker, 1963). 

Swallenia alexandrae (Swallen) Soderstrom & Decker forms small, 
dense, somewhat isolated colonies around the lower one-third of the dune 
(fig. 1). The grass is a vigorous, almost bamboo-like plant up to five feet 
tall, with stiff, sharp-pointed, distichous leaves. In the first week of April 
these plants were just coming into bloom and material was fixed in 3:1 
ethyl alcohol-glacial acetic acid for cytological and epidermal studies. 
This material is the basis for the following observations. 

Metcalfe (1960) did not note the presence of bicellular microhairs on 
the abaxial leaf surfaces of Swallenia. Material studied by the present 
author clearly shows the presence of occasional bicellular microhairs 
(fig. 2A). These are very fragile, the delicate terminal cell often collaps- 
ing in preparation of the slide. Other epidermal features observed agree 
with those reported by Metcalfe. 


202 MADRONO [Vol. 17 


Fic. 1. Colony of Swallenia alexandrae growing on sand dune at south end of 
Eureka Valley, Inyo Co., California. Last Chance Range in background. 


B 


Fic. 2. A, Abaxial epidermis of the uppermost culm leaf; B, microsporocyte 
showing ten bivalents. 


The chromosome number of Swallenia (Anderson 2406, UC), previ- 
undetermined, is 2n—20. This number was obtained by acetocarmine 
squashes of anthers. The chromosomes are small, averaging about lw in 


1964] NOTES AND NEWS 203 


length at diakinesis (fig. 2B). Pairing appeared normal in all cells ex- 
amined. 


Division of Biological Sciences, Humboldt State College, Arcata. 


LITERATURE CITED 


MetTcatre, C. R. 1960. Anatomy of the monocotyledons. I. Gramineae. Oxford Univ. 
Press. 


Muwz, P. A. and J. C. Roos. 1955. California miscellany. III. Aliso 3:111-129. 


SODERSTROM, T. R. and H. F. DECKER. 1963. Swallenia, a new name for the California 
genus Ectosperma (Gramineae). Madrofio 17:88. 


NOTES AND NEWS 


SAXIFRAGIA ESCHSCHOLTZII STERNB.—Although not described as such in the more 
recent Alaskan floristic works (E. Hultén, Flora of Alaska and Yukon, 913, 1945; 
J. P. Anderson, Flora of Alaska and adjacent parts of Canada, 290, 1959; I. L. Wig- 
gins and J. H. Thomas, A flora of the Alaskan arctic slope, 240, 1962), this species 
is dioecious. Plants of both sexes are present in a collection made at Point Hope, 
July 3, 1962 (Maxcine Williams 191, 191A, DS, OSC), from which the accompany- 
ing drawings were made (fig. 1). This type of sexual dimorphism is rare in Saxifraga 
and although it was discovered and illustrated long ago by Engler and Irmscher 
(Das Pflanzenreich, IV, 117(1): 164-165, 1916), it seems to be easily overlooked. The 
staminate flowers shrivel and become inconspicuous following anthesis, while the 
ripening pistillate ones retain their staminodia and might be thought to be perfect. 
According to field notes accompanying these collections the two types of flowers 
are quite distinct in form and color at anthesis. The staminate flowers are greenish- 
yellow, except for the red tips of the sepals and the vestigial styles; the sepals are 
sharply recurved. The pistillate flowers are principally red; their sepals are spread- 
ing or cupped upwards. In both sexes the small, narrow petals are yellow (not 
white, roseate, or hyaline, as suggested by Hultén, loc. cit., or Wiggins and Thomas, 
loc: Cit.) 

At Point Hope, S. eschscholtzii grows near sea level, in gravel and thin turf on 
the fringes of the Eskimo settlement. This seems to be an unusual habitat for the 
species, as it contradicts specific statements that the plant is limited to mountainous 
regions (Hultén, Joc. cit.; Wiggins and Thomas, loc. cit.; N. Polunin, Circumpolar 


Fic. 1. Flowers of Saxifraga eschscholtzii, X 10: a, pistillate; b, staminate, with 
petals removed. 


204 MADRONO [Vol. 17 


arctic flora, 260, 1959). A report by Porsild (Rhodora 41:141-183, 1939) lists the 
species among 88 which, in the Bering Sea region, are restricted to hills and moun- 
tains above 1000 feet elevation. Perhaps the explanation of the occurrence at Point 
Hope lies in the suggestion by Wiggins and Thomas (loc. cit.) that S. eschscholtzii is 
a calciphile. The prolonged human occupancy of a site on the coast may create 
edaphic conditions suitable for certain calciphilous species not adapted to the sur- 
rounding tundra.~—KeEeNTon L. CHAMBERS, Department of Botany and Plant Pathol- 
ogy, Oregon State University, Corvallis. 


The following publications are of interest: 


Experimental Studies on Species Relationships in Ceanothus. By Matcoitm A. 
Nos. vili + 94 pp. Carnegie Institution of Washington Publication 623. Washington, 
D.C. 1963. 

California Range Brushlands and Browse Plants. By ARTHUR W. SAMPSON and 
BERYL S. JESPERSEN. viii + 162 pp., illustrated wtih photographs, line drawings, and 
maps. University of California College of Agriculture Manual 33. 1963. $2.00. 

A Flora of Wyoming. By C. L. Porter. Part 1, 39 pp., Part 2, 16 pp. Bulletins 
402 and 404, Agricultural Experiment Station, University of Wyoming, Laramie. 
1962, 1963. 

101 Wildflowers of Acadia National Park. By GRANT and WENONAH SHARPE. 40 
pp., 102 line drawings. University of Washington Press, Seattle. 1963. $1.00. This 
booklet deals with 102 species of flowering plants which are to be found in Acadia 
National Park, Maine. 

A Checklist of Woody Ornamental Plants of California. By MitprEp E. MaTuias 
and ErizaBeETH McCLiInTock. 65 pp. University of California College of Agriculture 
Manual 32. 1963. $0.75. The 1957 addition to the Agricultural Code of California 
stated that ornamental plants offered for sale (with the exception of roses and annual 
and herbaceous perennial plants) had to be labeled with the botanical name. This 
checklist will make it easier for the members of the nursery trade to comply with the 
law and will be of considerable use to gardeners and professional botanists. 

Drawing of British Plants. By SteLLa Ross-Craic. Part XVIII. Compositae (4). 
41 plates. G. Bell & Sons, London. 1963. 10/6. 

Comparative Morphological Studies in Theaceae. By Hsuan KENG. University of 
California Publications in Botany 33:269-384. University of California Press, Berkeley 
and Los Angeles. 1962. $2.50. 

The Morphology and Classification of Some Ceramiaceae and Rhodomelaceae. 
By Max H. Hommersanp. University of California Publications in Botany 35:165— 
366. University of California Press, Berkeley and Los Angeles. 1963. $4.00. 

Selected Botanical Papers. Edited by IrviNc W. KNosBLocu. xiv + 311 pp. Prentice 
Hall, Englewood Cliffs, New Jersey. 1963. $3.95. 

A Flora of San Clemente Island, California. By PETER H. RAVEN. Aliso 5:289-347. 
Rancho Santa Ana Botanic Garden, Claremont, California. 1963. 

On the Flora of the Cascade Mountains. By WM. BripGE CooKE. Wasmann Jour- 
nal of Biology 20:1-—67. Published by the University of San Francisco. 1962. 

A Flora of the Tiburon Peninsula, Marin County, California. By JAVIER PENALOSA. 
Wasmann Journal of Biology 21:1-74. Published by the University of San Francisco. 
1963. Copies may be obtained from the Landmarks Society, P. O. Box 134, Belvedere- 
Tiburon, California. 

A California Flora. By Puitie A. Muwnz in collaboration with Davin D. Keck. 
viii + 1681 pp., 134 figs. University of California Press, Berkeley and Los Angeles. 
Second printing, 1963. $11.50. 

Alpine Wildflowers of Rocky Mountain National Park. By BETTIE E. WILLARD 
and CHESTER O. Harris. 24 pp., illustrated with color photographs. Rocky Mountain 
Nature Association, Estes Park, Colorado. 1963. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume I, 1916-1929 . . . . =. $5.00 
Volume II, 1930-1934. . . . . 5.00 
Volume ITI, 1935-1936 ees eg S100) 
Volume IV, 1937-1938. . . . . 5.00 
Volume V, 1939-1940 (2 2 2 9:00 
Volume VI, 1941-1942 Pe ee ee 60.0) 
Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 Sata 2 sae thes S00 
Volume X, 1949-1950 . . . . = . = 7.00 
Volume XI, 1951-1952 Se Ge eS 00 
Volume XII, 1953-1954 . . . . 8.00 
Volume XIII, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 aw ae UE ht lO.08 
Volume XVI, 1961-1962 . . . . 12.00 


Single numbers: 


VolumesIandIl. =. +. |. “S58 .50 
Volumes II through XIV. . ..... 1.00 
Volume XV°-< °°. 04% 4s ee 1.25 
Volume XVi 2 eceiiiget sew ae 1.50 


The subscription price of MApRONo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


fo 


ON 


MADRONO 


[oe Dylan 


| VOLUME 17, NUMBER 7 JULY, 1964 
| Contents 

PAGE 
THE POLLEN GRAIN MORPHOLOGY OF COLLOMIA AS A 

| Taxonomic Toot, Alfred R. Loeblich, III 205 


A NEw SPECIES OF PINE FRoM Mexico, Egon Larsen 217 


Two NEw SPEcIES RELATED TO CLARKIA UNGUICU- 
LATA, Frank C. Vasek 219 


TAXONOMIC NOTES ON THE CHRYSOTHAMNUS VISCIDI- 
FLORUS COMPLEX (ASTEREAE, COMPOSITAE), 
Loran C. Anderson Dae 


Davip DoUGLAS AND THE DIGGER PINE: SOME QUES- 
TIONS, James R. Griffin 227 


BARK PHOTOSYNTHESIS IN OcoTILLo, H. A. Mooney 
and B. R. Strain 230 


REviEws: Stewart L. Udall, The Quiet Crisis (John 
Pelton); Forest Shreve and Ira L. Wiggins, Vegeta- 
tion and Flora of the Sonoran Desert 
(Reed C. Rollins) 233 


NoTES AND NeEws: EDWIN BINGHAM COPELAND; 
NOTES ON THE F Lora oF Arizona. III, Charles T. 
Mason 235 


A WEST AMERICAN JOURNAL OF BOTANY 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madronio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


EDGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN Benson, Pomona College, Claremont, California 
HERBERT F. CopELAND, Sacramento College, Sacramento, California 
Joun F. Davipson, University of Nebraska, Lincoln 
WALLACE R. ErnsT, Smithsonian Institution, Washington, D.C. 
Mitprep E. Matutas, University of California, Los Angeles 
ROBERT ORNDUFF, University of California, Berkeley 
Marion OwnBEY, Washington State University, Pullman 
REED C. RoLiins, Gray Harbarium, Harvard University 
Ira L. Wiccins, Stanford University, Stanford, California 


Editor—JoHn H. THoMAS 
Dudley Herbarium, Stanford University, Stanford, California 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: G. Ledyard Stebbins, Department of Genetics, University of California, 
- Davis. First Vice-President: Annetta Carter, Department of Botany, University of 
California, Berkeley. Second Vice-President: Carl W. Sharsmith, Department of 
Biological Sciences, San Jose State College. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding 
Secretary, Margaret Bergseng, Department of Botany, University of California, 
Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State 
College. 


1964] LOEBLICH: COLLOMIA 205 


THE POLLEN GRAIN MORPHOLOGY OF COLLOMIA 
AS A TAXONOMIC TOOL 


ALFRED R. LOEBLICH, III 


There is a rich variety of pollen grain forms within the family Pole- 
moniaceae, and a species is commonly recognizable by the pollen grain 
characters. Furthermore, these grains are abundant—each plant produc- 
ing many more grains than leaves or fruits—and the grains are more 
readily preservable as fossils because of their small size and resistance to 
deterioration. Hence a well illustrated and comparative account of the 
modern pollen morphology may lead to the recognition of fossil members 
of the Polemoniaceae. 

There have been only three reports of fossil remains of the family 
Polemoniaceae, a Pleistocene fruit of a Phlox (Chaney and Mason, 1936), 
a Pleistocene Polemonium pollen grain (Godwin, 1956), and a subfossil 
pollen grain of Collomia (Hafsten, 1960). The lack of paleontological 
evidence renders suspect the validity of the conclusions that have been 
made concerning the evolution of this family (Wherry, 1944; Grant, 
1959). It is questionable whether megafossils will ever yield much in- 
formation about the evolutionary trends within this family, since preser- 
vation of recognizable fossil remains of these herbaceous plants or of 
their fruits or flowers is unlikely except under unusual circumstances. 
If a more abundant and more readily preserved part of the plant, such 
as the pollen, could be found to characteristic of a species or genus, paleo- 
botanists and palynologists would be aided in recognizing fossil occur- 
rences of the Polemoniaceae. 

When used in conjunction with other features of a plant, pollen char- 
acteristics should aid in its taxonomic classification. According to Grant 
(1959, p. 35), “The Polemoniaceae is a taxonomically complex family 
in which generic and specific divisions have frequently been the subject 
of confusion and dispute.” As will be evident from this study of the genus 
Collomia Nutt., pollen morphology may be as useful a taxonomic tool 
for the neontologist as it is for the paleontologist. 

Yet almost as sparse as the information of the fossil record of the Pole- 
moniaceae, is that concerning pollen morphology of the included genera. 
To date only four significant references have been made to the pollen of 
the genus Collomia. Erdtman (1952) briefly described and figured that 
of one form, C. grandiflora f. axillaris (Nels.) Wherry. Hafsten (1960) 
figured a subfossil pollen grain and identified it as the form illustrated by 
Erdtman. In 1959 Grant made a general survey of the pollen grain types 
within the family, but gave neither specific descriptions nor comparisons. 

The most comprehensive study of the pollen of the Polemoniaceae has 
been presented by Marticorena (1961), who was concerned only with the 
Chilean species, and treated both of the known South American species 


Maprono, Vol. 17, No. 7, pp. 205-236. July 23, 1964. 


206 MADRONO [Vol. 17 


in Collomia. Thus, to date, the pollen has been studied only of C. grandi- 
flora f. axillaris and the two Chilean species out of the total of 14 species 
in Collomia. No comparative studies of the pollen of all the species of this 
genus have yet been published. 


MATERIAL AND METHODS 

Poilen was obtained from University of California herbarium speci- 
mens and was acetolyzed according to the micromethod of Punt (1962). 
The collectors and collection numbers and herbarium numbers of the 
specimens are given in the graph of equatorial diameters (fig. 1). Data 
for Fig. 1 were obtained by measuring 50 fully expanded pollen grains 
from each herbarium sheet. Grains whose orientation on the prepared 
slide allowed a polar diameter measurement were scarce, therefore fewer 
polar than equatorial diameter measurements were obtained. The termi- 
nology used in the description of species follows that of Erdtman (1952). 

DESCRIPTIONS OF THE POLLEN GRAINS IN SPECIES OF COLLOMIA 

C. biflora (Ruiz et Pav.) Brand. Average equatorial diameter: 57.9 p, 
range: 50.8 to 63.5; average polar diameter: 44.5; average ratio of 
equatorial to polar diameter: 1.30:1 (subspheroidal, suboblate). Steph- 
anoporate, (7)—8—(9) apertures, composite, ovate, length in nexine 7.7 p, 
width 3.3 u. Exine 2.9 in thickness, sexine only slightly thicker than 
nexine, the nexine swelling at the apertural margins; surface of sexine 
with subparallel and irregular ridges locally breaking up into reticula- 
tions. Figs. 20, 21. 

C. cavanillesta Hook. & Arn. Average equatorial diameter: 56.2 p, 
range: 53.3 to 66.7 4; average polar diameter: 48.2 4; average ratio of 
equatorial to polor diameter :1.17:1 (subspheroidal, suboblate) . Stephan- 
oporate, (6)—(7)—8—-(9) apertures, composite, ovate, length in nexine 
7.7, width 5.5 w. Exine 3.5p in thickness, sexine 1.5 times as thick as 
nexine, nexine swelling at apertural margins. Sexine pattern composed of 
continuous meandriform ridges. Figs. 22, 23. 

C. debilis (S. Wats.) Greene ssp. debilis. Average equatorial diameter: 
50.3 w, range: 40.6 to 58.2; average polar diameter: 41 u; average 
ratio of equatorial to polar diameter: 1.23:1 (subspheroidal, suboblate). 
Stephanoporate, (6)—7—(8)—(9) apertures, composite, ovate. Aperture 
measurements (for species of C. debilis sensu lato): length in nexine 
7.7-8.8 », width 4.4—5.5 ». Exine 3.3 in thickness, sexine 1.5 times as 
thick as nexine, nexine swelling at apertural borders. Sexine pattern con- 
sists of subparallel ridges, whose orientation resembles the pattern of 
lines of force in a magnetic field whose poles are in the position of the 
apertures. Figs. 30. 31. 

C. debilis ssp. ipomoea (Pays.) Wherry. Average equatorial diameter: 
51.9, range: 45.6 to 64.34; average polar diameter: 42.7; average 
ratio of equatorial to polar diameter: 1.22:1 (subspheroidal, suboblate). 
Stephanoporate, (7)—8—(9) apertures, composite, ovate. Exine 3.3 » in 
thickness, sexine 1.5 times as thick as nexine, nexine swelling at the aper- 


1964] LOEBLICH: COLLOMIA 207 


STEPHANOPORATE GRAINS 


30 35 40 45 50 55 GO G5 TO 
C. tenella ! ! | | \ 1 \ \ ' 


U.c.165 G85 
Nelson and Macbride 1240 
C. macrocalyx 
U.C.8TSTOT 
Peck 21721 
C. Anearis Le 
U.C.9655390 
‘Mason 14396 
C. mazama 
U.C.53596 
Coville ARS Applegate 315 
C. AHvers/folia ae) . see 
uc. A 191132 
Mason 14519 
C. dversifolia ssincumaiaiieiah 
U.C.963990 
C. larsens a ' 
U.C.M094326 
C. cbs45 ssp. trifiola mae ' 
U.C. 968 
Hieeneote and Muhlick 11148 Sees 
C eebils 83p.c%bilis re 
U.C.TO0883 
Hitchcock and Muhlick i426 
C. eebsiis ssp. jpomoea * mt — 
U.C.eT9864 
C. rawsoniana aaah E 
U.C.1T435T7TS 
Brandegee 237 
C. granc/f/ora : 
U.C. 197855 
Tracy 278 
C. grandiflora 
U.C. M18 TO32 
Hesse 2691d 
C. &/#/ora 
UC. Be3ITSC 
Werdermann /1392 
& cavani/les// 


UC. Mis5e50 
wtchison PANTOPORATE GRAINS 


C. Reterophy//a 
U.C. 582686 
Tracy I5S296 

C. tracy, 


U.C.9T6535 
Tracy 14466 


C. trnctorva 


U.C. 133632 
Tracy 2712 


bd Only 25 measurements were made. 


Fic. 1. Graph showing equatorial pollen diameters of the various species of 
Collomia. 


tural margins. Sexine pattern in equatorial view of grain consists of sub- 
parallel radiating ridges whose orientation resembles the pattern of lines 
of force in a magnetic field having the poles at the ends of the apertures. 
Figs. 26, 27. 

C. debilis ssp. trifida (Pays.) Wherry. Average equatorial diameter: 
47.9 wu, range: 43 to 58; average polar diameter: 40y; average ratio 


208 MADRONO [Vol. 17 


of equatorial to polar diameter: 1.20:1 (subspheroidal suboblate). Steph- 
anoporate, (7)—8—(9) apertures, composite, ovate. Exine 3 » thick; sex- 
ine only slightly thicker than nexine, nexine swelling at the apertural 
borders. Sexine pattern composed of ridges subparallel in an orientation 
resembling the pattern of lines of force in a magnetic field hav:ng poles 
at the ends of the apertures. Figs. 28, 29. 

C. diversifolia Greene. Average eaua‘orial diameter: 49.6. range: 38 
to 61; average po'ar diameter: 43 4; average ratio of equatorial to 
polar diameter: 1.16:1 (subspheroidal, suboblate). Stephanoporate, (5)- 
6—(7) apertures, composite, ovate, length in nexine 6.6, width 4.4 pu. 
Exine 3.3 u thick, sexine 1.5 times as thick as the nexine, nexine swe'ling 
at apertural margins. Sexine pattern composed of slightly continuous 
subparallel ridges in a swirling design. Fig. 11. 


C. grandiflora Dougl. ex Lindl. Average equatorial diameter: 56.4 1, 
range: 50.7 to 61m; average polar diameter: 43.6; average ratio of 
equatorial to polar diameter: 1.29:1 (subspheroidal, suboblate). Steph- 
anoporate, (7)—8—(9) apertures, composite, ovate, length in nexine 11 pn, 
width 4.4 u. Exine 3.3 4 thick, sexine 1.5 times as thick as the nexine, 
nexine swelling at apertural margins. Sexine pattern composed of beaded 
ridges radiating from the poles and continuing as paral'el ridges along 
the aperture margins. Figs. 8, 9. 

C. heterophylla Hook. Average diameter: 44.3 uw, range: 38 to 50.7 u 
(spheroidal). Pantoporate, 10-16 apertures, simple, circular, diameter in 
nexine 3 ». Exine 3.3 » in diameter, sexine 1.5 times as thick as the nexine, 
nexine swelling at apertural margins (fig. 32). Sexine ornamentation 
composed of sharply curved ridges tending to a somewhat reticulate pat- 
tern. Figs. 6, 7. 

C. larsentt (Gray) Pays. Average equatorial diameter: 47.7 u, range: 
38 to 544; average polar diameter: 40.3 1; average ratio of equatorial 
to polar diameter: 1.18:1 (subspheroidal, suboblate). Stephanoporate, 
(6)—7—-8 apertures, composite, ovate, length in nexine 8 p, width 4.4 pu. 
Exine 2.6 » thick, sexine only slightly thicker than nexine, nexine swelling 
at apertural margins. Sexine pattern composed of subradiating ridges 
passing parallel to one another around the apertures. Fig. 10. 


C. linearis Nutt. Average equatorial diameter: 45.6 u, range: 40.5 to 
50.8 wu; average polar diameter: 40.3 1; average ratio of equatorial to 
polar diameter: 1.31:1 (subspheroidal, oblate spheroidal). Stephano- 
porate, 7-8—(9)—(10) apertures, composite, ovate, length in nexine 7.7 p, 
width 3.3 ». Exine 3.6 » thick, sexine twice as thick as the nexine, nexine 
swelling at apertural margins (fig. 33). Sexine ornamentation composed 
of sharply curved ridges tending to a somewhat reticulate pattern. Figs. 
14,15. 

C. macrocalyx Leib. ex Brand. Average equatorial diameter: 44.1 p, 
range: 38 to 50.8 »; average polar diameter 35.2 w; average ratio of equa- 
torial to polar diameter: 1.25:1 (subspheroidal, suboblate). Stephano- 


1964] LOEBLICH: COLLOMIA 209 


Fics. 2—7. Pollen grains of C. tinctoria, tracvi, and heterophylla; 2, 3, sexine pat- 
tern and arrangement of apertures of C. tinctoria, both x 640; 4, 5, sexine pattern 
and distribution of apertures of C. tracyi, x 1600 and 640, respectively; 6, optical 
section of C. heterophylla, apertures appear as light areas in center and one is shown 
in side view at the lower right periphery, * 1600; 7, surface view of C. heterophylla 
showing sexine ornamentation and scattered apertures, < 640. 


porate, (7)—8-(9) apertures, composite, ovate, length in nexine 5.5 u, 
width 2.2 ». Exine 3 » thick, sexine 1.5 times thicker than nexine, nexine 


210 MADRONO [Vol. 17 


Fics. 8-13. Pollen grains of C. grandiflora, larsenii, diversifolia, and rawsoniana; 
8, 9, polar views showing sexine ornamentation of C. grandiflora, x 1600 and 640 
respectively; 10, polar view showing sexine ornamentation of C. larsenii, x 640; 
11, polar view showing sexine ornamentation of C. diversifolia, < 640; 12, polar 
view showing sexine pattern of C. rawsoniana, X 640; 13, equatorial view of C. raw- 
soniana showing pattern of the sexine around two apertures, xX 1600. 


swelling at apertural margins. Sexine pattern composed of rather contin- 
uous subparallel ridges in a swirling design. Figs. 16, 17. 
C. mazama Cov. Average equatorial diameter: 46.6», range: 43.2 to 


1964 | LOEBLICH: COLLOMIA PANG 


Fics. 14-19. Pollen grains of C. linearis, macrocalyx, and mazama; 14, 15, poiar 
views showing sexine pattern of C. linearis, —K 1600 and 640 respectively; 16, 17, 
polar views showing sexine pattern of C. macrocalyx, both « 640; 18, 19, polar views 
showing sexine pattern of C. mazama, < 640 and 1600 respectively. 


50.8 «; average polar diameter: 38 «; average ratio of equatorial to polar 
diameter: 1.23:1 (subspheroidal, suboblate). Stephanoporate (7)—8—(9) 
apertures, composite, ovate, length in nexine 5.5, width 3.3 yu. Exine 
2.6 thick, sexine as thick as the nexine, nexine swelling at apertural 


PAD MADRONO [Vol. 17 


Fics. 20-25. Pollen grains of C. biflora, cavanillesii, and tenella; 20, slightly tilted 
polar view showing sexine pattern of C. biflora, x 640; 21, polar view showing 
sexine pattern of C. biflora, < 1600; 22, 23, polar views showing sexine pattern of 
C. cavanillesii, < 640 and 1600 respectively; 24, 25, polar views showing sexine 
ornamentation of C. tenella, both * 640. 


margins. Sexine pattern composed of rather subparallel, continuous ridges 
in a swirling design. Figs. 18, 19. 

C. rawsoniana Greene. Average equatorial diameter: 50.5 uw, range: 43 
to 56 w; average polar diameter: 41.2 w; average ratio of equatorial to 
polar diameter: 1.23:1 (subspheroidal, suboblate). Stephanoporate, 7--8— 


1964] LOEBLICH: COLLOMIA 213 


(9) apertures, composite, ovate, length in nexine 8.8 », width 3.0 pw. Exine 
3.6 p thick, sexine 1.5 times as thick as the nexine, nexine swelling at 
apertural margins (fig. 34). Sexine pattern composed of subparallel ridges 
in an orientation resembling the pattern of lines of force in a magnetic 
field having poles at the ends of the apertures. Figs. 12, 13. 

C. tenella Gray. Average equatorial diameter: 39.5 p, range: 33 to 
47.4 »; average polar diameter: 32 w; average ratio of equatorial to polar 
diameter; 1.23:1 (subspheroidal, suboblate). Stephanoporate, (6)—7—(8) 
epertures, composite, ovate, length in nexine 5.5 », width 2.2 uw. Exine 
2.6 » thick, sexine twice as thick as nexine, nexine swelling at the aper- 
tural margins. Sexine pattern composed of rather continuous subparallel 
ridges in swirling design of meandriform, curving ridges. Figs. 24, 25. 

C. tinctoria Kell. Average diameter: 47.7 p, range: 43.2 to 50.8 j 
(spheroidal). Pantoporate, 19 to 24 apertures, simple, circular, diameter 
in nexine 3.9 w. Exine 3.3 uw thick, nexine thinnest in area between pores 
where sexine is thickest (fig. 35). The nexine swells to 1.3 » at the aper- 
tural margins. Each aperture is at the center of a small, deeply sunken 
pentagonal or hexagonal area, bordered by a broad radially grooved 
ridge, the slightly irregular radial grooves also extending across the poly- 
gonal area but less prominent there than on the marginal ridges. Figs. 
Die. 

C. tracyt Mason. Average diameter: 44.2 uw, range: 38 to 50.8 w (spher- 
oidal). Pantoporate, apertures 17 to 28, simple, circular, diameter in 
nexine 3.9 uw. Exine, 3.3 m» thick, the nexine thinnest in area between 
apertures where the sexine is thickest. The nexine swells at the apertural 
margins. Each aperture is at the center of a small, deeply sunken penta- 
gonal or hexagonal area, bordered by a broad radially grooved ridge, the 
slightly irregular radial grooves also extending across the polygonal area 
but less prominent there than on the marginal ridges. Figs. 4, 5. 


KEY TO SPECIES OF COLLOMIA BASED ON POLLEN MORPHOLOGY 
Pantoporate. 
Apertures at center of sunken pentagonal or hexagonal areas. 
C. tracyi, C. tinctoria 
Apertural margin flush, not sunken; sexine with surface of narrow vermiform 
TCL Oe oma Se sete web foe. Me tee gsees, pe ace ete t a t W eas C. heterophylla 
Stephanoporate. 
Sexine with subparallel ridges whose orientation resembles the pattern of lines of 
force in a magnetic field having poles at the ends of apertures. 
C. rawsoniana, C. grandiflora, C. larsenii, C. debilis 
Sexine with flowing or swirling meandriform ridges. 
Sexine pattern rather continuous with subparallel ridges in a swirling design. 
C. diversifolia, C. mazama, C. tenella, C. macrocalyx 
Sexine ornamentation of numerous sharply crested, narrow ridges of irregular 
orientation. 
Pollen oblate spheroidal .......................00..... ee eee C. linearis 
Pollen suboblate. 
Sexine with subradial and irregular ridges locally breaking up into reticu- 
| FEV. (0) 00) ae ee Sore Re oP OE er EOD aE Bare ERT Is C. biflora 
Sexine with continuous, meandriform ridges.................0..0......... C. cavanillesii 


214 MADRONO [Vol. 17 


RELATION OF POLLEN MORPHOLOGY TO TAXONOMY 


In stephanoporate pollen grains the polar diameter is always less than 
the equatorial diameter. The range of the ratios of equatorial to polar 
diameters is 1.13:1 to 1.30:1. On the basis of these ratios Erdtman 
(1952) would consider the stephanoporate grains all to be subspheroidal. 
Within this category all but C. linearis are suboblate; C. linearis is oblate 
spheroidal. The three species with pantoporate pollen grains are, of 
course, sperical. 


Fics. 26-31. Pollen grains of C. debilis; 26, polar view showing sexine ornamen- 
tation of ssp. 7pomoea, xX 640; 27, view of area slightly above apertures of ssp. 
ipomoea, X 640; 28, 29, polar views of sexine ornamentation of ssp. trifida, both 
x 640; 30, 31, polar views of sexine crnamentation of ssp. debilis, both x 640. 


The apertures of pantoporate grains are circular and simple, whereas 
the apertures of stephanoporate grains are composite and ovate. The 
stephanoporate apertures are intermediate between pori (circular aper- 
tures) and colpi (apertures at least twice as long as they are wide) (Erdt- 
man, 1952). In the nexine the apertures are ovate, but the sexine projects 
over each side of the ovate opening, narrowing it longitudinally to an 
elongated subrectangular opening in external view. The sexine in most 
species is composed of small pegs with the tops fused to form ridges. These 
ridges are seen as the sexine pattern in the external view (figs. 13, 27). 
The nexine in all the species swells to some extent at the margins of the 
apertures. This swelling produces light circles around the apertures (figs. 
2,334, 5, 13,20, 279). 

There is some degree of correlation between flower length and grain 
size (small-flowered plants tend to have small pollen grains), but there 
is less correlation between grain size and plant size. 


1964 | LOEBLICH: COLLOMIA 215 


A comparison of the species groups based on pollen grain characteris- 
tics as found in the above key and the accepted sections of this genus 
based on other features should show the relation of pollen morphology to 
taxonomy. The following sections and their species are currently accepted 
(Wherry, 1944; Grant, 1959): 

Collomiastrum Brand: C. mazama; C. rawsoniana; C. larsenii; C. 
debilis. 

Courtoisia (Reichenb.) Wherry: C. heterophylla; C. diversifolia. 

Collomia: C. cavanillesu; C. grandiflora; C. linearis; C. macrocalyx; 
C. tinctoria; C. tenella; C. tracyt; C. biflora. 

Some discrepancies are immediately evident in a comparison of these 
sections and the key to species based on pollen morphology. The pollen 
grain morphology seems to indicate that the section Collomiastrum stands 
well as a unit except for the inclusion of C. mazama, whose grains are 
somewhat smaller and whose sexine pattern is not as striate as C. grandi- 
flora. Although C. grandiflora has slightly larger grains, it seems to fit 
better into this section because its sexine ornamentation is almost iden- 
tical to that of the other species of Collomiastrum. 

The section Courtoisia is not at all homogeneous according to pollen 
morphology. Collomza diversifolia was formerly not recognized as differ- 
ing from C. heterophylla, but the pollen grains of C. heterophylla are 
pantoporate whereas those of C. diversifolia are stephanoporate, and fur- 
thermore, the sexine patterns are different, strongly supporting their 
separation into two species. The sexine ornamentation of C. heterophylla 
closely resembles that of C. linearis, but spherical shape, pantoporate 
apertures and sexine pattern of C. heterophylla differ from other Collomia 
species, even including C. diversifolia. Thus, Wherry’s statement (1944) 
that C. diversifolia deserves perhaps only subspecies status under C. het- 
erophylla is not supported by pollen morphology. 


Three species of the Collomia section, 1.e., C. cavanillesu, C. biflora, 
and C. linearis, have very similar pollen morphology, but the other species 
assigned to this section are dissimilar in differing degrees. C. tenella and 
C. macrocalyx do not have the sharply curved ridges in the sexine of C. 
cavanillesii, C. biflora, and C. linearis. C. tenella and C. macrocalyx 
have a sexine pattern similar to C. diversifolia and C. mazama. C. tenella 
has a smaller ratio of equatorial to polar diameter and a smaller grain 
size than does C. macrocalyx; furthermore, the number of apertures is 
usually smaller. Some grains of C. tenella have six apertures, but never 
nine, while C. macrocalyx never has six apertures, but sometimes has nine. 


The remaining species of the Collomia section, C. tinctoria and C. 
tracyi, present an interesting problem. The pollen grains of these two 
species do not resemble any other pollen types in this genus, except that 
they have a pantoporate arrangement of simple apertures in common with 
C. heterophylla. However, the sexine pattern of C. heterophylla shows 
no resemblance to those of C. tinctoria and C. tracyi. The peculiar pollen 


216 MADRONO [Vol. 17 


a ; 


Fics. 32-35. Diagrammatic optical sections of apertural regions; 32, C. hetero- 
phylla; 33, C. linearis; 34, C. rawsoniana; 35, C. tinctoria. The upper layer is the 
sexine: the lower, the nexine. 


morphology of the latter two species leaves doubtful their inclusion in 
this genus. 

It is perhaps more than coincidence that C. linearis, one of the two 
North American species with the most southerly range, more closely re- 
sembles the two South American species (C. biflora and C. cavanillesii) 
in potlen sexine pattern than does any of the other North American 
species. Pollen of C. grandiflora, the second southward ranging North 
American species, most closely approximates that of the two South Ameri- 
can species in size and shape, and has a ratio of equatorial to polar diame- 
ter almost identical to that of C. diflora, but this species differs in sexine 
pattern. Thus the geographically closer species also show greater agree- 
ment with respect to their pollen grain morphology. 

Possibly the similarities and dissimilarities in pollen morphology of the 
various species might suggest a restudy or reevaluation of additional 
characters as a step in determining more closely the natural relationships. 


Department of Botany, University of California, Berkeley 
LITERATURE CITED 


CuHaney, R.W. and Mason, H. L. 1936. A Pleistocene flora from Fairbanks, Alaska. 
Am. Mus. Nov. 887:1-17. 

ErpTMAN, G. 1952. Pollen morphology and plant taxcnomy. Angiosperms. Almqvist 
and Wiksell, Stockholm. 

Gopwin, H. 1956. The history of the British flora. Cambridge Univ. Press. 

Grant, V. 1959. Natural history of the Phlox family. Martinus Nijhoff, The Hague. 

HarstTENn, U. 1960. Pleistocene development of vegetation and climate in Tristan da 
Cunha and Gough Island. Arbok. Univ. Bergen, Mat.-Naturv. ser. n. 20:1-48. 

MarTIcorENA, C. 1961. Morfologia de los granos de polen de las Polemoniaceae 
Chilenas. Gayana Bot. 2:1-12. 

Punt, W. 1962. Pollen morphology of the Euphorbiaceae with special reference to 
taxonomy. Wentia 7:1-116. 

Wuerry, E.T. 1944. Review of the genera Collomia and Gymnosteris. Am. Midl. 
Nat. 31:216-231. 


1964] LARSEN: PINUS 217 
A NEW SPECIES OF PINE FROM MEXICO 
EGON LARSEN 


Pinus martinezii Larsen, sp. nov. Arbor modica vel magna mole, 
altitudine m 25 vel pluri, diametro cm ca 60 (supra terram metros 1.30). 
Maturis coma rotundata est et rarior; veterum ramos crassos, horizon- 
tales-gravidos, raro tamen satarum sine ordine vel singulos vel involutos 
usque ad terram productos invenies. Foliorum bases praecoce quodam 
et deciduo epidermati sunt tectae, quo ramuli asperrimi fiunt. Hae in 
veteribus ramis et in superiore stipite velut scabies permanent. Cortex, 
quae in recentissimis pruinosa et quasi viridis conspicitur, in veteribus 
ramulis in colorem brunneum sed non saturatum mutatur. In imo trunco 
necnon in veteribus ramis cortex crassa et aspera et altis in longitudinem 
positis fissuris penetrata, in quibus sane flava conspicitur. Cortex omnino 
mitis est et squamosa. Folia in fasciculis quinis, senis, septenis, interdum 
octenis, plerumque senis invenies, longitudine cm 20-28, plerumque 23, 
dura, recta, intus glauci colores qui postea in viridem obscurum mutatur, 
marginibus aciformis et serratis, dentibus parvis et frequentibus in- 
structa; in omnibus faciebus stomata, quorum tres vel quattuor ordines 
in superficie superiore adsunt, in aliis sex vel septem. Hypodermatis 
crassitudo variata, quod dum saepe in chlorenchyma alte penetret, 
musquam tamen ad endoderma pervenit. Canales resiniferos tres et 
quidem medios habent. Exteriores endodermatis parietes crassissimi. 
Sunt fasces fibro-vasculares duo, conferti sed satis distincti. Vaginae 
recentium fasciculorum co'ore brunneo sunt, qui postea in cinereum 
mutatur, et longitudinem cm 1.2—2.5 habent. Gemmae conoides, colore 
brunneo non saturatae; non resiniferae. Strobili oblongi vel conoides, 
subsymmetricales, leniter incurvati et reflexi, coloris brunnei non saturati 
et in pedunculis longitudinis cm ca 1.2 positi, mensibus decembre et 
lanuario maturescunt, maturatique aperiuntur. Semidecidui sunt, nam 
etiam cum strobilus cecidit, pedunculi eius cum paucis squamis adhuc 
in ramo manent. Strobilorum squamae durae satis ac validae, longitudine 
cm ca 2.5 vel paulo maiores, latitudine cm ca 1.2 ap‘ce rotundato et 
aliquando inaequali instructae. Apophyse on forma est varia: nunc fere 
plana, nunc protuberans, aliquando denique reflexa. Umbones sublati 
et colore quam reliqua apophysis nigriore distincti aculeo deciduo et 
invalido armati sunt. Semina flava cinerascentia nigro maculata plerum- 
que, sed quaedam co'lore aliquo modo rubro aequaliter tecta sunt. 
Pleraque conspicuis in longitudinem factis canalibus notata sunt. Semi- 
num fere 48000 pro kg. Alae seminum sine hamo basali, qui totam seminis 
peripheriam amplectitur, longae cm 1.2—1.9 sunt. 

Type: Approximately 4 mi S of Paracho between km markers 45 and 48 on road 
from Carapan to Uruapan, Michcacan, Mexico, Lat 19° 17’ N, Long 102° 04’ W, 
altitude 7800 ft, H. V. Hinds & E. Larsen 65, Jan. 10, 1960 (herbarium of the Forestry 


and Timber Bureau, Canberra, no. A.F.S. 13084/A—holotype; CANB, no. 133708; 
Forest Research Institute, Rotorua, New Zealand, no. F.R.J. 60/1277; MEXU; UC). 


218 MADRONO [Vol. 17 


This species is named in honor of Professor Maximino Martinez of 
the University of Mexico, world famous authority on Mexican conifers. 
It was collected for the New Zealand Forest Service during a visit 
to Mexico during 1959-1960 while I was in the employ of the Forest 
Research Institute, Rotorua. 

In general appearance P. martinezi resembles P. douglasiana Mart. 
from which it differs in having its needles predominantly in fascicles of 
six, by having hypoderm which never extends to the endoderm, by its 
cone-peduncle which, with a few scales, remains on the branch when 
the cone falls, and by the rough bark on young trees and upper part 
of the stem. 

In addition to P. douglasiana it resembles P. michoacana Mart. var. 
quevedoi Mart. in several respects, the six needle fascicles in particular, 
but differs in having much shorter cones and needles, by the glaucous 
inner surface of the needles, and by having only three resin canals; also 
the buds are not resinous. From P. durangensis Mart. which is the only 
other species with six needles per fascicle it differs in having glaucous 
needles 8-11 in. long, while those of P. durangensis are non-glaucous 
and only 4-9 in. Pinus durangensis has short but strong and persistent 
cone scale prickles, while those of P. martinez are very small, weak, 
and deciduous. 

In the Uruapan area the species grows in association with 
P. montezumae Lamb., P. teocote, C. & S., P. leiophylia Benth, P. 
pseudostrobus Lindl., and Quercus species. It generally occupies the drier 
ridges where it equals P. montezumae in growth, while P. pseudostrobus 
is found mainly on the slightly better sites. It is easily distinguished 
from these species by its stiff, erect, dark green to glaucous needles. 

The site is rather poor, forming the higher part of a large undulating 
plateau. The soil is loose, coarse volcanic ash containing many particles 
and large outcrops of vesicular lava. The forest is mostly second growth 
of varying ages, semi-open with a dense cover of grass and tussock in 
many places. It is 2800 ft higher than Uruapan where the mean annual 
rainfall is 65 in and the mean annual temperature 67°F with the means 
for the coldest and warmest months 61°F and 72°F respectively. 

The trees are generally of good form and branches, while heavy on 
open-grown trees, respond well to close spacing to give long, clean 
boles. The rate of height growth is approximately 2 ft per year. 


Forest Research Institute, Canberra, A. C. T., Australia 


1964 | VASEK: CLARKIA 219 


TWO NEW SPECIES RELATED TO CLARKIA UNGUICULATA 
FRANK C. VASEK 


Clarkia ungutculata is common and widespread in the coast ranges 
and Sierra Nevada foothills of California. It is morphologically quite 
variable with regard to leaf size and shape, bud shape, sepal color, petal 
shape, size. and color, and pubescence type (Lewis and Lewis, 1955). A 
portion of this variation has been resolved as a closely related species, 
C. exils (Lewis and Vasek, 1954; Vasek, 1958; 1960). The latter occurs 
in the low elevation foothill wood!and of the southern Sierra Nevada of 
Kern and Tulare counties and is distinguished from C. unguiculata by 
its more slender habit, narrower leaves, smaller flowers, by the position 
of the stigma near the anthers which facilitates self-pollination, by its 
more rapid development, and by the absence of long hairs on the ovary 
and calyx. 

Plants from the inner coast ranges of Alameda, San Benito, and west- 
ern Fresno counties have been included in C. unguiculata but were noted 
to have leaves consistently narrower than usual (Lewis and Lewis, 1955). 
The same plants also lack long hairs on the ovary and calyx (e.g. Corral 
Hollow, Eastwood & Howell 5291, POM), and appear to have flowers 
somewhat smaller than usual. All of these characteristics suggest a mor- 
phological similarity to C. exilis. Consequently, an investigation was ini- 
tiated to deterine the relationships of the inner coast range plants with 
C. unguiculata and C. exilis. Plants from a population near Springville 
in Tulare County were included in the investigation because of their 
somewhat narrow leaves and their lack of long hairs on the ovary and 
calyx. 

This investigation led to the conclusion that four distinct, morphologi- 
cally recognizable entities occur, each differentiated from the others by 
extensive chromosome repatterning, hybrid sterility, and in the case of 
sympatric distributions, by reduced crossability (Vasek, 1964). All four 
species are morphologically similar and closely related. In all probability, 
C. exilis, C. tembloriensis, and C. springvillensis have each evolved inde- 
pendently from C. unguiculata. The details of the relationships, and a 
distribution map, are published elsewhere (Vasek, 1964). 

I am grateful to Harlan Lewis for criticizing the manuscript and to 
Douglass S. Parker for assistance with the Latin diagnoses. 

All four species are endemic to California and may be identified with 
the aid of the following key. 


Some long hairs present on the ovary and calyx; leaf width %4 to % the length; 
petal width 4% to 34 the length; leafy bract width, % to %8 the length; style 
usually well exserted; widespread in Coast Ranges and Sierra Nevada foothills 
from Lake and Plumas counties to San Diego County; usually in oak wood- 
1s 01 01.0 ace a a pe aPC eo On a eT eI PE C. unguiculata 

Long hairs absent; leaf width less than 4 the length; petal width usually less than 
Y% the length; leafy bract width usually less than 1% the length. 


220 MADRONO [Vol. 17 


Style usually well exserted; sepals dark red-purple; petals usually with a large, 
dark, red-purple spot at the base of the blade; often with 5 or 6 flowers 
open on one stem at the same time; near Springville, Tulare County; in 
foothill woodland: 2.225 ee en ee ee C. springvillensts 


Style seldom well exserted; sepals mostly green or only slightly reddish; petal spot, 
if present, small and well defined or if large, not sharply defined; usually 
only 1-3 flowers open on one stem at the same time. 


Style equalling or only slightly exceeding the anthers; leaves usually bright 
green; petals pink, with or without a small purple spot at the base of blade, 
or white; Sierra Nevada foothills, southern Tulare and northern Kern 
counties, in or at the low elevation margin of foothill woodland........ C. exilis 


Style equalling the anthers to well exserted; leaves usually gray-green; petals 
pink, sometimes with a darker blotch at the base of the blade, the claw often 
shorter but sometimes longer than the blade; or petals reduced, sepal-like, 
unexpanded and wrinkled, the claw very short, scarcely distinguishable 
from the blade which may be only 1-2 mm wide; arid inner coast ranges 
from eastern Alameda and western San Joaquin counties to western Kern 
and eastern San Luis Obispo counties; usually with Haplopappus lineari- 
folius, sometimes at the valley grassland margin and sometimes with Juni- 
perus californica or at the dry margin of foothill woodland....C. tembloriensis 


Clarkia springvillensis Vasek, sp. nov. Herba erecta, altitudine ad 
1 m; caulibus simplicibus vel ramosis, glabris et glaucis; foliis 2-9 cm 
longis, 5-20 mm latis; calycis tubo 3-4 mm; calycis limbo 12-16 mm 
longo, 3-4 mm lato, rubido, puberulento; petalis unguiculatis, 13-16 mm 
longis; petali unguiculo gracili, 7-9 mm longo; petali limbo 6-8 mm 
longo, 7-10 mm lato, roseo, in basi saepe macula roseo-purpurea; ovario 
10-17 mm longo; stylo 14-20 mm longo, quam staminibus longiore. 

An erect herb to 1 m tall; stems simple or usually branched, glabrous 
and glaucous; leaf blades 2-9 cm long, 5-20 mm broad; hypanthium 
3—4 mm; sepals 12-16 mm long, 3-4 mm wide, puberulent, usually dark 
red; petals 13-16 mm long, including a narrow red claw 7-9 mm long 
and a limb 6-8 mm long, 7-10 mm broad, lavendar-pink usually with a 
dark purplish spot at the base of the limb; stamens 8, filaments red; 
ovary 10--17 mm long; style 14-20 mm long; exceeding the stamens. 


The description is based on flowers from eight plants collected at the 
type locality in 1958, preserved in alcohol, and measured in the labora- 
tory. The gametic chromosome number is n= 9 (Vasek, 1960; 1964). 
The plants usually bloom in May. 

Type: Tulare Co.: Balch Park Road, 1.8 mi N of Springville Ranger Station, 
Vasek 630522-1A, May 22, 1963 (LA). Topotypes: Vasek 630522-1B (LA), Vasek 
630522-1C (UCR = University of California, Riverside), Vasek 630522-1D (DS), 
Vasek 630522-1E (DS), Vasek 630522-1F (RSA), Vasek 630522-1G (LA), Vasek 
630522-1H (UC), Vasek 630522-1I (RSA), Vasek 630522-1J (RSA), Vasek 630522- 
1K (UCR), Vasek 630522-1L (UC), Vasek 630509-1 (UCR). 


Specimens examined. Tulare Co.: above Springville, H. & M. Dearing 2666 
(SBBG) ; N of Springville, Purpus 1319 (UC). 


Clarkia tembloriensis Vasek, sp. nov. Herba erecta, altidudine ad 
8 dm; caulibus simplicibus vel ramosis, glabris et glaucis; foliis 2-7 cm 


1964] VASEK: CLARKIA 221 


longis, 3-13 mm latis, glaucis; hypanthio 2 (3) mm longo; calycis limbo 
9-16 mm longo, 2-3 mm lato, puberulento; petalis expansis, unguiculatis, 
13-17 mm longis, unguiculo gracili, 5-11 mm longo, limbo 5—9 mm longo 
et 4-9 mm lato; vel petalis non expansis, 7-14 mm longis, unguiculo et 
limbo (1) 2 mm lato: stylo 9-18 mm longo, quam staminibus longiore 
vel longitudine aequa. 

An erect herb to 8 dm tall; stems simple or sometimes branched, gla- 
bous, glaucous; leaf blades gray-green, 2—7 cm long, 3-13 mm wide; 
hypanthium 2(3) mm long; sepals 9-16 mm long, 2—3 mm wide, puberu- 
lent; petals expanded 13--17 mm long, including a narrow claw 5-11 mm 
long and a limb 5—9 mm long; 4-9 mm wide; or petals not expanded, 
7-14 mm long, (1) 2 mm wide, with the limb scarcely wider than the 
claw; style 9-18 mm long, exceeding, or usually not exceeding the 
stamens. 

The description is based on 39 plants from three sites in Carneros 
Canyon and nine plants from scattered localities as far north as Panoche 
road. Flowers and bracts of wild plants were preserved in alcohol, then 
dissected and measured in the laboratory. The gametic chromosome num- 
ber is n = 9 (Vasek, 1964). The plants bloom in April and May. 

Type: Kern Co.: Carneros Rock, Temblor Range, Lewis 1028, April 15, 1956 
(LA). 

Specimens examined. Alameda Co.: Corral Hollow, Eastwood & Howell 5291 
(POM). Fresno Co.: N of Coalinga, H. & M. Lewis 911 (LA, RSA); Alcalde, East- 
wood 13566 (CAS); 4.1 mi W of Coalinga, Vasek 620505-5 (UCR); 8.7 mi W of 
Coalinga, Vasek 62505-6 (UCR); Panoche Road, 9.7 mi W of P.G.& E. Panoche 
Substation, Vasek 620505-3 (UCR); Diablo Range, Cantua Creek Road, near State 
Highway 33, Vasek 620505-1 (UCR); Vasek 620505-2 (UCR). Kern Co.: Hodges 
Canyon, 1.7 mi S of United States Highway 466, Vasek 620425-3 (UCR); Cedar 
Canyon, Alec Cook Canyon, Twisselman 1189 (CAS). Kings Co.: Kettleman Hills 
near Avenal, Hoover 3301 (RSA, UC). San Benito Co.: 6.3 mi S of Panoche, 
McKaskill 440 (DAV); Idria Road, 4.6 mi S of Panoche Road, Hesse s.n. (UC); 
Call Mounties, 18 mi N of New Idria, Raven 10866 (RSA); 4 mi S of Paicines, 
Constance & Morrison 2267 (POM). San Joaquin Co.: Coral Hollow, Arnold s.n. 
(CAS) ; near Corral Hollow, Hoover 3358 (RSA). San Luis Obispo Co.: Red Hills, 
Kelly’s Canyon, Twisselman 2855 (CAS); 2 mi NE of Cholame, MacMillan 150 
(LA) ; Caliente Range, 8.1 mi S of Simmler, Vasek 620502-1 (UCR). Stanislaus Co.: 
Mount Hamilton Range, Arroyo del Puerto, Sharsmith 1753 (DS). 


University of California, Riverside 


LITERATURE CITED 


Lewis, H. and MArGareT E. Lewis. 1955. The genus Clarkia. Univ. Calif. Publ. 
Bot. 20:241-342. 


Lewis, H. and F. C. VAsEK. 1954. Clarkia exilis, a new Californian species. Ma- 
drono 12:211-213. 


VASEK, F. C. 1958. The relationship of Clarkia exilis to Clarkia unguiculata. Amer. 
Jour. Bot. 45:150-162. 


. 1960. A cytogenetic study of Clarkia exilis. Evolution 14:88-—97. 


. 1964. The evolution of Clarkia unguiculata derivatives adapted to rela- 
tively xeric environments. Evolution 18:26-42. 


202 MADRONO [Vol. 17 


TAXONOMIC NOTES ON THE CHRYSOTHAMNUS 
VISCIDIFLORUS COMPLEX (ASTEREAE, COMPOSITAE) 


LoRAN C. ANDERSON 


The genus Chérysothamnus, consisting of about 13 species, ranges over 
most of western North America. These shrubs, commonly called rabbit- 
brush, are related to section Macronema or section Ericameria of Haplo- 
peppus and more distantly to Solidago. For the past several years I have 
been studying the genus Chrysothamnus. New names and notes in the 
C. viscidiflorus complex are presented here to enable their use in future 
publications on the anatomy and cytology of the genus. In addition to 
my 550 collections of Chrvsothamnus and observations in the field and 
herbarium, I have utilized experimental gardens and greenhouse space 
at Logan, Utah (elevation 4850 ft) and Claremont, California (eleva- 
tion 1350 ft) in this study.! 

Selected features of the inflorescence were studied in at least three 
collections of each of the eight subspecies of C. viscidiflorus and in the 
related species discussed below. Averages for these features for some of 
the taxa grown in the gardens are presented in Table 1 to illustrate varia- 
tion produced in different environments. Variation in these features can 
also be seasonal; i.e., samples taken early in the flowering season fre- 
quently average more flowers per head than do samples from the same 
plant taken late in the season. Measurements from young flowers (pre- 
anthesis) could introduce error, since stigmatoid tissues mature later 
than style appendages. Heads of dried materials were selected for rela- 
tive maturity, then softened several days in 50% ethyl alcohol to restore 
original size and for handling ease. For features of the involucre, 10 
heads from each sample were studied; one flower from each head was 
used for floral measurements. 

In some instances involucral length, bract number, corolla length, and 
average flower number per head vary within a subspecies, but these 
features still have diagnostic value when used with other characteristics. 
The width-length ratio of the involucre and ratio of stigmatic appendage 
length to total style branch length appear to be reasonably constant 
(fig. 1). Blake (1940) questioned the value of stigmatic features as diag- 
nostic criteria; however, his position may have been influenced by the 
relationship he proposed for C. molestus. He described this taxon as a 
variety of C. viscidiflorus, but since C. molestus is not a close relative 
of C. viscidiflorus, the style branches of the two species need not nec- 
essarily be similar. 

Chrysothamnus molestus (Blake) L. C. Anderson, comb. nov. C. vis- 
cidiflorus var. molestus Blake, Jour. Wash. Acad. Sci. 30:368. 1940. 


1 Contribution no. 626, Department of Botany and Plant Pathology, Kansas Agr. 
Expt. Station, Manhattan, Botany serial no. 785. 


1964 | ANDERSON: CHRYSOTHAMUS 223 


Blake (1940) named this taxon as a variety of C. viscidiflorus (Hook. ) 
Nutt. of section Typici, but compared it with C. pulchellus (Gray) 
Greene, C. depressus Nutt., and C. vaseyi (Gray) Greene, all of section 
Pulchelli, the section to which C. molestus properly belongs. The glandu- 
lar achenes of this species are similar to those of C. depressus and C. 
vase\1. Heads of C. molestus are longer than those of C. viscidiflorus 
(table 1) and contain 21-24 phyllaries and 5—7 flowers; whereas heads 
of the latter usually have 10-19 phyllaries and 2—5 flowers. 


This species was previously known only from the San Francisco Mountains in 
Coconino Co., Arizona. The following collections extend the known range of this 
species in Coconino Co.: 16 air mi S of Grand Canyon Village, T28N, R2E, Ander- 
son 1839 (KSC) ; 1840 (KSC) ; National Tank, Haulapai Indian Reservation, Dar- 
row 3135 (ARIZ). 


CHRYSOTHAMNUS VISCIDIFLORUS (Hook.) Nutt. ssp. planifolius L. C. 
Anderson, ssp. nov. Frutex e radice crassa caudiceque multiramoso lig- 
noso. Caules plures, 1.2—2.5 dm alti, foliosi ad inflorescentiam, rigide 
erecti vel ascendentes, infra simplices, inflorescentiis ramosis, glabres. 
Folia (10)14—20 mm longa, 1—2 mm lata, anguste lanceolata, cuspidata, 
plana, aliquanto crassa, integria. Capitula in inflorescentiam corymbosam 
planam vel convexam disposita; involucris (4) 5—5.6 mm altis, cylindricis; 
phyllariis 15-17, stramineis, apicibus leviter crassis; floribus 3-4(5) ; 
corollis flavis, 3.5—4.5 mm longis, dentibus, 1.4—-1.7 mm longis; antheris 
1—1.5 mm longis, apicibus attenuatis 0.3—0.4 longis; stylus ramis 1.4—1.7 
mm longis, appendice lineis stigmaticis breviore; achaenis 3—3.5 mm 
longis, pubescentibus. 

Type: Arizona, Coconino Co.: dry sandy soil of sparse juniper slopes (Cedar 
Ridge) at 5500 ft, 16.5 mi N of The Gap, 11.5 mi S of Bitter Springs, Anderson 
1747 (ARIZ, KSC, MSC, UC—holotype, US, UTC). 

Additional specimens examined: Arizona, Coconino Co.: Box Canyon, Wupatki 
National Monument, Jones 28 in 1939 (US); 1 mi E of The Gap, Anderson 1851 
(KSC) ; 4.5 mi N of The Gap, Anderson 1852 (KSC) ; type locality, Anderson 1749 
(KSC); 11 mi N of Cedar Ridge Trading Post, Darrow 2886 (ARIZ); 6 mi S of 
Bitter Springs, Anderson 1853 (KSC); 1902 (KSC). Kearney & Peebles 12813 
(ARIZ, US), collected 20 mi W of Cameron, represents a geographically peripheral 
population with pubescent stems. 

This subspecies is related to C. viscidiflorus ssp. elegans (Greene) Hall 
& Clem., from which it differs by having flat, thickened leaves, glabrate 
stems, and smaller heads. It is perhaps distantly related to C. viscidiflorus 
ssp. stenophyvllus (Gray) Hall & Clem.; the two grow together at the 
type locality, but no intergradation has been observed. Chrysothamnus 
viscidiflorus ssp. stenophylius differs by having narrower, twisted leaves 
and larger heads. Twisted leaves appear to be only modifications to the 
xeric environment as indicated by greenhouse studies; however, all sub- 
species, except C. viscidiflorus ssp. planifolius, have twisted leaves to 
some degree in native habitats. 

CHRYSOTHAMNUS VISCIDIFLORUS ssp. HUMILIS (Greene) Hall & 
Clem. Keck (Munz, 1959) reduced this subspecies to synonymy under 


224 MADRONO [Vol. 17 


TABLE 1. DATA FOR SELECTED FEATURES OF INVOLUCRE AND FLOWER 


| 
| 


| 


; 3 
S 5 

S e ass 

> Ex | 

ps = a Ue a= FE) 

Ge Fe - 6) Se aul 

aS Shes oa © > = SH - Bel 

Collection oe 26 2s oe ae ae ie el 

Original locality R ge] © iS es =~ % SE] 

Taxon Garden progeny £ = Es ie Es Se So sient ay Sel 

C. molestus Anderson 1840 (KSC) 22,0. MUleL” 23:4 BeZ 81 12.7 3.2 37a 

Coconino Co., Ariz. | 

Claremont garden 2353 OID (25-4 AS ie Se 2.9 452) 

C. linifolius Anderson 1487 (KSC) 13.2 45, 4933 cw 49 26.1 1.7 404 

Grand Co., Utah | 

Logan garden 14.9 5.7 43.6 5.4 Sel eZ 99 125 417 

Anderson 1860 (KSC) 12.3 5.5. 46.3. 5.26.0 37.5 2A ATG 

Kane Co., Utah : 

Claremont garden lioes be2 642.3 be Sle 42-5 2.0 474 

C. viscidifiorus Anderson 1926 (KSC) 14.6 6.9 29.5 4.1 5.32 2925 ons 312 

ssp. elegans Juab Co., Utah 

Claremont garden 13.4 5.9 35.4 4.6 Age) 2753 22 33.0 

C. viscidiflorus Anderson 1617 (KSC) 14.8 Of 2222 2,8 doe LOR 3.1 53m 
ssp. humilis Modoc Co., Calif. 

Claremont in 1961 11.8 82-257 SZ 6.8 17.8 35 52.3 

Claremont in 1962 15 9.6 22.7 es (i Pe! 2.7 514i 

C. viscidiflorus Anderson 1747 (KSC) 15.2 525. 3.5 3.9 AS | 3158 1.5 203i 

ssp. planifolius | Coconino Co., Ariz. 

Logan garden 15.4 50. 35.1 AG. 3.8 (638.3 1.8 205m 

| 

C. viscidiflorus Anderson 885 (KSC) 14.9 50 ais S55) ASS 72587 235 2 

ssp. pumilus Ircn Co., Utah 

Logan garden 13% 6.2 28.0 3.9 5108-5129 Sol 118 

© vicidificrus Anderson 605° KSC) 152 65 254. 25) Sales soe 

ssp. stenophyllus Coconino Co., Ariz. | 

Claremont garden 15.0 6.1 275 5149) 5.0. 36:0 2.6 36.69 

Logan garden 122 6:5 © 25.8 3.6 Salt) 32 7 2.4 38.4 

C. viscidiflorus Anderson 1701(KSC) 163 74 29.7 4.5 62 283 2.9 204% 

ssp. viscidiflorus Cocenino Co., Ariz. | 

Claremont garden 14.4 fhe, ee!) 5.0 6.1 30.9 2.5 29.2% 

Logan garden 17.6 9.9 22.7 40 63 262 2.9 em 


C. viscidiflorus ssp. puberulus (D. C. Eat.) Hall & Clem. Intergrada- 
tion between the two taxa where their ranges overlap is indicative of their 
infraspecific relationship; nonetheless, farther north C. viscidiflorus ssp. 
humilis is one of the most distinctive units in C. viscidiflorus. Such dis- 
tinguishing features as the long, narrow involucre, few flowers per head, 
style branches included or barely surpassing the corolla lobes (fig. 2), 
and long stigmatic appendages (the five circles to the right in fig. 1 rep- 


1964] ANDERSON: CHRYSOTHAMUS 225 
50 A 
A 
45 A 
s 
a4 
40 = 
A 
mf & 
35 e@ o e 
ng @ eo — 
r) ee ®@ | 
30 ee ec ® . “8 
© @ 
ry & @ 
ee ® 
@ e @ 
r) eo 
20 25 30 35 40 45 50 55 60 65 70 


Fic. 1. Scatter diagram for selected floral features expressed in percentages. 
Horizontal axis is the ratio of style appendage to total style branch length; vertical 
axis is the width-length ratio of the involucre. Circles represent C. viscidiflorus ; 
triangles C. linzfolius; and squares C. spathulatus. 


resent this subspecies) were maintained when the plants were grown in 
experimental gardens (table 1). 


CHRYSOTHAMNUS AXILLARIS Keck. Populations from Inyo County, 
California, and adjacent Nevada for which this name has been pro- 
vided (Keck, 1958) do not appear to be sufficiently distinct from C. vis- 
cidiflorus to justify taxonomic recognition with the evidence at hand. 
These plants represent forms of C. viscidiflorus ssp. stenophyllus having 
involucral bracts with attenuate tips arranged in very pronounced ver- 
tical ranks. Anderson 1480 (KSC) and 1483 (KSC) from Emery and 
Grand Co., Utah, respectively, are collections of this subspecies with 
similar involucral features. 


The following are collections with similar bract alignment that are referable to 
other subspecies: to C. viscidiflorus ssp. vicidiflorus: Inyo Co., California, Ander- 
son 2000 (KSC) ; Anderson 2004 (KSC) ; 2007 (KSC) ; Coconino Co., Arizona, An- 
derson, 1881: to C. viscidiflorus ssp. elegans(?): Mono Co., California, Anderson 
2015 (KSC): to C. viscidiflorus ssp. puberutus: Inyo Co., California, Anderson 2014 
(KSC); Riverside Co., California, Anderson 1933 (KSC); 1935 (KSC); 1936 
(KSC), bracts also acuminate. 


The geographically isolated population from Riverside Co., California, 
is very different from other populations of C. viscidiflorus ssp. puberulus 
by having broader leaves, heads congested in the inflorescences, and 
17-21 phyllaries per head (an unusually high number for C. viscidi- 
florus). The disposition of this population cannot be determined until 
additional data are available. 


226 MADRONO [Vol. 17 


RAE ar 


Cott pod 


=H sone e 
See 
SSS ae ony erie 
bY Se = 


Se 
< -+ 


Fic. 2. Line drawings of flowers with stamens removed; a, C. viscidiflorus ssp. 
viscidiflorus (Anderson 1701, KSC); b, C. viscidiflorus ssp. humilis (Anderson 
1617, KSC); c, C. viscidiflorus ssp. planifolius (Anderson 1747, KSC); d, C. vis- 
cidiforus ssp. stenophyllus (Ferris 6924, POM, isotype of C. axillaris) ; e, C. lini- 
folius (Anderson 1860, KSC); d, C. spathulatus (Anderson 2052, KSC). 


CHRYSOTHAMNUS LINIFOLIUS Greene. Hall (Hall and Clements, 1923) 
treated this taxon as a subspecies of C. viscidiflorus; however, I recog- 
nize it as a separate species. In addition to differences in plant form and 
habital preference pointed out by Hall, C. lénifolius has these distin- 
guishing features: 1) heads are slightly turbinate rather than narrowly 
cylindric, are much wider in relation to their length (fig. 1), and have 
more flowers than C. viscidiflorus; 2) the point of staminal departure 
from the corolla tube is not readily apparent externally (fig. 2); whereas 
in C. viscidiflorus the tube is somewhat swollen at this point and often 
darkly colored; 3) the bases of the style branches are free; they are fre- 
quently fused for 0.5-1 mm in C. viscidiflorus; 4) the style appendages 
are longer than those of C. viscidiflorus, excepting the unique C. viscidi- 
florus ssp. humilis; 5) the plants have lateral roots from which adven- 
titious shoots arise as far as 4 dm from the main axis of the plant; in this 
respect C. linifolius differs from all other species in the genus. Even 
though plants of this species usually grow in sandy soil, they are soboli- 
ferous when grown in experimental plots of very fine clay loam. 


Chrysothamnus spathulatus L. C. Anderson, sp. nov. Frutex e radi- 
cibus multiramosis lignosis. Caules plures, 7-15 dm alti, foliosi ad in- 
florescentiam, puberulentes. Folia 3.5-4.7 cm longa, 5-10 mm lata, ad 
inflorescentiam parviora, spathulata, apicibus rotundatis apiculatisque, 
plana, subtile puberulenta. Capitula in inflorescentiam corymbosam plus 


1964 | GRIFFIN: DIGGER PINE ZL, 


minusve planam disposita; involucris 6.3—7.5 (8.5) mm altis, cylindricis; 
phyllariis 13-16, stramineis, apicibus pallide viridibus; floribus 3—4 (5) ; 
corollis flavis, 4.6-5.4 mm longis, dentibus, 1.7—2.6 (3.0) mm longis; 
antheris 2.1—2.3 mm longis, appendicibus 0.5 mm longis; stylus ramis 
2.2-3 mm longis, appendice lineis stigmaticis longiore; achaenis 2.5—3 
mm longis, sparse pubescentibus, pilis 0.1—0.25 mm longis, raro glabris. 
Cotyledones 7 mm longae, 3 mm latae, spathulatae. 

Type: New Mexico, Otero Co.: shaded loamy soil in Pinyon-Juniper-Oak Asso- 
ciation on the “Upper Burro Flats” at 6000 ft between LaLuz and LaBorcita can- 
yons, 7 mi NE by road from town of LaLuz, T15S, R10E, Sec 14 and 15, Anderson 
2052 (KSC, MSC, NMC, UC—holotype, US, UTC). The type collection was propa- 
gated at Claremont, California, as transplants taken from the type locality; Ander- 
son 1905 (KSC) represents immature specimens from the type locality. 

Additional specimens examined: New Mexico, Otero Co.: Sacremento Moun- 
tains, Rehder 331 (US); 332 (US); High Rolls, Vaughn 2155 (ARIZ); NW of 
High Rolls, Jackson 8083 (NMC). Socorro Co.: Mt. Oscuro at 6000-7000 ft, Dunn 
& Lint 4030 (NMC). 


This species, isolated by 150 mi from its near relatives, is distinct by 
having spatulate cotyledons (found in no other Chrysothamnus ), spatu- 
late to oblanceolate leaves, long style appendages, achenes with few, very 
short hairs, and unlike other members of the C. viscidiflorus alliance, the 
broken twigs emit a fragrance similar to that of C. nauseosus. The species 
can be further distinguished from C. viscidiflorus (of which ssp. lanceo- 
latus probably is the nearest relative) by its height, free style branches 
with long appendages, and lack of swelling at the point of staminal de- 
parture from the broader corolla tube (fig. 2). 


Department of Botany, Kansas State University, Manhattan 


LITERATURE CITED 


BLAKE, S. F. 1940. New species and new names among Arizona Asteraceae. Jour. 
Wash. Acad. Sci. 30:467-472. 

Harr, H. M. and F. E. CLrements. 1923. The phylogenetic method in taxonomy. 
The North American species of Artemisia, Chrysothamnus, and Atriplex. Car- 
negie Inst. Publ. 326:1-355. 

Keck, D. D. 1958. Taxonomic notes on the California flora. Aliso 4:101-114. 

Munz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley. 


DAVID DOUGLAS AND THE DIGGER PINE: 
SOME QUESTIONS 


JAMES R. GRIFFIN 


While collecting in the central coast region of California in 1831, 
David Douglas described Pinus sabiniana Dougl. in a letter written at 
Mission San Juan Bautista (Douglas, 1833). Descriptive passages in this 
letter—and sketches later made from the specimens—leave no doubt that 
Douglas had studied P. sabiniana cones, seed, and foliage at San Juan 


228 MADRONO [Vol. 17 


and sent them to England. He made no specific mention of where the 
material had been gathered, and his geographic notes are of little help 
in suggesting the locality. The collection may have come from the Gabi- 
lan hills southeast of San Juan, as Jepson (1910) speculated, or it might 
have come from the easily accessible Pine Canyon stand to the southwest 


(fie. 1), 


San owreia 
Bautista 


MM P coulteri 10 Miles 
P sabiniana 


Fic. 1. Present distribution of Pinus sabiniana and P. coulteri in the San Juan 
Bautista area of central California. (Adapted from unpublished vegetation type maps 
of the Pacific Southwest Forest and Range Experiment Station, U.S. Department 
of Agriculture, Berkeley, California.) 


1964] GRIFFIN: DIGGER PINE 229 


Other parts of the description, however, raise the question of whether 
Douglas had ever carefully observed P. sabiniana trees in the field or 
not. Did he possibly confuse this species with some other local pine? 
Paradoxically, he never mentioned the striking character of irregular 
crown branching so typical of the species. Instead he used such phrases 
as “The trees are of a tapering form, straight, and of regular growth, 40 
to 120 feet in height, 2 to 12 feet in circumference, clothed with branches 
to the ground, when standing far apart or solitary.” The dimensions 
quoted are reasonable, but “tapering”, “straight”, and regular growth” 
are terms with little relevance to this species. 

What other pine might have been involved in this ambiguous descrip- 
tion? Perhaps it was Pinus coulterti D. Don. During this period Douglas 
climbed the Gabilan range near San Juan to take geodetic observations. 
He may well have encountered P. coutleri on the higher ridges, for it can 
still be seen there from the vicinity of the mission. Stands of P. coulteri 
are more conspicuous and accessible to one climbing the hills near San 
Juan than those of P. sabiniana (fig. 1). Yet, Douglas made no mention 
of P. coultert in any of his writings which are still available to us. If he 
missed P. coulteri on the Gabilan range, he should have seen it later when 
he traveled through the Santa Lucia mountains to the south. In any case, 
he did send P. coulteri specimens and seed to England in 1832. This col- 
lection was apparently labeled as a variety of P. sabiniana (Little, 1948). 
When Loudon (1838) eventually looked at this material, he questioned 
the P. sabiniana label. After consulting Don he decided it was the same 
species that Don had received from Thomas Coulter and named P. 
coultert. 

If the P. sabiniana variety label on the P. coulteri collection accurately 
summarized Douglas’ views on the two pines, those who are acquainted 
with these relatively distinct pines are left with an unsatisfying feeling 
about the affair. Why did a botanist of such competence not describe 
P. sabiniana more clearly or why did he not discuss P. coulteri even if 
only as a variety within P. sabiniana? Only a few years before Douglas 
had repeatedly risked his life to track down a new pine. In this case he 
appeared to have lumped together two easily available species. Douglas 
did not seem to be inclined to create broad tree species, and with consid- 
erable perception he separated several new fir species differing only in 
rather subtle characters. I can only emphasize the difficulties here, for 
no answers are now available. But it would be interesting to know if 
Douglas combined some of his field observations of P. coulteri with some 
P. sabiniana specimens which had been given to him when he composed 
the San Juan letter. It would also be very interesting if we could go back 
in time to Monterey in 1832 and hear Douglas discuss these pines with 
his colleague Thomas Coulter. 


Pacific Southwest Forest and Range Experiment Station, Redding, California 


230 MADRONO [Vol. 17 


LITERATURE CITED 


Dovuc tas, D. 1833. Description of a new species of the genus Pinus. Trans. Linn. Soc. 
16:747-749. 

Jepson, W. L. 1910. The Silva of California. Mem. Univ. Calif. Vol. 2. Univ. Calif. 
Press, Berkeley. 

Littie, E. L. 1948. David Douglas’ new species of conifers. Phytologia 2:485-490. 

Loupon, J. C. 1838. Arboretum et fruticetum Brittannicum. Vol. 4. Longman, London. 


BARK PHOTOSYNTHESIS IN OCOTILLO 
H. A. Mooney AND B. R. STRAIN 


Ocotillo (Fouquieria splendens Engelm.) is a unique plant of the 
Sonoran Desert in respect to its physiology and candelabra growth form. 
The rapidity of leaf development following an increase in soil moisture 
after a drought period has been the object of numerous investigations. 
In only a few days after rain, leaves may fully develop on bare stems 
(Cannon, 1905). The problem of survival during extensive drought peri- 
ods has been studied also. As early as 1905, Cannon noted that, 
‘Although seemingly lifeless during the drought the plant is not dormant, 
since beneath its gray exterior there is a chlorophyllous bearing tissue 
which enables the photosynthetic process to go on, even if in a feeble 
manner...” Later, Scott (1932) described the anatomy of this bark 
chlorenchyma and noted its association with water storage cells and leaf 
primordia. 

The objective of this study was to determine if bark chlorophyllous 
tissue contributes to the photosynthetic economy of this plant. Bark 
photosynthesis during leafless periods could be of adaptive significance 
in respect to extended drought tolerance and might also be involved in 
the rapidity of ephemeral leaf production. 

Photosynthesis and respiration measurements were made in the field 
on portions of stems of two mature plants growing in Deep Canyon near 
Palm Desert, California.’ Measurements were made when the plants 
were in full leaf in March and when leafless subsequent to drought in 
May. A cylindrical double-walled plexiglass chamber was placed on the 
stem and sealed at both ends (fig. 1). Air temperature within the cham- 
ber was controlled by water flowing through the jacket from a constant 
temperature bath. The COz content of air passing through the chamber, 
and of free air, was determined with a Beckman model 15A infrared gas 
analyzer. Air flow rate was maintained at 120 liters per hour. 

When the plants were in leaf, measurements were made in the light 
and in a darkened chamber. Then, all leaves were removed and addi- 

1 We would like to express our appreciation to Lloyd P. Tevis for information 


relevant to ocotillo behavior in Deep Canyon as well as the personnel of the Philip 
L. Boyd Desert Research Center for assistance and the use of facilities. 


1964] MOONEY & STRAIN: OCOTILLO 23 


Fic. 1. Photosynthesis chamber on ocotillo stem. 


tional measurements were made on the leafless stem in light and dark. 
Measurements were performed also on the leafless stems subsequent to 
natural leaf loss. 


232 MADRONO [Vol. 17 


Additional measurements were made in the laboratory on a potted 
transplant which had been in a gallon container for over a year prior to 
use in the study. As in the field, measurements were made when the plant 
was in leaf; when the leaves had been hand-removed; and when leafless 
subsequent to drought. 

All measurements were performed at chamber temperatures between 
20-25°C and at light intensities parallel to the chamber between 4000 
to 6000 foot candles. 

The results of measurements on all plants are shown in Table 1. The 
figures represent apparently stable rates for the stated conditions and 
are expressed on a square decimeter stem surface basis. 


TABLE 1. GAS EXCHANGE MEASUREMENTS ON OCOTILLO 


With leaves 


Deep Canyon Plants Potted Plant 
no. 1 no. 2 
light —5.231 —6.45 —1.98 
dark +1.11 +0.86 +1.24 
Leaves stripped 
light +0.59 +0.53 +0.54 
dark -+1.04 +0.78 +0.90 
Bark photosynthesis —0.45 —0.25 —0.36 


1 Carbon dioxide production or absorption in mg/dm2 stem surface/hr. 


All plants had bark photosynthetic activity to varying degrees when 
in a hydrated leafy condition. However, in no instance was bark photo- 
synthesis great enough to attain the compensation point. That is, when 
hydrated leafless stems were illuminated, carbon dioxide was produced, 
but to a lesser extent than when the stem was darkened. Bark photo- 
synthesis then is expressed as the difference between carbon dioxide 
production of illuminated and darkened stems. Bark photosynthesis ot 
the three test plants varied between 0.25 and 0.45 mg CO2z/dm* stem 
surface/hr, a small fraction of the photosynthetic contribution of the 
leaves. 

There was no measurable bark photosynthesis on any of the three 
plants when they were in a drought-induced leafless condition. 

The chlorophyllous bark tissue of ocotillo was found to be photo- 
synthetically active when plants were in a non-drought condition. How- 
ever, even during such periods of photosynthetic activity, compensation 
was never attained under the test conditions. Bark photosynthesis, there- 
fore, probably contributes little to the overall seasonal photosynihetic 
economy of the plant. This small contribution, however, may be of some 
survival significance. 

Scott (1932) presumed that the rapid leaf development of ocotillo 
after rain depends upon efficient water conduction and high meristematic 


1964] REVIEWS 233 


activity, the latter implying a readily available respiratory substrate. 
Bark photosynthetic activity subsequent to precipitation may be of some 
importance in maintaining localized substrates utilized in leaf produc- 
tion. Knowledge of the precise adaptive significance of bark photosyn- 
thesis, if any, awaits more detailed studies. 


Department of Botany, University of California, Los Angeles 


LITERATURE CITED 


Cannon, W. A. 1905. On the transpiration of Fouquieria splendens. Bull. Torrey 
Club 32:397-414. 

Scott, F. M. 1932. Some features of the anatomy of Fouquieria splendens. Am. Jour. 
Bot. 19:673-678. 


REVIEWS 


The Quiet Crisis. By StTEWart L. UDALL. xii + 209 pp, 32 plates. Holt, Rinehart, 
and Winston, New York. 1963. $5.00. 


This should be an influential book in contemporary United States conservation 
literature. A wide audience is assured by the author’s position as Secretary of the 
Interior, the literary style, the brevity, the biographical approach, and the fresh 
insights into the history and future of the conservation movement in America. 

The contrast between the American Indian’s land ethic and that of the white 
settler is vividly protrayed. Gradually and sporadically a different set of values 
gained a toehold, with naturalists as the Bartrams and Audubon, the historian 
Parkman, and the philosophers, Emerson and Thoreau. George Perkins Marsh, 
and his international classic, Man and Nature, is given full credit as the intellectual 
“fountainhead of the conservation movement.” It is a rewarding feature of this book 
that these men who had little immediate practical influence on conservation are 
accorded nearly as much space as those later ‘men of action” who usually dominate 
United States conservation treatises. 

But “The Raid on Resources” had just begun, based on the “Myth of Super- 
abundance,” and expedited by the “Great Giveaway” of land as a federal policy. 
Men of action were needed, and their strengths and weaknesses are succinctly 
evaluated: Schurz and Powell, Pinchot and Muir, Mather and the two Roosevelts, 
Olmstead and Rockefeller. Curiously, Hugh Bennett and the Soil Conservation 
Service is accorded only eight lines, a neglect perhaps accounted for by the fact 
that the research staff of the Department of Interior, who are acknowledged by the 
author as collaborators, timidly avoided crossing departmental lines to cover 
Department of Agriculture affairs. 

Unusual emphasis for volumes of this sort is afforded the role of the private 
foundation, city planning, and the importance of wilderness. The approach in general 
is aesthetic and biographical rather than ecological and technical. The author’s 
acknowledgements to the works of Leopold, DeVoto, Krutch, Stegner, Atkinson, 
and Mumford, and his relatively full treatment of the naturalists and philosophers, 
clearly show a recognition of non-material as well as material benefits of conservation 
philosophy This emphasis serves to counterbalance that powerful segment of the 
present schizoid conservation movement which desires nothing more than a continu- 
ous sheet of highly productive crops, hybrid trees, or cows over a fully domesticated 
and manicured earth’s surface. The controversy, described by Udall, which developed 
between Pinchot and Muir on this issue continues unabated today, as shown by 
current debate on the Wilderness Bill in Congress. Possibly this point could have been 
given more than an aesthetic foundation alone by incorporating more ecology — 


234 MADRONO [Vol. 17 


for example, the value of relatively undomesticated ecosystems interspersed in space 
and time with the inevitable but productive monccultures, in order to provide 
greater homeostatic buffering in the form of high diversity against pest outbreaks 
and soil deterioration. The avoiding of oversimplification of food webs and inadequate 
circulation within vital biogeochemical cycles is a corollary of diversity which also 
can help weld aesthetic and practical considerations together. The space afforded 
the population problem is not in proportion to its significance. 

These criticisms do not detract from Udall’s accomplishment, which can perhaps 
best be summarized by his own words in the forward: “Each generation has its 
own rendezvous with the land, for despite our fee titles and claims of ownership, 
we are all brief tenants on this planet. By choice, or by default, we will carve out 
a land legacy for our heirs. We can misuse the land and diminish the usefulness of 
resources, or we can create a world in which physical affluence and affluence of the 
spirit go hand in hand.” — Joun PELTON, Butler University, Indianapolis, Indiana. 


Vege'ation and Flora of the Sonoran Desert. By ForEST SHREVE and Ira L. 
Wiccins. Vol. I, x + 1-840 pp., 37 plates, 27 maps; Vol. II, v + 841-1740 pp. 
Stanford University Press, Stanford, California. 1964. $22.50. 


The Sonoran Desert, one of the most arid areas of the North American continent, 
has received a botanical treatment in these two volumes that rivals the best account 
for any similar land surface. Nearly all of Part I of Volume I we have seen before 
as Carnegie Institution of Washington Publicaticn No. 591 but the bulk of the work 
as a whole is entirely new. The decision to reprint the contribution of the late 
Forest Shreve in the present work is a fortunate one, for it puts into the reader’s 
hands an out-of-print publication that deserves to be a part of the package, as it 
was originally intended by the authors. The fold-out maps of the Shreve contribution 
have been reduced to page size in the reprinted portion without loss of clarity. 
These maps, Map i and Map 2, appear on pages 6 and 7 and they are very effectively 
used on the back and front covers and adjoining pages of both volumes. Looking 
at Map 2, showing the routes taken during the many field trips of the authors 
throughout the Sonoran Desert, permits one to visualize the vitality of the work 
they have produced. But one has to remember that the map dates back to the late 
1940’s and that the many trips made by Wiggins since then are not shown. 

The work is vital because it represents and reflects the first hand, on the spot 
knowledge gained by numerous contacts with the living plants in their natural 
settings. This is not to imply that the deliberative, scholarly activities of the 
laboratory, herbarium and library have been neglected. By no means is this the case. 
The work on many of the richly represented genera of the flora are minor or major 
monographs in themselves. It is difficult, if not impossible, for the non-taxonomist 
to appreciate how much sustained effort goes into a publication of this nature. I am 
thinking particularly about the flora portion written by Wiggins. Here in the final 
product, we see what has become in floristic works generally, a shortened telegraphic 
style that follows a rigidly formal pattern. The information is packed into limited 
space with a uniformity that obscures the torment and labor required to produce it. 
Yet to do it differently, in a legitimately expanded and more informal way, would 
have required three fat volumes instead of two. 

The treatment of genera and species is moderately conservative and will have 
a lasting influence on subsequent works dealing with the general region because of 
the reliability achieved. As an example, one finds in the handling of the Cactaceae, 
where it would be easy to include many more genera by following recent reckless 
literature, a balanced position is taken with 23 genera recognized. In this family, 
Opuntia is one of the genera with a large number of species represented in the flora, 
having a total of 47 in all. Mammilaria, with its 39 species, further emphasizes the 
diversity at the specific level within the Cactaceae. 

The Sonoran Desert flora is a distinctive one, with many taxa of restricted 
distribution, many with unusual diversity, and others made up of plants with strange 


1964] NOTES AND NEWS 239 


growth forms. The number of genera and species of such major groups as ferns, 
gymnosperms and monocotyledons is low when compared to floras of other areas. 
The eighteen species of the genus Cheilanthes make it the largest of the fern genera 
present, and the peculiar genus Ephedra, with only six species, has two more than 
the total for Taxodium and Juniperus, the two other genera of gymnosperms repre- 
sented in the flora. There are 61 genera of grasses, about two thirds of the number 
in Munz’s A California Flora or in Gray’s Manual, and many of these are represented 
by but a single species. From this, it is obvious that the grass representation is not 
well developed. As expected, a genus like Agave is diversified, having 21 species, 
but on the whole the monocotyledons are not the prominent elments of the flora. 
The predominant role is played by dicotyledons, the major group that exceeds all 
others in abundance and diversification. Such genera as Dalea and Euphorbia match 
or exceed genera of the Cacti in numbers of species. I am always impressed by the 
heavy reprsentation of families such as the Malvaceae in the desert flora but the 
real shockers are in little known families such as the Fouquieriaceae with its bizarre 
Idria columnaris or the peculiar Pachycormus discolor of the Anacardiaceae. One 
expects the many shrubs and small trees of the Leguminosae, but personally I was 
much impressed with the woody members of families such as the Euphorbiaceae and 
Capparidaceae, whose temperate members tend to be herbaceous. We tend to think 
of such giant Cacti as Carnegiea and Pachycereus, and the tree Yuccas in connection 
with the Sonoran Desert but they are only a small part of the great complex of 
annuals, perennials, succulents, small trees and above all shrubs of many kinds and 
descriptions that make up the whole. 

It is true, the desert has a fascination that is hardly describable and I can 
understand the repeated beckonings that have pulled the authors of these fine 
volumes again and again within its borders. Even one trip of six weeks duratjon into 
Sonora with Professor Wiggins left an indelible impression on me. I often recall the 
vivid sunsets, the long, quiet dusk and the grotesque shapes of plants and horizons 
as the night came on, the tin cups set on the roof of the Model A station wagon 
to cool their liquid contents, or the treachery of an otherwise innocent playa after 
a sudden downpour of rain. The books under review do not tell of these things but 
they do contain the content that makes the desert appreciable when their storehouse 
is appropriately tapped. 

The volumes are well designed, printed and manufactured. The Stanford 
University Press is to be congratulated for their interest and leadership in producing 
these and other finely executed botanical volumes. It is trite to say that these excel- 
lent books are a must for everyone interested in the Sonoran Desert. They are that 
and more. They contain information not easily obtained elsewhere and provide a 
standard of comparison for work on other deserts of the world. They bring together 
for the first time and make comprehensible the ingredients of knowledge necessary 
for an understanding of many aspects of the plant cover of a large and very 
interesting area. The groundwork has been carefully laid bare by the authors, who 
have prepared the way for others to continue the process of building our fund of 
knowledge of this great desert. — REED C. Rotiins, Gray Herbarium of Harvard 
University, Cambridge, Mass. 


NOTES AND NEWS 


EpwIN BINGHAM CoPpELAND.— Dr. Copeland died in Chico, California, on 
March 16, 1964 at the age of 91. He was well known to members of the California 
Botanical Society and served as its president in 1944. Dr. Copeland received his 
A.B. from Stanford University in 1895 and his Ph.D. from the University of Halle 
the following year. Prior to going to the Philippines in 1903, he taught briefly at 


236 MADRONO [Vol. 17 


Chicago, Wisconsin, Stanford, California State Normal School at Chico, Indiana, 
and West Virginia. During his stay in the Philippines he served as dean and professor 
at the Philippines College of Agriculture. He was manager of the herbarium at the 
University of California, Berkeley, from 1928 to 1932 and in 1935 he was appointed 
Research Associate in Botany. In agriculture circles, Dr. Copeland is remembered 
as an expert in tropical agriculture, particularly with respect to the growing of rice. 
Among taxonomic botanists he is remembered as a great pteridologist and his Genera 
Filicum is present in the libraries of most of them. 


NOTES ON THE F Lora OF Arizona. IIJ.—Since the publication of the Howell and 
McClintock supplement to Kearney and Peebles’ Arizona Flora in 1960, some inter- 
esting additional plant species and range extensions have come to my attention. This 
paper is journal article no. 838 of the Agricultural Experiment Station, University of 
Arizona. 

Nuphar polysepalum Engelm. (Nymphaea polysepala Greene) was collected in 
Woods Canyon in the stream below the dam of Woods Canyon Lake, Sitgreaves 
National Forest, Coconino County (Mason, Phillips, & Niles 2273, ARIZ). The speci- 
mens represent the first and so far only record of the Nymphaeaceae in Arizona. Our 
attention was directed to the area by an inquiry from Margaret Schmidt. 


Hypericum anagalloides C. & S. (Mason, Phillips, & Niles 2270, ARIZ) and Viola 
palustris L. ssp. brevipes Baker (Mason, Phillips, & Niles 2269, ARIZ) were collected 
along the stream also in Woods Canyon. This small prostrate Hypericum is abundant 
along the stream. It is a northern species ranging along the Pacific Coast to British 
Columbia and eastward to Montana. This collection is the second record for Arizona; 
the previous report is a series of collections by L. N. Goodding from V. T. Park on 
the North Rim of the Grand Canyon, about 250-300 airline mi to the northwest. 
Russell reported the first collection of Viola palustris from Arizona (Rhodora 65:49. 
1963). The collection from Woods Canyon represents a second locality for this 
species. Not much extension of range is involved, however, for the two areas are 
only about three miles apart and are within the same drainage system. 


Asclepias cryptoceras Wats. was collected along the road from Kaibito to Inscrip- 
tion House, 15 miles from Inscription House at the base of White Butte (Mesa), 
Coconino County (Mason & Phillips 1942, ARIZ). The previous single record of this 
species in Arizona was from Pipe Springs, Mohave County. The new collection repre- 
sents an eastward extension of the range of about 100 mi. 


Arnica foliosa Nutt. (A. chamissonis Less. ssp. foliosa (Nutt.) Maguire) was 
collected at Crescent Lake, White Mountains, Apache County (Haskell s.n., July 13, 
1958, ARIZ). The previous reports of this species in Arizona were from the Kaibab 
Plateau and the North Rim of the Grand Canyon. The new location represents 
a range extension of about 250-300 airline mi eastward; however, a collection is 
known from Washington Pass, Chuska Mountains, San Juan County, New Mexico 
(McKnight 58080209, ARIZ), a distance of about 150 mi northeast of Crescent 
Lake——Cnwar_es T. Mason, University of Arizona Herbarium, Tucson. 


| 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

Membership in the California Botanical Society is normally considered 


a requisite for publication in MaApRONo. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume [, 1916-1929 : 2 ame. 4-95-00 
Volume IT, 1930-1934. . . . . 5.00 
Volume III, 1935-1936 ee i re S00) 
Volume IV, 1937-1938. . . . =. 5.00 
Volume V, 1939-1940 . . . . . ~ 5.00 
Volume VI, 1941-1942 ai. he eee OF OO) 
Volume VII, 1943-1944 oa ee, a EE OO 
Volume VIII, 1945-1946 . . . . ~ 5.00 
Volume IX, 1947-1948 oI an tee OL OO 
Volume X, 1949-1950 . . . . =. ~~ 7.00 
Volume XI, 1951-1952 o, eer ees O0 
Volume XII, 1953-1954 . . . . 8.00 
Volume XITI, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 er aie MOOS) 
Volume XVI, 1961-1962 . . . = . 12.00 


Single numbers: 


Volumes I and Il 2) 2) 5.) =) a ee 50 
Volumes II through XIV. . .. 1.00 
Volume acy <3 29 gee eee LS) 
Volume XVI... °.2 <3), eee 1.50 


The subscription price of MApDRONo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


ADRONO 


VOLUME 17, NUMBER 8 OCTOBER, 1964 
Contents 
PAGE 
THE GENUS XEROCOMUS QUETLET IN NORTHERN CALI- 
FORNIA, Harry D. Thiers 237 


SURVIVAL OF TRANSPLANTED CUPRESSUS AND PINUS 
AFTER THIRTEEN YEARS IN MENDOCINO COUNTY, 


CALIFORNIA, Calvin McMillan 250 
A PECULIAR CASE OF HEMLOCK MISTLETOE PARASITIC 
oN Larcu, Job Kutjt | 254 
LYONOTHAMNOXYLON FROM THE LOWER PLIOCENE OF 
WESTERN NevapA, Virginia M. Page Zo, 
| DoCUMENTED CHROMOSOME NUMBERS OF PLANTS 266 


THE HoRDEUM JUBATUM — CAESPITOSUM — BRACHY- 
ANTHERUM COMPLEX IN ALASKA, W. W. Mitchell 


| and A. C. Wilton 269 
| THE GENUS ESCHSCHOLZIA IN THE SOUTH COAST 

: RANGES OF CALIFORNIA, Wallace R. Ernst 281 
: 


NOTES AND News: A NOTE ON THE TYPE LOCALITY OF 
TETRACOCCUS ILICIFOLIUS, H. Thomas Harvey 268 
CONDALIA MEXICANA SCHLECHT. VAR. PETALIFERA 
M. C. JOHNSTON, VAR. NOV., Marshall C. Johnston 280 
ECHINOCHLOA ORYZICOLA IN CALIFORNIA, Beecher 
Crampton; NEW COMBINATIONS IN WESTERN 
NortH AMERICAN VIOLETS, J. Clausen 294 


A WEST AMERICAN JOURNAL OF BOTANY 


INDEX 296 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 

1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price 

$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


EDGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 
HERBERT F. COPELAND, Sacramento College, Sacramento, California 
Joun F. Davinson, University of Nebraska, Lincoln 
WALLACE R. ERnsT, Smithsonian Institution, Washington, D.C. 
MitpreD E. Matuias, University of California, Los Angeles 
ROBERT ORNDUFF, University of California, Berkeley 
Marion OwnBEY, Washington State University, Pullman 
REED C. RoLiins, Gray Harbarium, Harvard University 
Ira L. Wiccins, Stanford University, Stanford, California 


Editor—JoHNn H. THomas 
Dudley Herbarium, Stanford University, Stanford, California 


Business Manager and Treasurer—Douglas M. Post 
Biology Department, San Francisco State College 
1600 Holloway Avenue, San Francisco 27, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: G. Ledyard Stebbins, Department of Genetics, University of California, 
Davis. First Vice-President: Annetta Carter, Department of Botany, University of 
California, Berkeley. Second Vice-President: Carl W. Sharsmith, Department of 
Biological Sciences, San Jose State College. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding 
Secretary, Margaret Bergseng, Department of Botany, University of California, 
Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State 
College. 


THE GENUS XEROCOMUS QUELET IN 
NORTHERN CALIFORNIA 


Harry D. THIERS 


During the past four years considerable attention has been given to the 
mushroom flora of the coastal redwood region of central and northern 
California. The areas studied most intensively include Jackson State 
Forest in Mendocino County and forested portions of Marin and Santa 
Cruz counties. The bolete flora of these areas has received special em- 
phasis. As the study progressed it became apparent that there were sev- 
eral groups of closely related species in which determinations were con- 
sistently difficult. One such group was composed of the bolete species 
belonging to the genus Xerocomus Quelet, and this paper presents the 
results of an attempt to more clearly delineate the California species 
belonging to that genus. 

Special gratitude is extended to J. E. Sindel, Forest Manager, Jackson 
State Forest, Forestry Division of California for his wholehearted co- 
operation and generosity in placing facilities of the state forest at our 
disposal during the collecting periods in the fall of the year. 

Macroscopically the chief distinguishing characteristics of this genus 
are the tomentose to velutinous to less commonly pruinose surface of the 
pileus which is typically dry except for Xerocomus badius (Fries) Kuhn. 
ex Gilb., common in the midwest and northeast, which is viscid. Also, the 
pileus surface frequently becomes conspicuously rimose to areolate, es- 
pecially with age. The flesh of the pileus is usually yellow or pallid and 
changes to blue upon exposure in several species. The taste of the flesh 
is typically mild or slightly acid. The hymenophore or tubes do not 
readily separate from one another and are some shade of yellow. They 
are typically depressed to arcuate around the stipe and frequently have 
large, compound pores which are concolorous with the body of the tube. 
The hymenophore, like the flesh, often changes to blue when bruised. 
The stipe is typically equal or subequal to clavate, never appearing truly 
bulbous, and the surface is typically dry and glabrous to occasionally 
reticulate. Although the surface may appear granulose, the granules are 
not similar to those on the stipe of Swzl/us S. F. Gray species. 

The microscopic features of these fungi include smooth or perhaps 
obscurely roughened to striate spores that are typically subfusoid to sub- 
cylindric in outline. The cystidia are often abundant but typically incon- 
spicuous because of the very thin wall and hyaline contents. At times, 
however, they may become incrusted, particularly around the pores, and 
stain ochraceous to brown when mounted in a solution of potassium 
hydroxide. According to Singer (1945b) the most distinctive microscopic 
feature is the obscurely divergent to subparallel tube trama which is in 
contrast to the noticeably divergent trama in other genera. As has been 


Maprono, Vol. 17, No. 8, pp. 237-300. October 29, 1964. 


238 MADRONO PVolei7 


Fics. 1-2. Cuticle of Xerocomus: 1. X. chrysenteron, X 500; 2, X. zelleri, X 500. 


1964] THIERS: XEROCOMUS 239 


pointed out by Snell and Dick (1958), however, there are all degrees of 
intergradation of this character and unless very young specimens are 
available it is of little value. The cuticle is usually highly differentiated 
and varies from a layer of interwoven, heavily incrusted hyphae (fig. 1) 
to more or less radially arranged hyphal tips to a distinct palisade of 
globose to vesiculose cells (fig. 2). The surface of the stipe is usually inter- 
woven but may possess caulocystidia and, in the apical portion, be fer- 
tile. Clamp connections have not been found. Results of some of the 
routine chemical reactions do not appear to be highly distinctive. 

Although numerous characters have been cited in the above discus- 
sion, the fact remains that generic distinction is not based upon the pres- 
ence of one or two major characters but rather by a combination of 
several characters. Perhaps the most important single character, at least 
for field recognition of the genus, is the nature of the tubes. Snell and 
Dick (1958) have pointed out that the tubes of Xerocomus species form 
a “faveolate” pattern and do not readily separate from one another. They 
compared their arrangement to that of a honeycomb, which is quite dif- 
ferent from the “tubulate” arrangement in other genera in which the 
tubes are easily separated from one another and which, according to 
them, ‘‘resemble a bundle of slightly cooked spaghetti.” This character 
has been found constant for the California species except for X. zelleri 
which often seems somewhat intermediate. They also mentioned that 
Xerocomus badius was an exception to this pattern. Species belonging to 
this genus are most frequently confused with those belonging to the genus 
Boletus Fries, and it is the opinion of many that they are not sufficiently 
distinct to be recognized as belonging to separate genera. 

An analysis of the presently known species of Xerocomus indicates 
that they quite probably represent an intergrading series between the 
genus Boletus and the genus Boletellus Murr. The characters in common 
with Boletus are obvious and include the general aspect of the fruiting 
bodies, color of the hymenophore, the absence of lacunose or shaggy re- 
ticulations on the stipe as well as the smooth spores which are not exces- 
sively long (i.e., less than 20 y.). On the other hand the group has an 
affinity with some of the rough spored species although it may be some- 
what less apparent than with Boletus. Snell, Singer and Dick (1959), for 
example, have reported that carpophores of X. zelleri examined by them 
had obscurely roughened spores. Although such a spore character has 
not been observed in carpophores collected in California by this author, 
such a spore would clearly indicate a close relationship with Boletellus. 
As a matter of fact they recently transferred X. zelleri to Boletellus 
because of the spores and a somewhat intermediate type of tube trama. 
Further proof of the close relationship of these two genera is seen in 
Boletellus mirabilis (Murr.) Singer. This species has a dry, tomentose 
pileus and smooth, although large, spores. These characters caused Singer 
(1940) to place this species in Xerocomus; however, because of the 
large spores and general aspect of the carpophores he (1945a) later trans- 


240 MADRONO [Vol. 17 


Fics. 3-4. Basidiospores of Xerocomus: 
senteron, x 1750: 


3, Xa truncatus; 51 /50554, Se Xechin 


1964] THIERS: XEROCOMUS 241 


ferred it to Boletellus which seems to be a more satisfactory disposition 
of the species. 

The number of species of Xerocomus occurring in the United States, 
according to Singer (1962), is twelve. The number which has been found 
in northern California is five, representing a relatively high percentage 
of the total number. The species collected here fall into two distinct 
groups. The first group is represented by X. subtomentosus and X. illu- 
dens and has yellow to ochraceous to olive brown pilei, large, compound 
pores and more or less yellow to pallid stipes. The other group is com- 
posed of X. chrysenteron, X. truncatus and X. zelleri and is characterized 
by very darkly pigmented (dark brown to almost black) pilei, relatively 
small pores that change to blue when bruised and red to reddish stipes. 

All of the species included in this paper have been collected in the rel- 
atively dense mixed forests of the coastal range although X. subtomen- 
tosus is often found in more open areas, particularly along road cuts. 
Four of the five species have been found to be relatively abundant but 
only one collection of X. dludens has been made. Xerocomus subtomen- 
tosus and X. gelleri are the most abundant species and the latter fruits 
for a considerably longer period than any of the other species. One addi- 
tional species, X. /enticolor, was described by Dick and Snell (1960) from 
collections made in the Mount Shasta area by W. B. Cooke. An examina- 
tion of the type has shown this species to belong to the genus Suillus, 
thus it is excluded from this paper. Also, Boletellus mirabilis has been 
collected but, as mentioned earlier, since it is no longer considered a 
member of Xerocomus it is likewise excluded. 

The collections are in the herbarium of San Francisco State College, 
San Francisco, California, and the author’s accession number is included 
for each collection studied. The colors appearing in quotations in the 
species descriptions are those of Ridgway (1912). 


KEY TO SPECIES 


Pileus yellow to ochraceous to occasionally umber; if darker (i.e. bister to sepia) 
then stipe coarsely reticulate. 
Stipe coarsely reticulate from %3 to entire length... 1. X. illudens 
Stipe smooth to granulose, reticulate only at apex if at all........... 2. X subtomentosus 
Pileus dark brown to black, usually with reddish, granulose but not reticulate stipe. 
Spores rounded or subacute at apex. 

Pileus tomentose to velutinous, dark brown to chestnut colored, frequently 
areolate with age; cuticle composed of interwoven to radially arranged 
hey go lea Nala Seen eer coke ees e ee cbr Nee tere eed 4. X. chrysenteron 

Pileus conspicuously pruinose at least when youpng; almost black in color; 
surface strongly pitted but usually not rimose; cuticle differentiated as a 
compact palisade of inflated hyphal tips...............2..220..22:::22:0--00--- 5. X. zelleri 

SOLES suuunCa Ler dita DeKese.ertte ee eee ee ee ee ree ee 3. X. truncatus 


1. XEROCOMUS ILLUDENS (Peck) Singer, Farlowia 2:293. 1945. Boletus 
illudens Peck, Rep. N.Y. St. Mus. 50:108. 1897. Ceriomyces iludens 
(Peck) Murr., N. Am. FI. 9:145. 1910. 


242 MADRONO [Vol. 17 


Pileus 4-7 cm broad at maturity; convex when young becoming 
plano-convex to broadly convex with age; surface dry, conspicuously 
tomentose to velutinous, but not with fibrillose fascicles or scales, often 
somewhat areolate near the margin; evenly colored near ‘“‘Saccardo’s 
umber” to “‘snuff brown” to occasionally as dark as “‘bister’’ to “‘sepia’’; 
margin incurved, entire. Flesh 1-1.5 cm thick, white, unchanging when 
bruised or exposed; taste mild, odor not distinctive. Tubes narrowly to 
broadly and deeply depressed around the stipe; colored ‘‘chalcedony yel- 
low” to “bright chalcedony yellow” during all stages of development, 
unchanging upon exposure, 1-1.5 cm long; pores very large, 1-3 mm, 
concolorous with tube, unchanging upon bruising. Stipe 6—7 cm long, 
1-2 cm broad at the apex; yellow at the apex, becoming “warm buff”’ to 
“light buff” toward the base, staining reddish brown when bruised; sur- 
face with broad, coarse reticulations extending the full length of the stipe 
or at least the upper two-thirds of the length, moist to dry; typically 
crooked; tapering to a point at the base; whitish to pallid mycelium at 
the base; solid; flesh white, unchanging upon exposure. Spores 11.2—13.5 
< 4.0-6.4 », pale yellow to pale ochraceous in KOH, ochraceous in Mel- 
zer’s reagent, subfusoid to subellipsoid, not truncate, walls smooth and 
moderately thickened; basidia 30-36 & 9-12 u, hyaline, 4-spored, rarely 
2-spored; cystidia 48-60 *« 6-10 vp, scattered to abundant, thin-walled, 
hyaline, cylindric to subfusoid to occasionally subcapitate or obscurely 
fusoid-ventricose; tube trama hyaline, obscurely divergent to subparallel 
to somewhat interwoven, hyphae up to 6 y. in diam; pileus trama loosely 
interwoven, hyaline, homogeneous, occasional hyphae up to 12 y. in diam; 
hypodermis not strongly differentiated, more compactly interwoven than 
pileus trama, hyphae up to 8 yp in diam; cuticle pale yellow in KOH, 
composed of a tangled, more or less interwoven mass of free hyphal tips 
which are septate but not distinctively enlarged or differentiated, con- 
tents appearing dull yellow in KOH, up to 7 y in diam, but narrowed 
somewhat at the septa; cuticle of stipe typically composed of irregular 
tangled masses of hyphal tips, the terminal cell of which is typically en- 
larged and more or less clavate in shape, up to 12 yw broad and 30 yw in 
length; no caulocystidia or caulobasidia seen; clamp connections absent. 

Chemical Reactions. KOH—flesh negative to pale yellow; cuticle dark- 
ening slightly; NH,OH—flesh negative; cuticle negative to darkening 
slightly; HCl—flesh negative; cuticle pink; HNOs—flesh negative to 
pale yellow; cuticle dark red to pink; Sulfoformalin—flesh negative; 
cuticle pink; FeSO,—flesh negative or becoming pale gray; cuticle 
negative. 

Material Studied. Santa Cruz Co.: Thiers 10796. 

Discussion. Only one collection of this species has been made and 
this in the open Ponderosa pine forest near Ben Lomond rather than in 
the Mendocino region where the other species have been found in relative 
abundance. Although only a single collection is available for examina- 
tion, there is little doubt of its identity because of the characteristic 


1964] THIERS: XEROCOMUS 243 


large, coarse network on the stipe. As has been pointed out by Coker 
(1943) this species is similar in appearance to X. subtomentosus with 
the most consistent difference being the reticulate stipe. In addition the 
flesh of X. subtomentosus frequently changes to blue when bruised or 
exposed while no change has been observed in X. illudens. The micro- 
scopic characters of the two species appear very similar. 

When compared with collections from the southeastern United States 
the California material is similar except that the carpophores are more 
heavily pigmented and are more olive or umber in appearance. The same 
difference has been noticed between eastern and western carpophores of 
X. subtomentosus. 

As far as can be determined this is the first report of this species west 
of the Rocky Mountains. 

2. XEROCOMUS SUBTOMENTOSUS (Fries) Quelet, Fl. Mycol. Fr. 418. 
1888. Boletus subtomentosus Fries, Systema Mycologicum 1:389. 1821. 
Ceriomyces subtomentosus (Fr.) Murr., Mycologia 1:153. 1909. Cerio- 
myces oregonensis Murr., Mycologia 4:97. 1912. 

Pileus 6-15 cm broad when fully expanded, convex when young, be- 
coming broadly convex to plano-convex to plane to shallowly depressed 
at maturity; surface dry, dull, even, conspicuously velutinous to sub- 
tomentose to matted tomentose during all stages of development; when 
young colored “Isabella color” to “tawny olive” to ‘‘clay color” to occa- 
sionally near “‘yellow ocher” to “antimony yellow” usually becoming 
“sayal brown” to “snuff brown” to “buckthorn brown” to “olive ocher”’ 
with age; no reddish pigmentation along the margin or in cracks in the 
flesh, usually not conspicuously areolate; margin entire, incurved to de- 
curved, sometimes staining dark brown when handled. Flesh 1—2.5 cm 
thick, firm, whitish to pale yellow, often concolorous with the cuticle near 
the surface, often changing to blue when exposed; taste mild to slightly 
acid; odor not distinctive. Tubes adnexed to arcuate-decurrent when 
young, typically becoming shallowly to deeply depressed with age, up to 
1 cm long; when young colored “‘antimony yellow” to “‘warm buff” chang- 
ing to “primuline yellow” to “amber yellow” to “honey yellow” to “olive 
ocher”’ with age, unchanging or sometimes becoming blue when bruised; 
pores large, 1-2 mm, angular to highly irregular in outline, often com- 
pound, concolorous with tube. Stipe 5—11 cm long, 1—2.5 cm broad at the 
apex, equal or sometimes narrowing at the base; surface dry, glabrous to 
granulose to appressed fibrillose or longitudinally ridged, sometimes ob- 
scurely reticulate at the apex; colored ‘“‘Naples yellow” at the apex be- 
coming “tawny” to ‘ochraceous tawny” in the mid-portion and “‘warm 
buff” at the base, occasionally the background color is yellow but heavily 
overlain with darker colored granulations; often with whitish to yellow 
mycelium at the base and yellow rhizoids arising from the base; solid; 
flesh whitish to pale yellow, typically changing to blue when exposed. 
Spores pale yellow-ochraceous in KOH, smooth, thin-walled, subfusoid 
to subcylindric, 11.5-16 & 3.5-5 w; basidia clavate, hyaline in KOH, 


244 MADRONO [Vol. 17 


4-spored, 27-32 * 7-9 uw cystidia scattered to numerous, hyaline, thin 
walled, fusoid to fusoid-ventricose, often with elongated, tapering apices, 
40-66 8-12 »; tube trama obscurely divergent to subparallel, with 
oleiferous hyphae interspered, hyphae + 5 » in diam; pileus trama in- 
terwoven, homogeneous except for interspersed oleiferous hyphae; cuticle 
pale ochraceous yellow in Melzer’s reagent and KOH, loosely interwoven 
with numerous free, septate hyphal tips, often strongly incrusted, hyphae 
5—7 yu. in diam; surface of stipe differentiated as a layer of free to loosely 
interwoven hyphal tips with occasional tufts or clusters of caulocystidia 
which are hyaline, fusoid to clavate, thin walled, 24-40 « 6-9 wu; clamp 
connections not present. 

Chemical Reactions. KOH—flesh negative to pale yellow; cuticle 
darkening slightly; NH,OH—flesh negative; cuticle negative to dark- 
ening slightly; HCl—flesh negative; cuticle pink; HNOs;—flesh nega- 
tive to pale yellow; cuticle dark red to pink; Sulfoformalin—flesh nega- 
tive; cuticle pink; FeSO,—flesh negative or pale gray; cuticle negative. 

Material Studied. Marin Co.: Thiers 10857, 10872, 11145. Mendocino Co.: 
Thiers 8180, 8183, 8396, 8405, 8817, 8818, 8876, 8877, 9061, 9262, 9292, 9309, 9337, 
9348, 9438, 9471, 9767, 10573, 10647, 10676, 11019. Santa Cruz Co.: Thiers 8694, 
10756, 10771, 10964. San Mateo Co.: Thiers 8670, 10926, 11198. 

Discussion. This species is common under conifers and hardwoods in 
the coastal forests and also fruits along road cuts and similarly disturbed 
areas. The yellow to olive pileus, large, compound pores and more or less 
yellow stipe distinguish it from members of the complex composed of 
X. chrysenteron, X. truncatus and X. zelleri. As was indicated for X. 
illudens it is most likely to be confused with that species except for the 
distinctly reticulate stipe. The stipe of X. subtomentosus may be typically 
longitudinally ridged or granulose but not reticulate except occasionally 
at the very apex. When compared with midwestern collections the Cali- 
fornia carpophores appear more heavily pigmented, particularly with 
olive tones. Also the western carpophores often appear considerably 
darker when dried than those from other parts of the United States. 

Early in this century Murrill (1912) described Ceriomyces oregonensis 
from a collection made in the vicinity of Newport, Oregon. An examina- 
tion of the type revealed no significant differences between it and X. 
subtomentosus, thus it is reduced to synonymy with that species. 


3. XEROCOMUS TRUNCATUS Singer, Snell, & Dick ex Snell, Singer, & 
Dick, Mycologia 51:573. 1959. 

Pileus 7.5-12 cm broad when mature; convex when young becoming 
plano-convex to plane at maturity; surface dry, conspicuously tomentose 
during all stages of development, relatively smooth and even to some- 
times conspicuously areolate; colored near ‘“‘tawny olive” to “buffy 
brown” when young darkening to “clove brown” to “olive brown” to 
“sepia” to “bister’ when older, exposed flesh in cracks usually showing 
reddish tints; margin entire, incurved. Flesh 1—1.5 cm thick, colored 
“warm buff” to “light buff,” unchanging or becoming blue in irregular 


1964] THIERS: XEROCOMUS 245 


areas; taste often acid and somewhat unpleasant; odor not distinctive. 
Tubes subdecurrent to deeply and often narrowly depressed around the 
stipe; colored “olive ocher” to ‘Isabella color” to “old gold,” bluing 
readily when bruised; pores angular, often more than 1 mm. broad, con- 
colorous with tubes, Stipe 8-12 cm long, 1.5—3 cm broad at the apex; 
equal to tapering slightly toward the base; colored “hydrangea red”’ to 
“dark vinaceous” at the apex, becoming colored near “‘warm buff’’ to 
“light buff” to “naphthalene yellow” toward the base, usually with lines 
or ridges colored near “eugenia red”’; bright yellow mycelium at the base; 
surface dry, glabrous to obscurely punctate or ridged; solid, flesh red- 
dish toward the base, pallid at the apex. Spores 12.5-15 & 4.5—-6.0 yu, 
tawny to cinnamon in KOH and Melzer’s reagent, ellipsoid to subventri- 
cose, conspicuously truncate, walls smooth, relatively thick; basidia 28— 
33 X 7-10 yp, hyaline, clavate, 4-spored; cystidia scattered to numerous, 
often staining tawny to ochraceous in KOH, incrusted or filled with 
amorphous content, clavate to elongated fusoid-ventricose, 36-67 > 10- 
14 ».; tube trama hyaline, obscurely divergent to subparallel, hyphae + 
6 » in diam; pileus trama loosely interwoven, hyaline, homogeneous, hy- 
phae 7 » in diam; cuticle differentiated as a layer of interwoven to more or 
less radically arranged hyphal tips, often incrusted, staining ochraceous to 
tawny in KOH, hyphae + 10 pw in diam; surface of the stipe differentiated 
as a layer of caulobasidia with numerous sterile hyphal tips interspersed, 
staining cinnamon to tawny in KOH, tawny oleiferous hyphae scattered 
throughout; clamp connections not found. 

Chemical Reactions. KOH—negative; NH,OH—negative; HCl]— 
flesh yellowish, cuticle negative; HNO:—negative; Sulfoformalin—flesh 
pink, cuticle negative; FeSO,—negative. 

Material Studied. Marin Co.: Thiers 7504, 10875, 11134. Mendocino Co.: Thiers 
8203, 8462, 8463, 9296. Santa Cruz Co.: Thiers 10755, 10786. San Mateo Co.: 
Peters 399. Sonoma Co.: Thiers 9412. 

Discussion. This species, which appears to fruit under both conifers 
and hardwoods in the coastal forests, is readily recognized by the con- 
spicuously truncate spores, which all carpophores produce, although not 
exclusively since a mixture of spores with truncate and with more or less 
rounded apices has been observed from the same carpophore. X. trun- 
catus belongs to the X. chrysenteron-X . truncatus-X . zelleri complex and 
appears most closely related to X. chrysenteron. As pointed out by Snell, 
Singer and Dick (1959) many workers in the past have confused the 
two species, and, as was done by Coker (1943), often made special men- 
tion that some collections of X. chrysenteron had truncate spores while 
others did not. A re-examination of the collections labelled X. chrysen- 
teron in the herbarium of San Francisco State College has revealed that 
several collections made in Michigan belonged to X. truncatus. 

It has generally been found that X. chrysenteron and X. truncatus 
cannot be distingiushed with certainty in the field. Fruit bodies of both 
species have darkly colored pilei which are usually conspicuously rimose 


246 MADRONO [Vol. 17 


and stipes that are red to reddish, particularly with age. The cuticle of 
both species is composed of interwoven to radially arranged, undifferen- 
tiated hyphal tips which distinguishes them from X. zelleri which has a 
palisade of inflated hyphal tips. 

Although this species was first described from collections made in the 
state of Washington this, as far as can be determined, is the first report 
of its presence in California. 

4, XEROCOMUS CHRYSENTERON (Fries) Quelet, Fl. Mycol. Fr. 418. 
1888. Boletus chrysenteron Fries, Epicr. Syst. 415. 1836-1838. 

Pileus 6—8 cm broad when fully expanded, convex when young chang- 
ing to broadly convex to plano-convex to plane at maturity; surface dry, 
dull, velutinous to subtomentose to tomentose, usually shallowly to con- 
spicuously areolate with age, especially near the margin; colored ‘“Sac- 
cardo’s umber” to ‘‘snuff brown” to occasionally near “‘sepia”’ to “‘bister”’ 
during all stages of development; flesh in cracks on disc pallid, often as- 
suming reddish tints toward the margin when older; margin entire, de- 
curved. Flesh 1—1.5 cm thick on the disc, colored “‘ivory yellow” to “light 
buff” to “pale chalcedony yellow,” unchanging or becoming blue in irregu- 
lar areas; taste not distinctive to somewhat acid and unpleasant; odor not 
distinctive. Tubes arcuate-decurrent to depressed, often narrowly and 
deeply so in older carpophores; up to 1 cm in length; colored near “‘reed 
yellow” to “old gold,” typically changing to blue when bruised; pores 
typically highly irregular in outline and large (1—1.5 mm), concolorous 
with the tubes. Stipe 7-10 cm. long, 1—1.7 cm broad at the apex; equal 
to tapering slightly toward the base; surface dry, glabrous but often 
longitudinally ridged or striate; usually colored ‘“‘primrose yellow” at the 
apex becoming “russet” to “tawny” to “Hays russet” toward the base, 
sometimes pallid at the base and reddish only in mid-portion or entirely 
pallid with only the striations or ridges becoming colored near “eugenia 
red”; whitish to yellow mycelium at the base; solid, flesh pallid to yel- 
lowish at the apex, typically changing to reddish toward the base, un- 
changing when exposed. Spores ochraceous in KOH and Melzer’s reagent, 
ellipsoid to subventricose to subcylindric, not truncate, smooth, 12—13.5 
< 5-6 w; basidia hyaline, 4-spored, clavate, 33-36 7-9 uw; cystidia 
scattered to numerous, common only on the tube mouths, fusoid to fusoid- 
ventricose, hyaline, thin-walled, 56-75 * 10-13 y.; tube trama hyaline, 
obscurely divergent to subparallel, hyphae + 8 y. in diam; pileus trama 
homogeneous, interwoven, hyaline, hyphae + 7 » in diam; cuticle staining 
dark cinnamon brown in KOH, composed of tangled masses of hyphal 
tips which may appear radially arranged, especially when young, often 
heavily and spirally incrusted, hyphae + 10 yp in diam; surface of the 
stipe interwoven, incrusted, staining dark cinnamon brown in KOH; 
clamp connections absent. 

Chemical Reactions. KOH—flesh negative to pinkish yellow, cuticle 
negative or darkening slightly; NH,OH—negative; HCl—flesh nega- 
tive to yellowish, cuticle negative to pinkish; HNO».—flesh and cuticle 


1964] THIERS: XEROCOMUS 247 


negative to pale pink; Sulfoformalin—flesh and cuticle negative to pale 
pink; FeSO,—negative. 

Material Studied. Marin Co.: Thiers 8210. Mendocino Co.: Thiers 8746, 8778, 
8820, 9294, 9343, 9432,9470, 9617 ; Largent 65,92. San Mateo Co.: Thiers 9452,10940. 

Discussion. This is another species found inhabiting the mixed conif- 
erous-hardwood forests along the northern coast, and is a member of the 
complex mentioned earlier. Like X. truncatus it apparently fruits less 
commonly than X. zellert but nevertheless is found relatively frequently, 
particularly in heavily wooded areas. Xerocomus chrysenteron is dis- 
tinguished macroscopically by the dark colored, conspicuously rimose 
pileus that often has some red pigment in the cracks or along the margin, 
and microscopically by possessing spores that have rounded (fig. 4) in- 
stead of truncate apices and a cuticle of interwoven to radially arranged 
hyphal tips instead of a compact palisade of inflated cells. For a further 
discussion of the relationships and characteristics of this species see the 
discussion for X. truncatus and X. gellert. 

5. XEROCOMUS ZELLERI (Murr.) Snell ex Slipp & Snell, Lloydia 7:43. 
1944. Ceriomyces zelleri Murr., Mycologia 4:99. 1912. Boletus zelleri 
Murr., Mycologia 4:217. 1912. Boletellus zelleri (Murr.) Singer, Snell, 
& Dick, Mycologia 51:575. 1959. 

Pileus 5-10 cm broad at maturity; convex to obtusely convex when 
young becoming convex to plane with age; surface dry to moist, conspic- 
uously white pruinose when young often becoming glabrous to obscurely 
tomentose to subtomentose with age; typically markedly rugulose to ver- 
rucose when young, becoming more or less even or smooth at maturity; 
not conspicuously areolate when young but sometimes becoming so with 
age; colored “dusky brown” to “‘fuscous” to “fuscous black” to ‘“‘bone 
brown,” sometimes fading to “army brown” to ‘“‘natal brown,” often with 
a reddish margin; margin entire, often becoming eroded with age, in- 
curved when young becoming decurved at maturity. Flesh up to 1.5 cm 
thick, whitish to pale yellow, unchanging or becoming blue when ex- 
posed; taste not distinctive or mildly acid, odor not distinctive. Tubes 
arcuate-decurrent to depressed, often deeply depressed with age, up to 
1.5 cm in length; colored “olive ocher” when young, changing to ‘‘honey 
yellow” with age, typically becoming blue when bruised or exposed; tube 
mouths highly irregular in outline, typically more than 1 mm; concolor- 
ous with the tubes. Stipe 5-8 cm long, 0.7—1.3 cm broad at the apex; 
equal to occasionally tapering slightly toward the apex; surface dry, typi- 
cally appearing granulose to punctate, especially toward the base; when 
young background typically colored near “warm buff” which becomes 
more or less obscured with granules colored “‘eugenia red,’ sometimes 
colored “acajou red” at the apex, becoming yellowish with red punctae 
toward the base, older stipes typically colored ‘‘vandyke red”’ to ““madder 
brown”; white to pale yellow mycelium at the base; solid; flesh yellow 
when young, typically red when older, sometimes changing to blue in 
irregular areas when exposed. Spores pale yellow to pale ochraceous in 


248 MADRONO [Vol. 17 


KOH, ochraceous tawny in Melzer’s reagent, subellipsoid to subfusoid 
to subventricose, smooth, 12-15 & 4.0—-5.5 mw (occasional giant spores 
measuring up to 24 vin length found in one collection) ; basidia hyaline, 
clavate, 4-spored, rarely 1-3 spored, 18-21 10-11 yw; cystidia rare to 
scattered to numerous, apparently lacking in some carpophores, clavate 
to obtusely fusoid to fusoid-ventricose to obscurely mucronate, hyaline 
to rarely yellowish in KOH, thin-walled, 40-85 « 10-13 uw; tube trama 
parallel to obscurely divergent, with a distinct central strand, hyaline, 
hyphae + 6 yp broad; pileus trama loosely interwoven, homogeneous, 
hyphae + 3 » broad; cuticle staining dark brown in KOH, differentiated 
as a turf of free, septate, erect, inflated hyphal tips, often appearing 
similar to pilocystidia incrusted, typically collapsing and avpearing as 
a tangled mass of hphal tips in older pilei; surface of stipe interwoven, 
heavily incrusted with oleiferous hyphae differentiated throughout; clamp 
connections absent. 


Chemical Reactions. KOH—negative; NH,OH—flesh greenish, cuti- 
cle negative; HCl—flesh yellowish; cuticle dark pink; HNO»—flesh and 
cuticle pink; Sulfoformalin—flesh yellowish, cuticle dark pink; FeSO,— 
negative. 

Material Studied. Marin Co.: Thiers 9825, 10874. Mendocino Co.: Thiers 8165, 
8202, 8290, 8435, 8619, 8790, 8870, 8871, 8993, 9050, 9256, 9289, 9342, 9349, 9722, 
9854, 10051, 10643, 10706, 11017, 11076; Motta 51, 247; Peters 90. Napa Co.: Lar- 
gent 324; Santa Cruz Co.: Thiers 9088; Peters 349. San Mateo Co.: Thiers 7449, 
11199. 

Discussion. As indicated earlier this is one of the most common species 
of Xerocomus in California and fruits from the onset of the rainy period 
in the fall until March and April. It was found most frequently in mixed 
forests, usually in areas where the coastal redwood, Sequoia sempervirens 
(Lamb.) Endl, was relatively abundant. In some instances it was found 
along the margin of slash burns as well as in restricted clearings within 
the forests. 


This fungus appears to be very closely related to X. chrysenteron, and 
microscopic examination of the cuticle is usually necessary to distinguish 
them. Generally, however, X. zed/eri has a darker colored, pruinose pileus 
which typically does not become strongly rimose-areolate. It has been 
found, however, that by far the most reliable distinction is in the differ- 
ence in the structure of the cuticle of the two species. In young pilei of 
X. zelleri the cuticle is differentiated as a closely packed palisade of 
noticeably enlarged hyphal tips (fig. 1) in which the subterminal cell is 
globose to pyriform and the terminal cell is somewhat pyramidal in out- 
line. In older pilei these cells sometimes collapse and the cuticle then 
appears interwoven to radially arranged. In X. chrysenteron, on the other 
hand, the terminal hyphal cells are more or less equal in size and inter- 
woven or more or less radially arranged. Another conspicuous feature 
of the cuticle of X. chrysenteron is the heavy incrustation of the hyphal 
walls which is often deposited in a spiral pattern. 


1964] THIERS: XEROCOMUS 249 


Recently Singer, Snell, and Dick (1959) transferred this species to 
Boletellus chiefly because of their observation that the spores have slight- 
ly roughened walls. Careful attention, including the examination of the 
spores of local collections under oil with phase attachments as well as 
under oil with apochromatic objectives failed to reveal any definite rough- 
ness or wrinkling of the spore wall. It is possible that their collections 
were from a different area and represented a different race, or that the 
roughness was overlooked in the local collections. In would seem, how- 
ever, that if the spores are so obscurely roughened as to escape detection 
after such careful examination that it is placing undue emphasis upon 
such a characteristic to use it as the chief basis for transferring the species 
not only to a different genus but also to a different family. It would appear 
preferable to retain the species in its present genus and emend the concept 
to include species in which the spores are occasionally obscurely rough- 
ened. Thus this species is presently retained in Xerocomus until further 
substantiation warrants placing it with the rough spored species. 

Department of Biology, San Francisco State College, San Francisco, California 


LITERATURE CITED 


Coxer, W. C. and A. H. Beers. 1943. The Boletaceae of North Carolina. Univ. 
North Carolina Press, Chapel Hill. 
Dick, E. A. and W. H. SNELL. 1960. Notes on boletes. XIII. Mycologia 52:444—-454. 
Murrit1, W. A. 1912. The Polyporaceae and Boletaceae of the Pacific coast. Myco- 
logia 4:91-100. 
Rivncway, R. 1912. Color standards and color nomenclature. Washington, D.C. 
SINGER, R. 1940. Notes sur quelques basidiomycetes. Revue Mycol. Paris 5:3-13. 
—. 1954a. The Boletineae of Florida with notes on extralimital species. I. 
The Strobilomycetaceae. Farlowia 2:97-194. 
. 1945b. The Boletineae of Florida with notes on extralimital species. II. 
The Boletaceae (Gyroporoideae). Farlowia 2:223-303. 
. 1962. The Agaricales in modern taxonomy. 2nd edition. J. Cramer, 
Weinheim. 
SNELL, W. H. and E. A. Dick. 1958. Notes on boletes. X. A few miscellaneous dis- 
cussions and a new subspecies. Mycologia 50:57-65. 
, R. Stncer and E. A. Dick. 1959. Notes on boletes. XI. Mycologia 51: 
564-577. 


SURVIVAL OF TRANSPLANTED CUPRESSUS AND PINUS 
AFTER THIRTEEN YEARS IN MENDOCINO 
COUNTY, CALIFORNIA 


CALVIN MCMILLAN 


Survival of Cupressus seedling trees transplanted to the habitat of the 
pygmy forest in Mendocino County was reported after seven years 
(McMillan, 1959). Subsequent observation in 1963 followed a further 
six-year interval. The survival patterns of Cupressus will be discussed 
with a previously unreported companion study in Pinus. 

The transplant site, on the coastal plateau between Little and Albion 
rivers, is within the narrow distribution of Cupressus pygmaea (Lemm.) 
Sarg. and Pinus bolanderi Parl. The pygmy forest of the area (McMillan, 
1956) represents one of the most unusual vegetational situations in Cali- 
fornia. Surrounding the pygmy forest and adjacent to the transplant site 
is the contrasting dense Sequoia-Pseudotsuga vegetation. 

Seedlings of the various Cupressus and Pinus strains (table 1) were 
transplanted from the greenhouse at Berkeley to the pygmy forest in 
November, 1950. Six seedlings of each strain were planted and cov- 
ered by two large wire screen-redwood cages. The seedlings remained 
protected throughout the entire study. Cupressus seedlings removed in 
1954 reduced the number of seedlings in the transplant plot to 5 per 
strain. Some Pinus seedlings transplanted from the Mendocino County 
beach were not protected from browsing animals. 

The local strain of Cupressus showed superior ability to survive in one 
of the most extreme soil situations (pH 3.8—4.0) in California. Table 1 
shows the virtual elimination of all strains except the local one. The only 
surviving plant originating outside the pygmy forest (from Anchor Bay 
in southern Mendocino County) had only one remaining green branch 
in 1963. The strains of Cupressus goveniana Gord., C. abramsiana Wolf, 
and C. sargenti Jeps. were completely eliminated during the study. 

During 1957-1963, height increases were shown by all five trees of 
the pygmy forest strain. Though considerable by pygmy forest stand- 
ards, the height increase was only 2—6 cm, averaging 3.7, for this six- 
year period. All five trees were vigorous and with many side branches 
and their overall height varied from 19.5-28.5 cm. Only one of the five 
trees had mature seed cones. The Anchor Bay seedlings, all surviving 
in 1957, showed little height increase during the 1957-1963 period. The 
condition of the dead Anchor Bay seedlings, most branches and leaves 
intact, suggested fairly recent elimination. 

Of the eliminated seedlings, those which survived the shortest interval 
were of a strain of C. sargenti from serpentine soils (pH 6.5) in Marin 
County. The C. abramsiana strain was from sandy soils (pH 5.2) near 
the town of Bonny Doone in the Santa Cruz Mountains. Both of the 


1964 | McMILLAN: CUPRESSUS AND PINUS Zot 


C. goveniana strains were from acid soils of the Monterey Peninsula. 
The strain of C. goveniana that survived the longer was from the more 
acid soil (pH 4.6), a sandy soil on Huckleberry Hill. The other C. 
goveniana strain, eliminated before 1957, was from a ridge between San 
Jose Creek and Gibson Creek where the yellowish sandy soils (pH 4.9) 
were derived from granodiorites. The Anchor Bay strain of C. pygmaea 
was from a shallow, yellowish sand (pH 4.8) over sandstone bedrock. 

While the reason for the superior fitness of the local strain is not 
readily apparent, it must include a combination of tolerances that is 
unique. Although the elimination of the various strains could not be 
attributed to an obvious biotic factor, some competitive effects of the 
roots of pines and cypresses surrounding the transplant plot were likely. 
Since roots are confined to approximately 6-8 inches above the cemented 
ortstein, it seemed probable that roots extended into the transplant plot 
with some nutrient interaction. During the 13-year period very little 
revegetation had occurred on the transplant plot. In 1963, six small 
seedlings of P. bolanderi were on the plot and several leafy branches of 
Gaultheria shallon Pursh were at the margin of the plot. Hundreds of 
cane-like and cone-bearing trees of C. pygmaea, mostly under 100 cm, 
surrounded the transplant plot, but no cypress germinated on the plot 

Previously unreported was a study of Pzmus seedlings transplanted in 
1950. Included in the series (table 1) were the local pygmy forest strain 
of P. bolanderi (with serotinous cones), the local beach strain of P. con- 
torta Loud. (with cones opening at maturity), a beach strain from 
Oregon, and 2 strains of P. murrayana Grev. & Balt. from the Sierra 
Nevada. In addition, several strains of P. muricata Don. were included: 
the local Mendocino strain from the margin of the pygmy forest, an 
Anchor Bay strain that grew with C. pygmaea in southern Mendocino 
County, and a strain from Monterey County that grew with C. goveniana 
at Huckleberry Hill. 

The pine strains were not as decisively separated in their ability to 
survive as were the cypresses. One complete elimination was of the beach 
strain of P. contorta from Mendocino County. Four seedlings of the 
beach strain that were surviving in 1954 showed poor growth and the 
lone survivor in 1957 had been eliminated by 1963. 

Of the surviving members of the P. contorta complex, those of the local 
pygmy forest strain of P. bolanderi were taller than the other survivors. 
Only two seedling trees of the local strain showed a high degree of vigor, 
one of these having reached a height of 35 cm and the other 19 cm. The 
remainder were under 11 cm. 

When these strains were grown for a year in Berkeley in a control 
soil, superior vigor was shown by the Oregon beach strain, intermediate 
vigor by the Mendocino beach strain, and poor growth by the pygmy 
forest strain (McMillan, 1959). In that study, the Oregon beach strain 
showed an average height increase during one year of 8.7 cm, the Men- 


252 MADRONO [Vol. 17 


TABLE I. SURVIVAL OF TRANSPLANTED CUPRESSUS AND PINUS IN THE 
Pycmy Forest oF MENDOCINO COUNTY, CALIFORNIA 


Strain Number of surviving seedling trees 
19501 1952 19542 1957 1963 

C. pygmaea 

Pygmy Forest 6 6 oe (S) 5 5 

Anchor Bay 6 6 61) 5 1 
C. goveniana 

Huckleberry Hill 6 6 6 (5) 4 ‘@l 

Point Lobos 6 6 6-5) 0) 0) 
C. abramsiana 

Bonny Doone 6 6 4 2 0 
C. sargentii 

Mt. Tamalpais 6 5 0 0 0 
P. bolanderi? 

Pygmy Forest #1 (ME-20) 6 6 6 4 2 

Pygmy Forest #2 (ME-20) 6 6 6 3 3 
P. contorta 

Mendocino (ME-101) 6 6 4 1 0 

Tillamook Co., Oregon (TL-107) 6 6 6 4 2 
P. murrayana 

Sierra Nevada (PA-102) 6 6 4 2 1 

Sierra Nevada (PA-103) 6 6 4 2 0 


P. muricata 
Mendocino (ME-21) 
Noyo (NO-110) 
Anchor Bay (AB-26) 
Huckleberry Hill, Monterey Co. 
(HH-28) 


DAW OD 
DN UN OW 


DnBRAAD 
awe a 
toe eh 


1 Transplanted from Berkeley greenhouse in Nov., 1950, except C. sargentii added April, POST. 
2 One seedling of each of the two C. pygmaea strains and of each of the two C. goveniana 
pins were removed from the plots in 1954 for doucmentation. 
3 Series 7/1 planted with cypress seedlings: series ##2 planted with pine seedlings. 


docino beach strain, 3.3 cm and the pygmy forest strain, 1.5 cm. The 
alteration of these growth patterns in the pygmy forest habitat sug- 
gests the selective significance of unique tolerances of the pygmy forest 
strain. 

In P. muricata, decisive superior fitness was not shown by any of the 
strains (table 1). Some seedlings of each of the strains were able to 
tolerate the extreme conditions of the transplant site for 13 years, though 
none was vigorous. The distribution of P. muricata at the margin of the 
pygmy forest, in deeper and less acid soils, suggested an inability to 
tolerate the pygmy forest soils. Large trees of the local strain occur 
within 30 feet of the transplant plot, though seedling trees are not pres- 
ent in the immediate vicinity of the plot. Of the five pine seedlings that 
germinated on the plot during the 13 year study, all were P. bolandert. 

In addition to the seedling transplants, young trees of the beach strain 
of P. contorta were transplanted to the pygmy forest habitat. Seven 
were transplanted with soil around the roots, and seven were transplanted 
without soil. Soon after transplanting in November, 1950, considerable 


1964] McMILLAN: CUPRESSUS AND PINUS BSS) 


damage to the 14 trees was caused by browsing animals. Trees of 
P. bolanderi in the immediate area were not browsed. 

After four years, all of those transplanted with soil around the roots 
were vigorously surviving. At the same time only four of the seven trans- 
planted without soil were surviving and none of these was vigorous. Only 
two of the seven, bare-root transplants were surviving in 1963. Neither 
was vigorous, one measuring 8 cm in height and the other 38 cm. 

Of the seven transplanted with soil, the recorded heights in 1957 were 
70-110 cm, averaging 95 cm. In 1963, they measured 90-140 cm, aver- 
aging 107 cm. In spite of the height increase shown by each of the seven 
plants from 1957 to 1963, one plant had died and three others were 
with large dead portions at the top of the plants. The three plants with 
seed cones reacted as plants in the beach population, the cones opening 
at maturity. 

The severity of the pygmy forest habitat has been a selective force 
that has shaped a unique vegetational cover. This 13-year survival study 
emphasizes the unique tolerance of the local pygmy forest strain of 
Cupressus. Although it is well known that C. pygmaea may become a 
large tree, the ability to sustain itself in the very acid soils of the pygmy 
forest is not shared by other strains of cypress. Although the 13-year 
study had not led to the complete elimination of all other strains of the 
P. contorta complex, the superior growth of the local strain suggests its 
possible lone success. The survival of the beach strain for 13 years when 
transplanted with its native soil suggests the restrictive role of the pygmy 
forest soils. The current lack of vigor of the beach pines suggests the 
eventual loss of the protective influence of its native soil. The survival 
of some seedlings of P. muricata for the entire 13-year period was least 
anticipated because of the natural restriction of this species from the 
most acid soils of the pygmy forest. 

Although this study had not been planned as a long-range evaluation 
of survival, it does suggest the value of projecting such studies. Often 
short-term soil evaluations suggest broad tolerance ranges by woody 
plants. This study suggests more critical plotting of populational survival 
by longer examination. 


Department of Botany and Plant Research Institute, University of Texas, Austin 


LITERATURE CITED 


McMittan, C. 1956. The edaphic restriction of Curpressus and Pinus in the Coast 
Ranges of central California. Ecol. Monogr. 26:177-212. 
. 1959. Survival of transplanted Cupressus in the pygmy forests of Mendo- 
cino County, California. Madrono 15:1-4. 


A PECULIAR CASE OF HEMLOCK MISTLETOE 
PARASITIC ON LARCH 


Jos Kuljt 


The British Columbia Forest Service in 1951 undertook to plant some 
Larix europaea in several localities in the Cowichan Lake area on Van- 
couver Island. One such locality was near the entrance of the Forest 
Experimental Station at Mesachie, where trees were planted in the 
immediate vicinity of old trees of Tsuga heterophylla severely infected 
with Arceuthobium campylopodum. Young hemlocks, part of the natural 
regeneration, also bore many vigorous infections or were about to become 
infected. In the spring of 1961 a small number of swellings were dis- 
covered on two of the larches. The largest swelling was a fusiform one 
of about 3 in. in length and 1 in. in thickness. One or two lateral branches 
which took their origin in the swollen area themselves showed some 
slight hypertrophy at the base. The striking thing at the time of the 
1961 visit was the fact that no mistletoe shoots were present on any of 
the swellings. Consequently, some doubt remained as to the identifica- 
tion of the hypertrophies. 

The area was revisited again two years later, in July, 1963. A total of 
10 swellings were discovered on three larches. Some of these doubtlessly 
escaped detection during the earlier visit, but others had clearly devel- 
oped since that time. The two largest infections had undergone little 
change except that the formation of a typical small broom was now under 
way in both cases (fig. 3). Short shoots on the swollen portion of the 
host branch had grown into long shoots (fig. 2), a process similar to 
that in some pines when attacked by A. campylopodum (Kuijt, 1960). 
A certain amount of necrosis and resin flow had occurred. Most sur- 
prisingly, a careful search revealed the complete absence of mistletoe 
shoots and buds. 

It has been established since that time that the swellings are inhabited 
by the endophytic strands of Arceuthobium. The extra-cambial tissues 
of the host are traversed by many “cortical” strands, and many sinkers 
extend at least two or three years into the xylem. Abnormal rays are 
very frequent (Srivastava & Esau, 1961); in fact, normal ones can be 
located only with difficulty. The cells of the parasite in these abnormal 
rays are crowded with starch grains. 

The most mystifying aspect of these infections is, of course, the com- 
plete absence of shoots or even the smallest buds on mistletoe individuals 


Fics. 1-3. Parasitism of Arceuthobium campylopodum on Larix europaea, 
Mesachie, British Columbia; 1, old infection with some indication of early brooming ; 
cracking and rough texture of host trunk may indicate presence of the parasite within 
it; 2, very young infection showing fusiform swelling and change from short shoot 
to long shoot; 3, well developed young broom. 


1964] KUIJT: HEMLOCK MISTLETOE 255 


256 MADRONO [Vol. 17 


which are estimated to be about 6 years old. Even in the case of the 
5-year life cycle of the species on Pinus monticola (Kuijt, 1961) mistle- 
toe buds were developed almost immediately after the hypertrophy was 
initiated, i.e., within two or three years following seed dispersal. In the 
larch infections, however, we are faced with infections even at the broom- 
ing stage without the slightest indication of buds. Although aerial mistle- 
toe shoots may still appear in the years to come, we may be concerned 
here with a completely endophytic (“latent”) mistletoe. The fact that 
the host branches of one infection had outgrown the swollen and some- 
what broomed portion may support the contention of sterility of the 
mistletoe on this host (fig. 1). 

The idea of latency in mistletoes is not a novel one. It has been alluded 
to both for Phoradendron juniperinum (Wagener, 1925), and Arceutho- 
bium. Gill & Hawksworth (1961), for example, say: ‘The ability of the 
haustoria to live for long periods without benefit of aerial parts is well 
known in Arceuthobium.” But in all such cases the period of latency 
seems to be a terminal, senescent one (Kuijt, 1960). This is implied by 
Gill (1935) when saying that in Arceuthobium “the endophytic system 

. may continue to live for years after shoot production has ceased.” 
The present larch infections may never produce shoots. 

These observations do not provide an unambiguous answer as to the 
question of host-specific races within A. campylopodum. It is true, on 
the one hand, that the hemlock mistletoe can become established on at 
least one species of larch. The physiological distinction, if any, separat- 
ing “‘f. daricis” and “‘f. tsugensis” is therefore by no means complete. On 
the other hand, the behavior of the hemlock mistletoe on the European 
larch appears to be such as to preclude self-perpetuation. We must keep 
in mind here the possibility that the response of L. europaea to infection 
by A. campylopodum may be quite different from that of L. occidentalis. 
In other words, it is conceivable that the life cycle on the latter is rela- 
tively short, while development of shoots is delayed on the former host. 


University of British Columbia, Vancouver 


LITERATURE CITED 


GILL, L. S. 1935. Arceuthobium in the United States. Trans. Conn. Acad. 32:111-245. 

Git, L. S. and F. G. HAwksworTtH. 1961. The mistletoes: a literature review. 
U.S. Dept. Agr. Forest Serv. Tech. Bull. 1242. 

KurjT, J. 1960. Morphological aspects of parasitism in the dwarf mistletoes (Arceu- 
thobium). Univ. Calif. Publ. Bot. 30:337-436. 

. 1961. Observations on the life cycle in Arceuthobium campylopodum. Leafl. 
West. Bot. 9:133-134. 

Srivastava, L.M. and K. Esau. 1961. Relation of dwarfmistletoe (Arceuthobium) 
to the xylem tissue of conifers. II. Effect of the parasite on the xylem anatomy 
of the host. Am. Jour. Bot. 48:209-215. 

WAGENER, W. W. 1925. Mistletoe in the lower bole of incense cedar. Phytopathology 
15:614—616. 


LYONOTHAMNOXYLON FROM THE LOWER PLIOCENE 
OF WESTERN NEVADA 


VIRGINIA M. PAGE 


At the northern end of Fish Lake Valley, Nevada, a desolate, arid val- 
ley between the White Mountains to the west and the Silver Peak Range 
to the east, there is a curiously eroded gully containing the remains of 
about seventy fossil trees (fig. 1). Although the trees are badly weath- 
ered and fragmented, it is possible to observe that all were preserved 
in situ, some in a standing position and others lying where they were 
felled by some agent such as a storm or flood. 

Although the existence of the trees has been known for many years, 
no published descriptions have been found in the literature, nor, as far as 
I can determine, has there been any attempt to identify the wood. Dur- 
ing a brief visit to the fossil locality in 1955 and a recent visit in 1964, 
fragments of wood were collected from each tree. The following account 
is based on a study of thin sections made from some of the fragments.* 


GEOLOGIC OCCURRENCE 


The fossil locality is in Esmeralda County, T 1 N and R 35 E, three 
fourths of a mile south of hill 6061 on the Davis Mountain topographic 
sheet of the U.S. Geological Survey in the vicinity of the University of 
California Museum of Paleontology locality V—2804. The trees are em- 
bedded in a thick layer of sandstone several feet below the surface layer 
of white vitric tuff which in that particular area is one to two feet thick. 
Both sandstone and tuff appear to have been stream deposited. The beds 
dip to the southeast at an angle of forty to fifty degrees from the hori- 
zontal (fig. 1). 

Several feet above the tree horizon and just below the layer of tuff the 
sandstone is green in color and contains mammalian bones. Stirton (1929; 
1939) considered the fauna lower Pliocene in age and referable to the 
early Clarendonian using standard North American geochronology. 
Everndon et al (1964) using potassium-argon dating techniques found 
that biotite in the vitric tuff just below a micro-mammal layer in the 
vicinity of the fossil tree horizon gave a reading of 11.4 million years 
before Present. According to recent observations (Richard Tedford, per- 
sonal communication) the dated biotite layer is, as far as can be deter- 
mined, the same unit which occurs directly above the stump horizon and, 
therefore, also directly above Stirton’s mammalian layer, thus corroborat- 
ing his assignment of an early Pliocene age. 

The fallen trees lie in a southeasterly direction. Some of the trunks are 
six or more ft. long, but no branches have been observed, nor is bark pres- 
ent. The upright stumps consist of the very base of the trunk and the 
basal root system. The surrounding matrix has been eroded away several 


1 This work was supported in part by National Science Foundation grant GB184. 


258 MADRONO [Vol. 17 


ft below the original ground level, so that now some of the stumps rest 
on tall sandstone columns as much as 10 ft in circumference and over 
15 ft high (fig. 2). Since all the trees lie at the same horizon in the sand- 
stone, it can be assumed that they were contemporaries. 


SYSTEMATIC DESCRIPTION 

For the most part the wood collected is not well preserved. Many of 
the specimens are weathered and chalky with little or no cellular struc- 
ture remaining. Even in those specimens which are reasonably well pre- 
served the secondary walls of most of the cells are absent. It is possible, 
however, to determine that all the specimens are of the same kind of 
wood. Since samples were collected from at least seventy trees, it looks 
as though all the trees were of one kind. 

In comparing thin sections of the fossil wood with sections of various 
modern woods it was found that the structural pattern of the fossil is 
similar to that of certain woody members of the Rosaceae. In detail of 
pattern the secondary wood of the monotypic genus Lyvonothamnus ap- 
pears to resemble the fossil most closely. 

Lyonothamnoxylon nevadensis gen. et sp. nov. (figs. 3-6). Growth 
rings narrow, averaging less than 1 mm. in width; detectable with hand 
lens. Diffuse porous to semi-ring porous. Pores solitary, rounded or slight- 
ly angular averaging 27 by 44. (14 & 22 — 37 & 55) in diameter with 
longer axis in radial direction. Vessel elements 184-300 y.in length. Vessel 
walls were difficult to measure because of the absence of secondary walls 
in most cells, but on the basis of a few fragmentary walls it appears that 
they are about 8 p. thick. Intervascular pitting small and alternate. In two 
vessels it is possible to observe what are undoubtedly remnants of spiral 
thickening. Gum plugs present in nearly all vessels. Perforation plates 
simple, almost horizontal in large vessels to oblique in small vessels with 
“tails” evident in some elements. Axial parenchyma metatracheal-diffuse, 
6-8 cells high; small pits on tangential and radial walls. Rays hetero- 
geneous and homogeneous 2-4 cells wide (characteristically 2 cells wide). 
Heterogeneous rays with 2—4 marginal rows of square cells. Cells of some 
rays irregular in size; marginal cells tend to be larger than those of multi- 
seriate part. Height averages 427 ». (some almost 1 mm). Frequently 
two rays appear to be joined vertically. Uniseriates 1-20 (mostly 6-8) 
cells high, cells isodiametric in tangenital view. Ray to vessel pitting 
small, alternate. Fibers or fiber tracheids make up bulk of ground tissue; 
average 430 y» in length. Bordered pits occur on both tangential and 
radial walls. A few cells are sufficiently well preserved to show that the 
walls are fairly thick. 


Holotype: Nevada, Esmeralda Co., Fish Lake Valley, V. M. Page 5567, July 
1955, Stanford Univ. Paleont. Type Coll. 8425. 


DISCUSSION 


Although the fossil compares well with the secondary wood of Lyono- 
thamnus (figs. 7-10), there are a few differences. These differences are 


1964 | PAGE: LYONOTHAMNOXYLON 259 


SAR eee at BS Ss 


Fic. 1. Northend of Fish Lake Valley. Just to right of center is a tall column 
topped by a fossil stump. Behind the stump and at the same horizon along a line 
7 mm below surface layer is a series of knobs which represent exposed stumps. 


not major and may reflect either environmental differences, position in 
the tree from which the specimens were derived, or genetic changes that 
have occurred in the genus during the time interval between the lower 
Pliocene and the present. The main differences can be found in the con- 
sistently broader rays of the extant species (mostly 3-4 cells wide), ap- 
parently more abundant axial parenchyma, and more pores per mm? with 
a tendency toward radial chains in spring wood. Considerable variation 
was observed in the samples of modern material available. Rays, for 
example, vary from predominantly two cells wide in slide 9329 from the 
Arnold Arboretum wood collection to eight cells wide in a small branch 
from a tree growing on the Stanford campus. Some samples are distinctly 
ring porous, whereas in others there is little difference in pore size be- 
tween early and late wood (figs. 7, 8). The amount of parenchyma also 
varies from one sample to another. Similar variations can be observed 
among the many fossil specimens collected, although for the most part 
the over-all structural pattern is quite uniform. Such uniformity is to be 
expected, since all the fossil trees presumably were subjected to the same 
environmental conditions, whereas the trees from which the reference 
samples were taken were growing in a variety of habitats. Although the 
degree of variation was greater among the modern samples, such varia- 


260 MADRONO [Vol. 17 


tion as has been observed is of the same kind in both the fossil and the 
extant species. 

No evidence of spiral thickening was found in the fibers of the fossil, 
nor were septae observed. Spiral thickening occurs in most fibers and 
septae in a few in the extant species of Lyonothamnus. Their absence in 
the fossil is to be expected because of the absence of the secondary wall 
in the majority of cells. Macerations of the secondary wood of L. flori- 
bundus A. Gray ssp. asplenifolius (Greene) Raven and another member 
of the Rosaceae, Heteromeles arbutifolia (Lindl.) M. Roem., show that 
there is a series of transitional forms between tracheids and vessel ele- 
ments on the one hand and tracheids and fibers on the other. These tran- 
sitional forms are difficult to recognize in sections, particularly in the 
fossils, but because the elongated xylary elements in the fossil bear evi- 
dence of conspicuously bordered pits, it is assumed that similar transi- 
tional forms are present; that is, some of the pitted structures may be 
fiber tracheids, some may be fibers, and others may be narrow vessel 
elements. 

There are several genera in the Rosaceae whose wood bears a general 
resemblance to that of the fossil. Most of these, however, can be elimi- 
nated from consideration after examining the composition of the Pliocene 
vegetation of the western Great Basin where only a few genera of the 
Rosaceae are represented in the fossil record, and these, with the excep- 
tion of Lyonothamnus have their modern equivalents in non-arborescent 
forms. It has been shown that by Pliocene time the floral elements now 
represented in eastern United States and eastern Asia and also those 
now in northern areas, which constituted a substantial part of the early 
Miocene floras, were largely eliminated from southwestern United States. 
Equivalents of most of the species present in the western Great Basin in 
early Pliocene time can still be found in southwestern United States, 
although their ranges have been somewhat restricted. It would follow, 
therefore, that woody plants now occurring in that area might be expected 
to appear in late Tertiary deposits of the same general region. In addi- 
tion to Lyonothamnus, Heteromeles arbutifolia is the only arborescent 
member of the Rosaceae found in southwestern United States whose wood 
bears a resemblance to the fossil. Thus far Heteromeles has not been 
found in any of the Pliocene or Mio-Pliocene deposits in the western 
Great Basin, whereas Lyonothamnus has appeared in three localities. 
All three of these localities (Aldrich Station, Middlegate, and Stewart 
Springs) are Mio-Pliocene in age, thus slightly older than the fossil wood 
locality, and in all three the genus is represented by leaf impressions. 
Leaf fragments ascribed to Lyonothamnus from the Aldrich Station and 
Middlegate floras (Axelrod, 1956) were subsequently thought to be 
Comptonia. They have now been returned to Lyonothamnus (Wolte, 
in press). 

Some differences between the wood of Heteromeles and the fossil indi- 
cate that close relationship between them is questionable. Vessel elements 


1964] PAGE: LYONOTHAMNOXYLON 261 


Fic. 2. Group of stumps resting on sandstone columns. Note that stump to the 
left appears to have two trunks. 


in the secondary wood of Heteromeles average 600 » in length, whereas 
those of the fossil average 270 wu. The latter figure is more consistent with 
the 312 uw average in Lyonothamnus. Fibers in Heteromeles tend to be 
longer (825 yw as opposed to an average of 430 yp. in the fossil and in 
Lyonothamnus), and the uniseriate rays tend to be shorter (3-8 cells 
high as opposed to 1—18 cells high in the fossil and in Lyonothamnus). 
Comparisons were made of samples of Heteromeles collected from plants 
on the Stanford campus, a root and stem from Panoche Pass, California 
and samples from three trees of L. floribundus ssp. asplenifolius growing 
on the Stanford campus. Although vessel length is of doubtful value in 
wood identification, it is felt that, because the feature is consistent among 
the several specimens sampled of each species, the difference is real and 
can, threfore, be used together with the disparity in fiber length and ray 
height to point out the difference between the wood of Heteromeles and 
the fossil while at the same time emphasizing the similarity between the 
wood of Lyonothamnus and the fossil. 

Growth habit provides another link between the fossil trees and Lyono- 
thamuus. As described by Millspaugh and Nuttall (1923), L. floribundus 
ssp. floribundus forms groves consisting of about 50 trees on canyon sides 
in its native habitat. That the fossil tree locality represents the remains 
of an extensive grove can be assumed from the evidence that all the trees 


262 MADRONO [Vol. 17 


. sa St dee! _ 
Fics. 3-6. Lyonothamnoxylon nevadensis: 3, transverse section showing slightly 
angular pores, semi-ring porous distribution; 4, transverse section showing rounded 
pores and diffuse porous distribution; 5, radial section; 6, tangential section; 4-6, 
from holotype. All are x 160. 


1964 | PAGE: LYONOTHAMNOXYLON 263 


Fics. 7-10. Lyonothamnus floribundus ssp. asplenifolius: 7, transverse section 
showing ring porous distribution of pores; 8, transverse section showing diffuse 
porous distribution; 9, radial section; tangential section. All * 160; 8, 9, and 10 
from one specimen, 7 from another. 


264 MADRONO [Vol. 17 


lie at the same horizon in the sediments, they were preserved where they 
grew, and only one kind of tree is represented. 

The fossil record shows that Lyonothamnus had a wide distribution 
throughout western United States during the middle and late Tertiary 
(Axelrod, 1940a; Wolfe, in press), and was particularly in evidence dur- 
ing the late Miocene in western Nevada only 50 miles north of Fish Lake 
Valley. Its maximum extension so far recorded was during the Miocene 
when it ranged from west-central Washington south to southern Califor- 
nia and eastward into western Nevada. For reasons not entirely under- 
stood, its range became greatly restricted since the mid-Pliocene. In- 
creased aridity and colder winters undoubtedly contributed to its dis- 
appearance in the eastern part of its range, and the increasing depression 
of winter temperatures towards the end of the epoch probably affected 
its northern distribution. At the present time it occurs naturally only on 
the channel islands off the coast of southern California. 

Morphological variations in the leaves of Lyonothamnus in different 
parts of its range through space and time suggest that there were genetic 
variants in the genus at least until late Pliocene; even now there are two 
subspecies (Raven, 1963). Axelrod (1958) has shown that leaves of the 
mid-Miocene L. mohavensis Axelrod were half the size of the modern 
L. florioundus ssp. asplenifolius and the upper Pliocene form from coastal 
central California was intermediate. Wolfe (in press) describes a dis- 
tinctly different form from the late Miocene Stewart Spring flora from 
west-central Nevada which he calls L. parvifolius. These leaves charac- 
teristically have 7—9 leaflets as opposed to five in the modern species. 
There is also a difference in lobation. Wolfe suggests that the northern 
segregates of the genus became extinct, while the southern form (or 
forms) gave rise to the modern species. 

Whether the trees in Fish Lake Valley belonged to the northern or 
southern segregates one cannot say on the basis of the information avail- 
able from the wood. Inasmuch as we are dealing with only one species 
whose associates remain unknown, we can only speculate as to where its 
affinities lie. 

The proximity of the Stewart Spring locality to that of the fossil trees 
may suggest a close relationship floristicly despite the slight difference 
in age. The Stewart Spring flora, according to Wolfe, is mesic in aspect 
and, although distinct from contemporary floras to the north and south, 
is more closely allied to the northern floras. The early Pliocene Esmeralda 
flora (Axelrod, 1940b) from the east flank of the Silver Peak Range a 
short distance east of Fish Lake Valley is arid in aspect and appears to 
have affinities to the south. All but two of the thirteen species described 
occur also in the mid-Miocene Tehachapi flora 200 miles to the south, 
whereas only five occur in the Stewart Springs flora. The Esmeralda flora 
is very small and may not give an adequate picture of the vegetation of 
the time. It is interesting to note that growth rings in the fossil wood 
fragments are narrow and average less than a mm in width, suggesting 


1964] PAGE: LYONOTHAMNOXYLON 265 


the possibility that the climate was somewhat dry during the lifetime of 
the trees. 

It is highly probable that the trees in Fish Lake Valley are not con- 
specific with the extant species of Lyonothamnus. It is quite possible, 
also, that more than one species existed in the southern Great Basin dur- 
ing late Miocene time and possibly into early Pliocene time. 

It is tempting to try to reconstruct the sequence of events that led to 
the preservation of this ancient grove of trees. In western Nevada 11 
million or so years ago the Sierra Nevada did not present the great bar- 
rier to circulation of moisture-laden air from the ocean that it does now. 
Narrow growth rings point to a fairly dry climate, but contemporary 
faunal beds show that the surrounding vegetation supported a wide vari- 
ety of animals such as camels, horses, dogs, cats, bats, beavers, rhinoceros, 
etc. (Stirton, 1939). That there was some topographic relief in the area 
where the trees grew is indicated by the depth of the alluvial sediments 
in which they are buried. 

The trees all appear to have fallen in the same direction as if felled 
at the same time and by the same agent, such as a flash flood or a wind 
or both. The position of the trees indicates that the destructive force 
came from the northwest. They were not deeply buried at first, for the 
absence of crown, bark, and secondary cell walls shows that a certain 
amount of destruction by microorganisms or other agents had occurred 
prior to final and complete burial to a depth at which aerobic saprophytes 
could not function. Further degredation was halted; the tissues subse- 
quently became infiltrated with silica-bearing water; and the process of 
preservation was begun. Later the whole area was uplifted. The soft sedi- 
ments offered little resistance to the erosion which resulted in the inter- 
esting sculptured effects that can be observed there today and in the 
exposure of the now thoroughly silicified stumps and fragmented logs. 


Department of Biological Sciences, Stanford University 


LITERATURE CITED 


AXxELrop, D. I. 1939. A Miocene flora from the western border of the Mohave 
desert. Carnegie Inst. Publ. 516. 
. 1940a. A record of Lyonothamnus in Death Valley. Jour. Geol. 48:526-— 
531, 
—.1940b. The Pliocene Esmeralda flora of west-central Nevada. Jour. Wash. 
Acad. 30:163-174. 
-- 1956. Mio-Pliocene floras from west-central Nevada. Univ. Calif. Publ. 
Geol. Sci. 33:1-322. 
. 1958. Evolution of the Madro-Tertiary geoflora. Bot. Rev. 24:433-509. 
EverNpoN, J. F., D. E. Savace, G. H. Curtis, and G. T. JAMEs. 1964. Potassium- 
argon dates and the Cenozoic mammalian chronology of North America. Am, 
Jour. Sci. 262:145-198. 
MitispaucH, C.F., and L. W. Nuttatt. 1923. Flora of Santa Catalina Island, Cali- 
fornia. Fieldiana Bot. 212V:1-390. 
RAven, P.H. 1963. A flora of San Clemente Island, California. Aliso 5:273-347. 


266 MADRONO [Vol. 17 


STirRTON, R. A. 1929. Artiodactyla from the fossil beds of Fish Lake Valley, Nevada. 
Univ. Calif. Geol. Sci. Bull. 18:291-302. 
. 1939. Nevada Miocene and Pliocene mammalian fauna as faunal units. 
6th Pacif. Sci. Congr. Proc. 2:627-640. 
Wo re, J. A. (in press). Miocene floras from Fingerrock Wash, southwestern Nevada. 
U.S. Geol. Survey Prof. Pap. 454N. 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 


Beginning with this issue, Documented Chromosome Numbers of Plants will 
appear in a new format. The reasons for this are several: 1. preparation of the man- 
uscript for the printer is easier and there is less chance for error; 2. a paragraph 
rather than a tabular arrangement is less expensive to set in type; and 3. additional 
kinds of information can be included more readily. At times it may be desirable to 
include a sentence or two explaining the significance of a particular count, to com- 
ment on it briefly, to include a photomicrograph or a figure, or to make a new 
combination. It is to be understood that the collector and the counter are the same 
unless otherwise noted. For further instructions see Madrono 9:257-259. 1948. 


Aquilegia nevadensis Boiss. 2n = 14. Spain, Sierra Nevada, Puerto de la Ragua. 
Merxmuiuiller & Grau in 1962, M. Counted by Grau. 

Artemisia nana Gaud. 2n = 18. Switzerland, Wallis, Saas. Zollitsch in 1961, M. 
Counted by Damboldt. 

Bouteloua breviseta Vasey. n= 20. Texas, Presidio Co. F. W. Gould 9718, 
TAES. 

B. rothrockii Vasey. n = 20. Arizona, Pinal Co. F. W. Gould 10028, TAES. 

Bromus macrostachya Desf. n= 28. Texas, Brazos Co. F. W. Gould 9513, 
TAES. 

Carduus argyroa Kze. 2n = 26. Italy, Sicily, Syracuse. Podlech in 1961, M. 

C. velebiticus Borb. 2n = 16. Yugoslavia, Dalmatia, Zadar. Podlech in 1960, M. 

Carex acutiformis Ehr. 2n = 78. Germany, Bavaria, Miinchen. Hertel in 1963, 
M. Counted by Dietrich. 

C. alba Scop. 2n = 54. Germany, Bavaria, Pupplinger Au. Klofat in 1963, M. 
Counted by Dietrich. 

C. argyroglochin Horn. 2n = 68. Germany, Bavaria, Garmish. Oberwinkler in 
1963, M. Counted by Dietrich. 

C. atrofusca Schkur. 2n = 40. Switzerland, Silvretta, oberes Fimbertal. Dietrich 
in 1963, M. 

C. austroalpina Bech. 2n = 38. Italy, Riveria, Alassio. Podlech in 1960, M. 
Counted by Dietrich. 

C. brizoides Jusl. 2n = 58. Germany, Bavaria, Reichenhall. Oberwinkler in 1963, 
M. Counted by Dietrich. 

C. camposii Boiss. & Reut. 2n = 68. Portugal, Sierra de Maraon. Poelt in 1962, 
M. Counted by Dietrich. 

C. cuprea (Kiik.) Nelm. 2n = 70. Nyasaland, Lake Kauline, Nyaka Plateau. 
Robinson in 1959, M. Counted by Dietrich. 

C. curvata Knaf. 2n = 58. Germany, Bavaria, Regensburg. Dietrich in 1963, M. 

C. durieui Steud. 2n = 52. Spain, Prov. Oviedo, El Pedregal. Merxmiuiller in 
1962, M. Counted by Dietrich. 

C. ericetorum Poll. var. approximata (All.) K. Richt. 2n = 30. Switzerland, 
Graubiinden, Ardez. Dietrich in 1963, M. 

C. flavella Krecz. 2n = 60. Italy, Dolomites, Prodoijoch. Podlech in 1961, M. 
Counted by Dietrich. 


1964 | CHROMOSOME NUMBERS 267 


Carex frigida All. 2n = 58. France, Dep. Alpes Maritimes, Tende. Poelt in 1963, M. 
Counted by Dietrich. 

C. grioletii Roem. 2n = 48. Italy, Riviera, Val Seborino. Patzke in 1963, M. 
Counted by Dietrich. 

C. halleriana Asso. 2n = 50. Italy, Riviera, Alassio. Podlech in 1963, M. Counted 
by Dietrich. 

C. mucronata All. 2n = 34. Austria, Lechtaler Alpen, Stockach. Oberwinkler in 
1963, M. Counted by Dietrich. 

C. ornithopoda Willd. var. elongata (Leyb.) A. & G. 2n= 54, Switzerland, 
Graubiinden, Ardez. Dietrich in 1963, M. 

C. parviflora Host. 2n = 54. Switzerland, Silvretta, oberes Fimbertal. Dietrich 
in 1963, M. 

C. pauciflora Lightf. 2n = 76. Austria, Silvretta, unteres Fimbertal. Dietrich in 
1963, M. 

C. praecox Schreb. 2n = 58. Germany, Bavaria, Regensburg. Dietrich in 
1963, M. 

C. supina Wahlenb. 2n = 38. Italy, Vintschgau, Laas. Wiedmann & Grau in 
1963, M. Counted by Dietrich. 

C. umbrosa Host. 2n = 62. Germany, Bavaria, Munchen. Oberwinkler & Diet- 
rich in 1963, M. Counted by Dietrich. 

Chaetopappa asteroides DC. n= 8. Kansas, Woodson Co. E. B. Smith 289, 
KANU. 

Crocidium multicaule Hook. n= 9,,. California, Trinity Co., 3 mi e of Burnt 
Ranch. R. Spellenberg 32, UC. 

Digitaria patens Henr. (Trichachne patens Swallen). n= 36. Texas, Mason 
Co. F. W. Gould 8684, TAES. 

Eleocharis acicularis R. & S. n= 10. Kansas, Harvey Co. Harms & McGregor 
1222, KANU. Counted by L. J. Harms, KANU. 

E. arenicola Torr. n= 10. Mexico, San Luis Potosi, near Tamazunchale. L. J. 
Harms et al. 730, KANU. 

E. atropurpurea (Retz.) J. & C. Presl. n= 10. Kansas, Harvey Co. Harms & 
McGregor 1228, KANU. Counted by L. J. Harms, KANU. 

E. compressa Sulliv. n = 9. Kansas, Clark Co. L. J. Harms 1121, KANU. 

E. engelmannii Steud. n = 5. Kansas, Cherokee Co. L. J. Harms 1175, KANU. 

E. nervata Svens. n = 5. Mexico, San Luis Potosi, near Tamazunchale. L. J. 
Harms et al. 731, KANU. 

E. obtusa (Willd.) Schultes. n = 5. Kansas, Bourbon Co. L. J. Harms 1091, 
KANU. 

E. parvula (R. & S.) Link var. anachaeta (Torr.) Svens. n= 4. Kansas, Re- 
public Co. L. J. Harms 1209, KANU. 

Eragrostis lehmanniana Nees. n = 20. Texas, Hudspeth Co. F. W. Gould 9535, 
TAES. 

Eriochloa punctata (L.) Desv. n= 18. Texas, San Patricio Co. F. W. Gould 
0507, LABS. 

Franklinia alatamaha Marsh. n= 18. Hort. Arnold Arb. R. Dudley 2728-3-C, 
AAH. Counted by L. Riidenberg, GH. 

Helianthus debilis Nutt. var. cucumerifolius (T. & G.) Gray. n=17. Texas, 
Wharton Co. EF. B. Smith 255, KANU. Counted by R. R. Johnson and E. B. Smith, 
KANU. 

Jaumea carnosa (Less.) Gray. 2n = 19,,. California, Humboldt Co., near Hum- 
boldt Bay. D. E. Anderson 2812, UC. 

Lithospermum cinereum DC. 2n = 26. Southwest Africa, Windhoek. Volk in 
1956, M. Counted by Grau. 

Lolium temulentum L. n = 7. Texas, Anderson Co. F. W. Gould 9521, TAES. 

Lycurus phleoides Kunth. n = 20. Texas, Pecos Co. F. W. Gould 9731, TAES. 

Moehringia dasyphylla Bruno. 2n = 24. France, Dep. Alpes Maritimes, Fontan. 


268 MADRONO [Vol. 17 


Grau in 1932, M. 

Moehringia markgrafii Merxm. & Guterm. 2n = 24. Italy, Brescian Alps, Vestone. 
Merxmiiller & Wiedmann in 1956, M. Counted by Grau. 

M. papulosa Bert. 2n = 24. France, Dep. Alpes Maritimes, Tende. Merxmiiller 
& Grau in 1962, M. Counted by Grau. 

M. tommasinii Marchesetti. 2n = 24. Yugoslavia, Istria, Osp. Merxmiiller & 
Wiedmann in 1960, M. Counted by Grau. 

Mucizonia hispida Berger. 2n = 16. Spain, Prov. Malaga, Grazalema. Merx- 
miiller & Grau in 1962, M. Counted by Grau. 

Omphalodes verna Moench. 2n = 48. Italy, Riviera, Alassio. Podlech in 1963, M. 
Counted by Grau. 

Onosma cinerascens Br. Bl. 2n = 28. Italy, Cottian Alps, Foreso. Merxmiiller & 
Grau in 1963, M. Couned by Grau. 

O. frutescens Lam. 2n = 14. Greece, Arkadia, Vitina. Rechinger in 1958, M. 
Couned by Grau. 

O. tridentinum Wettst. 2n = 28. Italy, Vintschgau, Kastelbel. Grau in 1963, M. 

Oxytropis amethystea Arv.-Touv. 2n= 16. France, Dauphiné, Col de Glaize. 
Gutermann in 1960, M. Counted by Damboldt. 

Palafoxia callosa (Nutt.) T. & G. n= 10. Texas, McCulloch Co. E. B. Smith 
232, KANU. Counted by E. B. Smith and R. R. Johnson, KANU. 

Panicum texanum Buckl. n = 27. Texas, Starr Co. F. W. Gould 8157, TAES. 

Paspalum distichum L. 2n = 20,,. California, Humboldt Co., Fish Lake. D. E. 
Anderson 2846, UC. 

Quincula lobata (Torr.) Raf. 2n = 11,,. Texas, Presidio Co. D. E. Anderson 
2574. UC: Texas, Val Verde Co. D. E. Anderson 2593, UC. 

Scleropoa rigida (L.) Griseb. n= 7. Texas, Walker Co. F. W. Gould 7519, 
TAES. This collection was reported earlier as Festuca octoflora Walt. (Gould, F. W. 
Am. Jour. Bot. 45:764. 1958). 

Sporobolus contractus Hitchc. n= 18. New Mexico, Eddy Co. F. W. Gould 
9541, TAES. 

S. virginicus (L.) Kunth. 2n = 10,,. Texas, Cameron Co. D. E. Anderson 2772, 
ee: 

Willkommia texana Hitchc. n = 30. Texas, San Patricio Co. F. W. Gould 8606, 
TAES. 


NOTES AND NEWS 


A NOTE ON THE TypE LOCALITY OF TETRACOCCUS ILICIFOLIUS.—The type locality 
of 7. itcifolius Cov. & Gilm. (Jour. Wash. Acad. 26:530. 1936), as stated in the 
original description, is in Fall Canyon on the west slope of the Grapevine Moun- 
tains, Death Valley, California, at 2000 ft elevation. The 15 plants located were 
“chiefly in crevices in the rock wall of the canyon which at this point consists 
chiefly of rhyolite,” and “the fewness of the plants and scarcity of fruit are evidence 
that the plant is in the process of extinction.”’ An isotype in the Death Valley Na- 
tional Monument herbarium is labeled in Gilman’s handwriting as having been col- 
lected at 4000 ft. On April 11, 1963 a search was made from 2000 ft to over 4000 ft 
elevation in Fall Canyon and the only plants of J. zlicifolius encountered were at 
3200 ft about four miles above the mouth of the canyon in crevices in the limestone 
wall. Estimates were made of the size of the plants: two had crown diameters of 
more than 4 ft, eight had crown diameters of 2—4 ft, and nine had crown diameters of 
less than 2 ft, a total of 19 plants. Judging from the distribution of shrub size, the 
stand appears to be maintaining itself, although no seedlings were observed and 
there was little evidence of fruiting. The vigor of the plants was comparable to that 
of individuals in a stand of several hundred shrubs in the Panamint Mountains. 
It is concluded that the stand at 3200 ft in Fall Canyon is the type locality. — 
H. Tuomas HArvey, Department of Biological Sciences, San Jose State College. 


THE HORDEUM JUBATUM—CAESPITOSUM— 
BRACHYANTHERUM COMPLEX IN ALASKA 


W. W. MITCHELL AND A. C. WILTON 
INTRODUCTION 


The wild barleys Hordeum jubatum L. and H. brachyantherum Nevski 
have, until recently, been considered separate species. The two are dis- 
tinguished principally on the basis of awn length, the former being long- 
awned and the latter short-awned (figs. 2 and 4). Plants intermediate in 
awn length (fig. 3) have been classified variously: i.e., 1. as the species 
H. caespitosum Scribn. (Rydberg, 1922; 1932; Anderson, 1959); 2. as 
H. jubatum var. caespitosum (Scribn.) Hitchc. (Hultén, 1941-1950; 
Covas, 1949; Hitchcock, 1950; and others); and 3. as a subspecies of 
H. jubatum (discussed below). 

The intermediate taxon has been reported over a wide area. Covas, in 
his treatment of the American species of Hordeum, assigned a number of 
specimens originating from Mexico northward to Washington and Mon- 
tana to H. jubatum var. caespitosum and postulated that they were the 
result of hybridization between H. jubatum and H. brachyantherum. 
Love (1959) listed H. jubatum var. caespitosum as a member of an ele- 
ment that has migrated into Manitoba from the West. Hultén (1962) 
represented the approximate, total range of the taxon as extending from 
northern Mexico through the western half of the United States and Can- 
ada to the coastal regions of southern and south-central Alaska. 

Extensive biosystematic work performed recently in Canada has firmly 
established the hybrid status of the intermediate taxon and has cast some 
doubt on the separation of the parental taxa. Rajhathy and Morrison 
(1959; 1961), from studies of karyotype, pairing behavior, and inter- 
fertility, reported that H. jubatum and H. brachyantherum were con- 
specific. Improved pairing in the F, was said to indicate “a tendency for 
true breeding and stabilization in early generations” (1959). 

More recently Bowden (1962) assigned all three taxa to H. jubatum 
(all n=14), conferring subspecific rank upon each as follows: H. juba- 
tum L. emend. subsp. jubatum, H. jubatum L. emend. subsp. breviaris- 
tatum Bowden (to replace H. brachyantherum), and H. jubatum L. 
emend. subsp.  intermedium Bowden (to replace var. caespitosum). 
For the purposes of this paper,’ however, the nomenclature of Hitchcock 
will be followed. Bowden reported hybrid swarms in the Canadian prairies 
that demonstrated clinal variation with the parental taxa and also re- 
ported the presence of advanced segregate-populations in North and 
South Dakota. 


1 The authors express their gratitude to R.W. Pohl for his review of the manu- 
script. This study was supported in part by The Rockefeller Foundation under 
Grant RF-61036. 


270 MADRONO [Vol. 17 


woe in 7 ¢. pen U bes a + 
j : wees \ 
1 OR ww 
ea \ \1 ¢ ) ry x 
4 Vint ; 
es E fast) ) of 
i _ ENG bas \ 
CLidhain f } Ves ois ; % 
apt Ae oF ’ rs ¥ 
ne (as ; e . n \ 
oe j a 9% $s a. a 
ne « ee mas a 
¢ ,f/ ° : we ®, of 
aU pe w . & 
Ss vy, ert . c » ; i x ’ 
a . -yfewg - x Seo henal a . : R 
4 NN im PEG) Geen fa ne Ee ricen Meera ; 
7 SEMAN . Fal = 1 \. ie ‘ 
ems OL é : ‘ 
oc. i * : 2 oa 
t. f 2 9 - 
~ . x nis F bea 
a) . : Se 
Soe Wy hoo © esi 4 SAS ee 
oy rad f si fy a ae He 
a: Fa : \ ak Ss 
o Be Sa \Y 
\ : i ap = } i Chichagof lI. 
“ ney ~ ff i s eee : 
? 4 pie paranct : ¢ gy \\ 
nie ie n 
-o . ef ! " tae o 
a © oe 
few? O Collections of H. yubatum 
A : Lalor ee Ua. 
SE SE de A Collections of H. brachyantherum 
: ae € me =e 
ear ~ t ‘ . 
ae ad @& Study area with both parent 
Pe ad and hybrid populations 


Fic. 1. Collection sites of H. jubatum, H. jubatum var. caespitosum, and H. bra- 
chyantherum. 


The present report concerns the status of the three taxa in their north- 
ernmost region of occurrence in the West. The principal study area was 
in the vicinity of Palmer, Alaska, where hybrid swarms were found oc- 
curring with the parental taxa. But in most cases where jubatum and 
brachyantherum were found together, hybrids were not apparent. In 
view of the disposition by Bowden more detailed analyses of these popu- 
lations were deemed pertinent. 


SOURCE OF MATERIALS 


The study was confined to Alaskan materials. Population samples of 
hybrid and parental types were collected around the Knik Arm of Cook 
Inlet northeast of Anchorage in 1962 and 1963. Other collections of the 
parental taxa were made in regions where no contact between the two 
was evident (fig. 1). The latter collections of brachvantherum were made 
in the region of Homer on the Kenai Peninsula, on Kodiak Island, and 
at Juneau, Chichagof Island, and Baranof Island in southeastern Alaska. 
Samples of jubatum were collected along the highway system of the Alas- 
kan interior: at Atlasta House Lodge, Mile 166, Glenn Highway; Tok 
Junction, Mile 1314, Alaska Highway; Fairbanks; Paxson Lodge, Mile 
186, Richardson Highway; and at Kenny Lake and Copper Center in 
the Copper River Valley. 


1964] MITCHELL & WILTON: HORDEUM 271 


Spikes for cytological analysis were collected from plants of caespito- 
sum and from adjacent plants of jubatum and brachvantherum growing 
at one of the Cook Inlet sites. 


DISTRIBUTION AND ECOLOGY 


The parental taxa differ markedly in their ranges of distribution in 
Alaska, although overlapping in a number of areas. The long-awned 
jubatum is widely distributed and well represented in the interior. Gen- 
erally it is less frequent to rare or absent in the coastal regions. Appar- 
ently it does not occur in the arctic region north of the Brooks Range. 
Though essentially weedy in character, occurring in abundance along 
roads and around settlements, as remarked by Hultén (1942, p. 267) 
‘“‘(it) must certainly be indigenous in the interior (of Alaska).” Hultén 
(1962) considers it native both to North America and northeastern Asia. 
Farrand (1961) reported that the remains of jwbataum were found in 
the analysis of stomach contents of a frozen woolly mammoth discovered 
in northern Siberia. 

The short-awned brachyvantherum is indigenous to North America and 
some coastal portions of eastern Asia (Hultén, 1941-1950). It is strictly 
coastal in its Alaskan occurrences, being found inland only along estu- 
aries and inlets. Unlike jubatum it is an important component of certain 
native plant communities that develop on beach meadows and tidewater 
flats (Hanson, 1951). It also behaves as a weed by invading disturbed 
eround around settlements. 

The hybrid caespitosum may be expected to occur wherever the par- 
ental taxa meet. We observed it in abundance only in the Cook Inlet 
region. 

The parental taxa demonstrate a distinctive difference in ecology where 
they meet on beach meadowlands. For instance, jubatum occurs on dis- 
turbed ground of a picnic area intruding upon meadowland of the Eklutna 
flats northeast of Anchorage. It is not present, however, in the native 
meadow community that is immediately adjacent and wherein brachy- 
antherum is an important component. Hanson (1951) described this com- 
munity in some detail. Although there is ample opportunity for pollen 
exchange between the two taxa in this area, only one small colony of 
hybrid plants was located. These occurred in the vicinity of an old aban- 
doned dwelling. 

The most extensive hybrid swarms were found 1. on a homestead that 
was established in the middle 1930’s (during the early days of agricul- 
tural colonization of the Matanuska Valley), and 2. within the town of 
Wasilla, which was founded about 1916. Several collections also were 
made in Palmer, the agricultural center of the valley and Matanuska vil- 
lage, a nearly abandoned settlement. 

Hyrbidization appears to have occurred mostly where both parental 
taxa have been in contact on disturbed ground for a number of years. 
Indications are that the hybrids are short-lived, so hybridization may 


272 MADRONO [Vol. 17 


be more frequent than is apparent. Some hybrid colonies located in 1962 
were not present in 1963, apparently having succumbed during the 
winter. 


MorRPHOLOGICAL ANALYSIS 


In the morphological analysis of the three taxa a node bearing its 
single sessil spikelet and two pediceled lateral spikelets was chosen be- 
tween one-fourth and one-half the distance up the inflorescence. Often, 
due to disarticulation of the rachis, only the basal half or less of the in- 
florescence remained, particularly on brachyantherum. The following 
characters of this group of spikelets were measured or scored: 1. lemma- 
awn length of the single floret in the central spikelet, always measured 
from a point on the lemma opposite the tip of the palea, a procedure which 
ensured consistency of technique but which probably exaggerated the 
actual length of the awn, 2. palea length of the floret in the central spike- 
let, 3. internode length of rachis joint attached to spikelet, 4. develop- 
ment of a staminate floret in at least one of the two lateral spikelets, with 
a well developed palea being accepted as evidence of its development. 
Other characters measured or scored were: 5. width of penultimate leaf, 
6. presence or absence of hairs on upper surface of leaf, 7. depth of cleft 
on culm at juncture of peduncle and inflorescence—the cleft resulting 
from a collar that forms an open-necked V of varying depths on one side 
of the culm at the origin of the inflorescence (fig. 8), 8. features of the 
lower epidermis of the terminal leaf. 

In the analysis of the epidermal features, the three taxa were found 
to differ principally in the shape and frequency of prickles. The prickles 
of jubatum were long-barbed and very numerous (fig. 5); those of brachy- 
antherum were short-barbed and infrequent (fig. 7); and those of caespi- 
tosum were intermediate in shape and frequency (fig. 6). While critical 
measurements were not made of these characters, observations of a num- 
ber of leaves indicated that brachyantherum could be more readily dis- 
tinguished by this means than could jubatum or caespitosum, since the 
latter two tended to converge in their epidermal features. 

Quantitative measurements of spikelets and leaves are summarized in 
Fig. 8. The delineation of the taxa according to awn length is particu- 
larly noteworthy in this summary. Short-awned brachyantherum is clear- 
ly separated from the other two taxa, and all three populations are widely 
disjunct in their major portions (1.e., the portion contained in two stand- 
ard deviations—theoretically, about 68%). In the other measurements 
the means differ but the ranges of variability overlap for all three taxa. 
The portions contained in two standard deviations generally overlap, 
with the most disjunct patterns being registered in leaf width and palea 
length. The hybrid caespitosum essentially is intermediate in its ranges 
and means for all measurements. 

Figure 8 indicates, however, that the extent of overlap in characters 
is no greater where the parents come in contact and hybridize than it is 


1964] MITCHELL & WILTON: HORDEUM 2.3 


for the population as a whole. Indeed, in the cases of palea length and 
internodal length the portions of the collections from mixed populations 
included in two standard deviations are even more widely separated than 
comparable portions of the total parental populations. 

The three taxa were further delimited by plotting awn length against 
palea length (fig. 9) and by determining correlation and regression co- 
efficients of the paired data (table 1). A high positive correlation between 
awn length and palea length in brachyantherum, within a very narrow 
range of variation in awn length and a broad range of variation in palea 


Fics. 2-7. Inflorescence (14 x) and leaf epidermis (125 x): 2, 5, H. jubatum; 
3, 6, H. jubatum var. caespitosum; 4, 7, H. brachyantherum. 


274 MADRONO [Vol. 17 


TABLE 1. CORRELATION AND REGRESSION COEFFICIENTS OF THE PAIRED VARIABLES 
LEMMA-AWN LENGTH AND PALEA LENGTH FOR HORDEUM JUBATUM, 
H. JUBATUM VAR. CAESPITOSUM, AND H. BRACHYANTHERUM 


Number of specimens Correlation Regression 

Taxon included in analysis coefficient coefficient 
H. jubatum 184 + .22 0.09 
H. jubatum var. caespitosum 69 + .32 0.39 
H. brachyantherum 210 + .62 atid 


length, distinguishes it in particular from the other two taxa. These have 
much lower correlation coefficients within much wider ranges of variation 
in awn length. 

An analysis of the combined data, i.e., treating the data as that of a 
single taxon, indicated an entirely different relationship between the two 
variables. Whereas the correlation coefficients were positive when treating 
the taxa separately, the combined data yielded a negative correlation 
with a high value of —.76 and a regression coefficient of .40. 

Differences in leaf pubescence contributed to the separation of the 
shorter-awned hybrid forms from brachyantherum. In Figure 9 all but 
one of the hybrid specimens nearest brachyantherum are pubescent on 
the upper leaf surface, as opposed to the consistently glabrous condition 
in brachyantherum. Both jubatum and caespitosum varied in this regard. 

Figure 9 also demonstrates that all three taxa varied in the develop- 
ment of staminate or sterile lateral spikelets and that plants with longer 
paleas tended to produce the staminate lateral spikelets. The correlation 
was much more pronounced in brachyantherum than it was in jubatum,; 
but because of the difference in magnitude of palea lengths of the two 
taxa, where the two overlapped in palea lengths they often differed in the 
character of their lateral spikelets (table 2). Plants of jubatum with 
paleas over 6.5 mm up to the maximum of 7.5 mm tended to produce 
staminate lateral spikelets, while plants of brachyantherum with paleas 
measuring in the same range generally produced sterile lateral spikelets. 
Palea measurements ranged up to 10.2 mm in brachyantherum. 


TABLE 2. PRODUCTION STAMINATE LATERAL SPIKELETS IN SPECIMENS OF HORDEUM 
JUBATUM, H. BRACHYANTHERUM, AND H. JUBATUM VAR. CAESPITOSUM 
WITH PALEAS MEASURING FROM 6.6 mm TO 7.5 mm 


Number Number 

of specimens with of specimens with 

Taxon sterile spikelets staminate spikelets 
H. jubatum 8 12 = 60% 
H. brachyantherum 48 a= OG 


H. jubatum var. caespitosum 18 16 = 47% 


1964] MITCHELL & WILTON: HORDEUM 215 


CYTOLOGICAL ANALYSIS 


Meiotic studies were conducted of floret material fixed in acetic alco- 
hol (3:1), smeared in acetocarmine, and the correct stages made perma- 
nent with Venetian turpentine (Wilson ,1945). Meiosis was found to be 
highly irregular in the hybrid (figs. 10-13). Because of this it was im- 
possible to obtain detailed pairing relationships between the parental 
genomes. Metaphase I of the hybrid was particularly disturbed; among 
120 cells examined there were none in which chromosome pairing ratios 
could be ascertained. Practically all were of the types illustrated. Multi- 
valent associations usually appeared (figs. 10 and 11), but neither the 
number of chromosomes in the association nor their frequency could be 
definitely determined. Further, it is not certain whether these associa- 
tions were always due to true pairing or to chromosome stickiness, as 
suggested by several investigators of interspecific and conspecific hy- 
brids in the genus Bromus (Walters, 1954; Nielsen, 1955; Hanna, 1961). 

Anaphase I showed some laggards that often seemed to be in multi- 
valent associations (fig. 13). These laggards were manifest as micro- 
nuclei at second divisions and in the quartet stage (fig. 12). The unequal 
timing of second division also is common in cells of the hybrid and may 
be a further manifestation of abnormal meiosis. 

Meiosis was mostly regular in jubatum and brachyantherum (fig. 14). 
The frequency of micronuclei, based on observations of 200 cells for each 
species, was 0.4% in jubatum and 0.5% in brachyantherum, compared 
to 13.1% in caespitosum. The meiotic irregularities of the hybrid were 
correlated with a high degree of sterility, as determined by counts of 
mature caryopses in the central spikelets (table 3). One hybrid plant 
was found to be quite fertile. The parental taxa were highly fertile. 


DISCUSSION 


We believe that the data presented here offer little support for com- 
bining the Alaskan populations of H. brachyantherum with H. jubatum. 
This study reveals 1. marked differences in ecology of the two taxa, 2. an 
apparent failure of the two taxa to hybridize on many sites where they 
are in contact, 3. meiotic irregularities and a high degree of sterility in 
the hybrid intermediate caespitosum, 4. a distinct morphological separa- 
tion of brachyantherum from both caespitosum and jubatum on the basis 
of awn length, 5. an association of other morphological differences with 
this difference in awn length, and 6. differences in relationships between 
characteristics that would appear to be based on fundamental differences 
in genetic systems. 

If brachyantherum and jubatum are to be considered members of the 
same species, then jubatum would be the long-awned phase and brachy- 
antherum the short-awned phase of a mutual genetic system. Accord- 
ingly, if awn length is plotted against another characteristic with which 
it shows correlation, we might expect that variation within one phase 


276 MADRONO [Vol. 17 


would be a natural extension of variation within the other phase. But 
when awn length was plotted against palea length, the specimens formed 


j- (189) NS cers cree te 


(92) 5 eee = eee 


c-(68) =e Awn Length (cm) 


» (84) che 
(209) 


ase ee 
j-(18 9) 


(92) ~ Uys 7 


Palea Length 
c-(68) an HH 


(84) yess) Se 


b-(209) a ee Folge aay 
,_ (188) Pe coos cox: Sees 


(92) pe oo Leof Width 


c-(64) SES - ie 


» 82) sie I ee oe ee ee 
(204) eS seers Om re a 


j_(189) _th_ 
(92) 1p- 


Internode Length on 
c-(686) Hh} 

Inflorescence (mm) 
»/84) th 
(209) “oO 


; W189) (cc co ee ee * 


Cleft on 
o-(68) a Peduncle (mm) 


O l 2 3- 4 Do 36 t 8 Se OR 


Fic. 8. Summary of morphological analysis of H. jubatum, H. jubatum var. 
caespitosum, and H. brachyantherum. Small letters: j = jubatum, c = caespitosum, 
b = brachyantherum. Figures in ( ) signify number of specimens analyzed for each 
character. Horizontal line encompasses total range of variability; vertical line denotes 
mean; blocked portion comprises two standard deviations. Solid line represents total 
sample analyzed for each character; dashed line includes only that portion of each 
parental sample obtained from mixed populations. 


1964] MITCHELL & WILTON: HORDEUM if 


@ Lateral Floret Staminate 
O Lateral Floret Sterile 


re) Leaves Pubescent 
O Leaves Glabrous 


-> Regression Line 


z= 9 
= ony 
H >} 
ao 
8 
OB 6 cach a 
vA rere _‘é 
a) : eo 
OSS 4 \ 
Pe | 4 entre ae eee \ 
7 t te Jo oe ‘ 
< 6 $° ° ae een oe 
o Sue 43 ne bo,” 5 48 2° paees 
“ (88 te gis pray Gesu o a ee oeieys 2 eet 
< & acs se”, lags 8, can 
A, 6 
. poets ° dot, 208 Based omc dus ave 4! 
oN $ Sauce of S83 °6 O- = 
ar as ies 
5 a ,° ae 
‘bo? 


LEMMA-AWN LENGTH (CM) 


Fic. 9. Graphical comparison of H. jubatum (long-awned), H. jubatum var. 
caespitosum (intermediate), and H. brachyantherum (short-awned). Dashed lines 
enclose all variants of parental taxa collected in regions where the two were not in 
contact. See Fig. 8. for numbers of specimens analyzed. 


three natural groupings with regression lines that could not possibly be 
interconnected. An attempt to analyze the data as that of a single taxon 
produced a result that clearly violated obvious character relationships. 
The high correlation between awn length and palea length in brachy- 
antherum, as opposed to a weak correlation in jubatum, and the very 
different regression coefficients certainly must reflect significantly differ- 
ent genetic systems. 

Difficulties of separation that did occur generally involved specimens 
resembling jubatum, whereas the brachyantherum component of mixed 
populations appeared quite distinct. Whatever introgression has occurred 
has been insufficient, at least at this stage, to have produced intergrada- 
tion. The propensity for hybridization between jubatum and brachy- 
antherum, moreover, is insufficient in itself as a criterion for lumping the 
two. For instance, H. jubatum appears to hybridize quite as readily with 
Agropyron trachycaulum (Link) Malte as it does with H. brachy- 
antherum in Alaska. 

Most specimens of caespitosum analyzed for fertility were completely 
infertile. The representative found to be the most fertile was a short- 
awned variant (labeled f in fig. 9) nearer to brachyantherum than to 
jubatum. The structure of the scatter diagram shows considerable room 
for the development of specimens with awns longer than in brachyan- 


278 MADRONO [Vol. 17 


therum and with paleas longer than in jubatum. The self-compatibility 
of these taxa enhances the possibility of a new species being formed in 
this morphological range, should appropriate reproductive barriers de- 
velop (Grant, 1963, pp. 451 ff.). This assumes, of course, that these char- 
acters reflect more than morphological differences. Biotic and physical 
factors of the environment, however, could preclude such a development. 


TABLE 3. PERCENT FERTILITY OF HORDEUM JUBATUM, H. JUBATUM VAR. 
CAESPITOSUM, AND H. BRACHYANTHERUM 


No. of % No. of Range in % 
Taxon Florets Fertility Plants Fertility Median 
A. jubatum 666 87.1 10 18.6-100* 89.2 
H. jubatum var. 
caes pitosum 1054 3.0 15 0.0-47.07 0.0 
H. brachyantherum 543 89.7 13 79.1-100 89.7 


1QOne plant of jubatum registered 18.6% seed set, otherwise the range was be- 
tween 86.7% and 100%. 

2 One plant of caespitosum registered 47% seed set, otherwise the range was be- 
tween 0.0% and 2.0%. 


Fics. 10-14. Meiotic configurations in Hordeum: 10-13, meiotic irregularities in 
H. jubatum var. caespitosum; 10, 11, Metaphase I; 12, second division; 13, Ana- 
phase II; 14, normal Telophase II in H. jubatum. 


1964] MITCHELL & WILTON: HORDEUM 279 


Apparently the intermediate taxon has achieved a measure of fertility 
in the Midwest and is able to migrate on its own. With the potential doubt- 
lessly varying from region to region throughout the extensive range of 
the hybrid, the possibility of its establishment as a species in some por- 
tion of its range seems worthy of more serious consideration. The out- 
come of this development in Alaska would appear as yet to be in doubt, 
but its present status does not warrant combining brachyantherum with 
jubatum for theoretical or practical reasons. 

In view of the above we recommend that the hybrid be referred to as 
Hordeum X caespitosum Scribn. (pro sp.). The descriptions of Bowden 
(1962) for distinguishing his three subspecies apply, with the following 
modifications on measurements of the floret of the central spikelet: 

H. brachyantherum ...lemma-awn 0.6—1.5 mm, palea 6.4—9.5—(10.2) mm 


H. X caespitosum ...lemma-awn 1.5—3.5 mm, palea (5.7)—6.5—9.0 mm 
A. jubatum ...lemma-awn (2.6)—3.5—7.5—(9.4) mm, palea 4.8—7.0—(7.5) 
mm 
SUMMARY 


Alaskan populations of Hordeum jubatum, H. brachyantherum, and 
their hybrid H. jubatum var. caespitosum were sampled and analyzed 
morphologically and cytologically. Irregular meiotic behavior in the hy- 
brid was coupled with a high degree of sterility, while the parental taxa 
were highly regular in meiosis and in most cases highly fertile. Differ- 
ences in ecology, morphology, and certain character relationships were 
cited as evidence favoring the continued separation of H. brachyanthe- 
rum and H. jubatum, in contraposition to a recent action combining the 
two. The possibility of the development of a new species in some por- 
tion of the extensive range of the hybrid was proposed. It was recom- 
mended that the hybrid be designated as Hordeum  caespitosum 
Scribn. (pro sp.). 


Agronomy Department, Alaska Agricultural Experiment Station, Palmer, 
cooperating with the United States Department of Agriculture, Uni- 
versity of Alaska 


LITERATURE CITED 


ANDERSON, J. P. 1959. Flora of Alaska and adjacent parts of Canada. Iowa State 
Univ. Press, Ames. 

BowpEn, W. M. 1962. Cytotaxonomy of the native and adventive species of Hor- 
deum, Eremopyrum, Secale, Sitanion, and Triticum in Canada. Canad. Jour. 
Bot. 40:1675-1711. 

Covas, G. 1949. Taxonomic observations on the North American species of Hor- 
deum. Madrono 10:1-21. 

FARRAND, W. R. 1961. Frozen mammoths and modern geology. Science 133:729-735. 

Grant, V. 1963. The origin of adaptations. Columbia Univ. Press, New York. 

Hanna, M. R. 1961. Cytological studies in Bromus species, section Bromopsis. 
Canad. Jour. Bot. 39:757-773. 

Hanson, H. C. 1951. Characteristics of some grassland, marsh, and other plant com- 
munities in western Alaska. Ecol. Monogr. 21:317-378. 


280 MADRONO [Vol. 17 


Hitcucock, A. S. 1950. Manual of the grasses of the United States. Ed. 2, revised 
by Agnes Chase. U.S. Dept. Agr. Misc. Publ. 200. 

Hu tten, E. 1941-1950. Flora of Alaska and Yukon. Lunds Univ. Arssk. 

. 1962. The circumpolar plants. I. Vascular cryptogams, conifers, mono- 
cotyledons. Kungl. Svenska Vetenskapsakademiens Handl. Pjarde Ser. 8:1-275. 

Love, D. 1959. The postglacial development of the flora of Manitoba: A discussion. 
Canad. Jour. Bot. 37:547-585. 

NiEtsen, E. L. 1955. Cytological disturbances influencing fertility in Bromus inermis. 
Bot. Gaz. 116:293-305. 

RaAjyHATuy, T. and J. W. Morrison. 1959. Cytogenetic studies in the genus Hordeum. 
IV. Hybrids of .H. jubatum, H. brachyantherum, H. vulgare, and a hexaploid 
Hordeum sp. Canad. Jour. Genet. Cytol. 1:124-132. 

1961. Cytogenetic studies in the genus Hordeum. V. H. jubatum and the 
New World species. Canad. Jour. Genet. Cytol. 3:378-390. 
RyvpsBerc, P. A. 1922. Flora of the Rocky Mountains and adjacent plains. New York. 
. 1932. Flora of the prairies and plains of central North America. New York 
Botanical Garden. 

Watters, M.S. 1954. A study of pseudobivalents in meiosis of two interspecific 
hybrids of Bromus. Am. Jour. Bot. 41:160-171. 

Witson, G. B. 1945. The Venetian turpentine mounting medium. Stain Tech. 20: 
133-135. 


NOTES AND NEWS 


CONDALIA MEXICANA SCHLECHT. VAR. PETALIFERA M. C. JOHNSTON, VAR. NOV.— 
Varieties a typica differens floribus petala parva caduca habetibus. Holotype: Mexico, 
Zacatecas, rd to Huejuquilla el Alto, Jalisco, 1 mi w of rd junction 18 mi s of 
Valparaiso on rd to Mezquitic, Jalisco, near Lat 22° 38’ N, Long 103° 48’ W, 
McVaugh 17675, Sept. 4-5, 1958 (MICH). This variety is treelike, 6 m tall with a 
trunk diameter of 20 cm. The fruit is reddish; the flowers are greenish. Only one 
tree was seen at the type locality. It grew near the summit of a pass at 2100 m in 
rocky oak-covered mountains. The discovery of this plant supports my recent 
deemphasis of the significance of the presence or absence of petals in this genus 
(Brittonia 14:332-368. 1962). It is extremely closely related to C. mexicana Schlecht. 
of the eastern Sierra (Tamaulipas to Guanajuato and Hidalgo, in southern Puebla, 
and in northern Oaxaca), but its flowers possess minute petals. Its locality, to the 
west of the range of C. mexicana var. mexicana and at a much higher elevation, 
supports its designation as a variety. It also differs from C. mexicana var. mexicana 
in the dark olivaceous color of the dried specimen and in the arborescent stature. 
A collection from Durango (Palmer 608, F, GH, UC, US) may also be referable 
to C. mexicana var. petalifera. This new variety emphasizes the similarities of C. 
mexicana to the rarely collected petaliferous populations near Guanajuato which 
have been called C. velutina I. M. Johnst. The Zacatecas plant, however, has the 
narrow bud scales, small fruits, and sparse pubescence of C. mexicana. — MARSHALL 
C. Jounston, Plant Research Institute and Department of Botany, University of 
Texas, Austin. 


THE GENUS ESCHSCHOLZIA IN THE SOUTH COAST 
RANGES OF CALIFORNIA 


WALLACE R. ERNST 


Six species of Eschscholzia occur in the South Coast Ranges, a moun- 
tainous complex extending southeast from San Francisco Bay to the 
Transverse Ranges that begin in Santa Barbara and Ventura counties 
(Munz, 1959). Three virtually are restricted to this region and three have 
wide range of distribution extending beyond the boundary of the state. 
The primary purpose of this paper is to clarify the identity of E. hype- 
coides. The other species are discussed briefly from the narrow viewpoint 
of their occurrence in this region. I am grateful to those who have assisted 
me in the preparation of this paper, among them Sir George Taylor, 
Director of the Royal Botanical Gardens, Kew, for the privilege of bor- 
rowing three types critical to this discussion, and also M. A. Canoso, 
Emily Reed, Velva E. Rudd, Isabelle Tavares, John H. Thomas, and 
Ernest C. Twisselmann. 


Eschscholzia hypecoides was one of four species of the genus described 
by Bentham (1834) from plants collected in California by David Doug- 
las. This one was distinguished by its branched leafy stems and resem- 
blance to Hypecoum grandiflorum Benth., a fumariaceous herb of the 
Mediterranean region. Douglas’s records of the gathering of the plants 
were lost but his itinerary in California probably was confined to the 
Coast Ranges (Jepson, 1933; McKelvey, 1955). 


Plants of Eschscholzia with branched leafy shoots are common in the 
Coast Ranges but the species usually attributed to this region normally 
bear no very close resemblance to Hypecoum, the genus to which Ben- 
tham alluded in naming EF. hypecoides. It seemed possible to me that 
some peculiar aspects common to both were not covered in the formal 
description of E. hypecoides and that these might be of assistance in 
rediscovering E. hypecoides since it is not recognized in the region from 
which described. 


To test this hypothesis I examined the specimens of Eschcscholzia in 
20 herbaria (A, ARIZ, CAS, DS, GH, LA, MO, ND, NY, OBI, ORE, 
OSC, POM, RSA, SBBG, SBC, SBM, SD, UC, US). The distribution of 
those which, by stretch of imagination, might resemble Hypecoum is 
plotted on the map (fig. 1). Almost all, bearing collective resemblance to 
one another and restricted in origin to the South Coast Ranges, were 
found in the covers with EL. lemmoni. These specimens clearly are con- 
specific with the holotype of E. kypecoides (fig. 2). In the modern liter- 
ature, however, they can be identified only as EL. lemmoni and the name 
E. hypecotdes is found in synonomy only under E. caespitosa (or as E. 
caespitosa var. hypecoides). 


Several characteristics of E. hypecoides are notable. The cotyledons 
are linear spatulate and undivided. The plants are annual, usually con- 


282 MADRONO [Vol. 17 


E 3 ¥ 0; 
: Be . ALAMEDA e 


ESCHSCHOLZIA 


e HYPECOIDES 


o LEMMONII 


* MINUTIFLORA 


o RHOMBIPETALA 


Fic. 1. South Coast Ranges region of California showing counties and the dis- 
tribution of four species of Eschscholzia. 


1964] ERNST: ESCHSCHOLZIA 285 


spicuously glaucous and pubescent with short, unicellular hairs. The 
amount of pubescence, variable from plant to plant in the same colony, 
sometimes nearly is uniform, or confined to the sepals, the petioles, or the 
fruits; occasionally the plants are glabrate. 

The earliest few flowers often are borne on rather long, sometimes 
leafless peduncles originating from the center of the caespitose plants. In 
all but the most depauperate specimens, there usually are several flowers 
(or young fruits) borne on relatively short pedicels inserted on determi- 
nate inflorescence shoots. The leaves on these shoots frequently are oppo- 
site on short petioles and the immature buds, at first clustered and almost 
always nodding, appear sessile among the small leaves. 

The petals usually are bright yellow and frequently show a diffuse 
orange spot near the base; in a few instances they are orange through- 
out. The receptacle, relatively to the corolla, is characteristically small 
(table 1). Flowering plants have been collected from early March through 
June on clay, diatomaceous earth, granite, gravel, serpentine, and white 
shale at altitudes of 700 (rarely 200) to 3900 ft. The habitat usually is 
open slopes or disturbed areas near woodland or chaparral associations 
from western Stanislaus Co. in the north to Santa Barbara Co. in the 
south (fig. 1). The gametic chromosome number is six (Ernst, 1959) ; 
reconfirmed in Ernst 539, 538, 762, DS) and the pollen have five or six 
colpae. 

The specimens from Monterey (especially in the Arroyo Seco) and 
San Benito counties most nearly match the holotype. The plants often 
occur on outcrops of white strata (the clay, diatomaceous earth, and 
white shale noted above) which seem sparsely colonized by other vege- 
tation. The frequency of pubescence, the branching habit often with 
opposite leaves, the nodding clustered buds, the small size of the recep- 
tacles, and the frequent association with the white strata of the South 
Coast Ranges distinguish E. hypecoides. 

At three localities I observed EF. Aypecoides and E. californica within 
a few feet. The populations of LE. Aypecoides, confined to the white strata 
in two instances, seemed normal. The nearby colonies of EL. californica 
consisted of an inordinate number of depauperate plants and became 
typical (in stature of plants and prominence of receptacle rims) at some 
distance from the E. hypecoides and away from the white strata. If these 
species are hybridizing, it would seem that only the morphology of E. cali- 
fornica is being affected. 

Only a few problems should arise in identifying EF. hypecoides. In 
western Stanislaus Co., the range coincides with that of E. caespitosa, 
and E. rhombipetala (as well as E. californica) and a few sheets seem 
difficult to determine. Some specimens collected between Coalinga and 
Parkfield, Fresno-Monterey county line (K. Brandegee s.n., 7 sheets, 
UC), almost are indistinguishable from E. minutiflora but my observa- 
tions in the field (and of the pollen) convince me that these are E. 
hy pecoides. 


~~ 
re 
= 
© 
- 
bed 


MADRONO 


284 


Loez(9r1) Ss] 


ACHAEA ACA 


GCC CCX L009) S¢ 


cx 


Z(LT)T X €(€°7)Z 


(ACS G SGA 


(On) syueld p-T 


DIDIJAGIQueoyA “YT 


[00€ ($'16Z)0LT] 


p(S7)z xX S(S'e)Z 


G(8iZ) 2 << 1869) ce 


SZ(OL)OL X SE (Fr 61T)OI 


OCC) Eee c) Sel 


9(9°S)b X 8SI(LIT)Z 


(Sq) sqzurfd TI-9 


DSO4UdSavno “FT 


loge (Ss STZ)06] 


S(r'r)€ X OL(T'8)9 


(8°Z)Z X O8(¢L£)S9 


6Z(T'EZ)ST X ZE(7'SZ)ST 


COS) x 6 CSS) 


O1CS:9)# X TZCUST) TT 


(DN) squetd ¢7-T1 


nNuUouULM ay “Y 


ALITIGVIaVA JO AONVY MOHS 
OL GALOATAS VIZIOHOSHOSY JO SNAWIONdS aassaug JO (SASTHL 
-NIUVd NI AOVAGAY) “WI NI SLNAWAXASVAYT HIGIM, X HLONAY ATAWVS ‘T Alavy 


[OOr (8° TTZ) $6] 


COL IE eX SCE 2) 2 


Z(ST)T X L9(S'Sh) LZ 


ST(T'OL) X OL (6TT)S 


CODECS pI 


OCS HS Si sys 


(DN) syueld 77-81 


Saplorad KY “F 


sjueld 
Jo 431I9H 


sajoe}das01 
ySa31e'T 


S}INay 
}SaB1e'T 


s[eyod 
dInjely 


stsoyjue 4 
appe}das0y 


spnq oainjeu 
JO xA[eZ 


The range of E. hypecoides overlaps that of E. lemmonu and some- 
times both occur on the same or nearby slopes. In these instances the 


petals of EL. hypecoides sometimes are larger and darker in color but the 


receptacles remain characteristically small. The striking degree of simi- 


1°64] ERNST: ESCHSCHOLZIA 285 


larity between these species is emphasized by the fact that plants unmis- 
takably equivalent to Bentham’s holotype of E. Avpecoides can be iden- 
tified in contemporary floras and manuals only as EL. lemmoni. The 
confusion surrounding the identify of EL. kypecoides will be traced briefly. 


7 . es : 
bikes? 2 pene, et tay 
Sey 

Vite Str § OI fh ok O 


t 


Uasiguches PSRs. Se 


3 8. Ries Sats pa 
\ ; PPR RE GES 
; fo ‘ S 
combines t MPM 5% Las gD 
% pes ; ie : 


REN OPE MOR SER: 


Bi BE: 


hGH ti is: 


Fic. 2. Holotype of E. hypecoides Benth. (K), collected in California by David 
Douglas. 


286 MADRONO [Vol. 17 


The first of Bentham’s four species of Eschscholzia, E. crocea, was 
published with a plate which showed this to be a form of E. californica. 
Next, and not illustrated, were E. caespitosa, E. tenuifolia, and E. hype- 
coides. These names are preceded chronologically only by E. californica 
Cham., the type species of the genus. The holotypes of E. caespitosa and 
E. tenuifolia (fig. 3) are rather young specimens that probably were 
collected before the first seeds matured. They cannot be matched easily 
with modern collections. Gray, after inspecting these holotypes reduced 
E. tenuifolia to synonomy under E. caespitosa and originated the com- 
bination E. caespitosa var. hypecoides (Benth.) Gray, a trinomial ap- 
pearing in many treatments of the California flora since 1887. The most 
recent of these, and one reflecting standard practice among California 
botanists, is that of Munz (1959), where it is said that “The typical 
form of the sp[ecies, i.e., E. caespitosa| has scapose stems, those with 
leafy stems constitute the var. hypecotdes, which is the more common 
form and occurs throughout the range.” 

In some instances, branching which gives rise to leafy inflorescence 
shoots may depend on the location or the degree of maturation of the 
plant, and perhaps on climatic conditions. The branched leafy plants 
separated in this way, furthermore, are not E. hypecoides. The generally 
accepted circumscription and range of EF. caespitosa, however, are not 
affected by removal from synonomy (or varietal staus) of the name 
E. hypecoides. Normally these species will not be confused. The primary 
affinities of E. hypecoides are not with E. caespitosa or E. tenuifolia but 
are with E. lemmonti, a species unknown to Bentham and to Gray. 

Greene examined the same holotypes in 1894. Later (1905), he sepa- 
rated E. caespitosa and E. hypecoides but E. tenuifolia disappeared and 
at least three new species were described which now seem indistinguish- 
able from E. hypecoides. The syntypes of two, viz. EL. eximia Greene (T. 
Brandegee, 30 March 1893, CAS 2633) and E. alcicornis Greene (T. 
Brandegee, 1891, CAS 2632), were collected at Alcalde, Fresno Co. The 
holotype of E. delitescens Greene ex Fedde was mounted on the same 
sheet with the syntype of E. alcicornis and probably was from the same 
collection. All were cited by Jepson (1922) as synonyms of E. lemmonu 
but, in my opinion, are morphological extremes of EL. hypecoides. The 
holotype of E. asprella Green (Eastwood, May 1897, CAS 2663), flower- 
less and nearly leafless, the basis for E. lemmonu var. asprella Jepson, 
probably is E. hypecoides. 

Bentham’s holotypes of Eschscholzia also were examined by Jepson 
who cited (1922) a number of specimens that he considered representa- 
tive of E. caespitosa var. hypecoides. Two of these (Jepson, May 1892, 
DS, JEPS, and 28 April 1893, JEPS), which he compared with the holo- 
type of E. hypecoides, do not seem to match it very well now. The speci- 
mens cited by him as referable to E. hypecoides mostly are from outside 
the geographical range of this species as I have determined it (fig. 1), 
and probably can be referred more satisfactorily to E. caespitosa. 


1964] ERNST: ESCHSCHOLZIA 287 


Eschscholzia lemmoni was described by Greene in 1887 from the col- 
lections of J. G. Lemmon. The type material was said to be pubescent 
throughout, the color of the petals orange, and the receptacle about one- 
quarter the length of the inch long corolla. The location of the gathering 
of the material was not given until 1905 when Greene mentioned (p. 290) 


/ 


Le Ah BT OMe EheV ir : ke cte 
eee oe 
hee i” ACF mi, Meat 2.064 Ring BS 


Caltlorima. Lipavins 183%. 
e oo Mos ee : f 
ceekte Acta BER ts Ty S take. top. : oe 
ae | a oe 
(hoe Baw lauleulunbuiliubueanlanta 
Califurnia, Pandas }sag. RYN. BLN. AMS Y, 


Fic. 3. Holotypes of Eschscholzia (K), both collected in California by David 
Douglas: left, E. caespitosa Benth. (two similar superposed plants) ; right, E. tenwz- 
folia Benth. 


2388 MADRONO [Vol. 17 


that ‘“The type of this remarkable species, obtained by Mr. Lemmon at 
Cholame, San Luis Obispo, Co., Calif., as long ago as 1887, has not yet 
been rediscovered.” At the same time he described two varieties (laxa 
and cuspidata) which now seem too indistinct to be maintained. 

A specimen of £. lemmonii collected by Lemmon cannot be located. 
An illustration (fig. 4) signed ‘““H. D. House, ’04” (US), conforms to the 
description of E. lemmonti. The drawing was inscribed ‘Eschscholtzia 
lemmonii Greene / San Luis Co., / Collected by J. G. Lemmon / 1887. 
Drawn from type / in Greene herb.” The specimen from which the draw- 
ing was made does not seem now to be in the Greene-Nieuwland Her- 
barium. This illustration did not exist when the description of the species 
was published and no other illustration of the whole plant is known; the 
one labeled E. lemmonii in Abrams (1944) is E. Aypecoides . 

A solitary bud at the California Academy of Sciences (CAS 2506) may 
be a fragment of the missing holotype. On the same sheet are three small 
plants of E. lemmonu (Brandegee, Alcalde, CAS 2505) bearing the anno- 
tation ‘“??? / E. L. G.” Another specimen on this sheet (Jared, May 
1893, CAS 2507) was annotated by Greene as ‘‘Eschscholtzia lemmonii, 
Green type ! var. cuspidata Greene.’”’ On another sheet (Hastwood, 
10 May 1893, CAS 2510), Greene wrote “Eschscholtzia lemmonii, Greene 
not typical; var. laxa Greene.” None of these annotations was dated and 
the varieties now do not seem distinct. All conform to my conception of 
E. lemmonii but none appears to me a suitable candidate for lectotype or 
neotype. 

In general aspect E. /emmonii is very similar to E. hypecoides. The 
cotyledons are linear spatulate and entire. The plants are annual, usually 
glaucous, the pubescence conspicuous (although some plants glabrate), 
and the flowers frequently on branched, leafy shoots with reduced, often 
opposite leaves on short petioles. Sometimes the plants are acaulescent 
with flowers on simple leafless peduncles. The buds mostly are fewer in 
number, larger in size, but almost always nodding when young. The petals 
usually are dark orange or red orange but sometimes yellow. Often there 
is a black spot at the base of the staminal filaments. The pollen have five 
or six colpae. The differences from E. hypecoides are the much larger 
buds and the proportionately greater size of the receptacles at all stages 
of development (table 1). At anthesis, the receptacles are opaque and 
glaucous but frequently become scarious or translucent. 

Flowering plants have been collected from February to May on adobe, 
calcareous sand, gravel, gypsum, heavy soil, loam, white clay, white shale, 
and sand at altitudes of 700-2900 ft. The habitat usually is open grass- 
land slopes in the eastern portion of the South Coast Ranges and the 
adjacent margin of the Central Valley from western Merced Co. in the 
north to Ventura Co. in the south (fig. 1). The gametic chromosome num- 
ber is six (Ernst, 1959; reconfirmed in Ernst 751, DS). This species 
mostly is distributed at somewhat lower altitude and further east than 
E. hypecoides. Near Alcalde, Fresno, Co., E. lemmontu and E. hypecoides 


1964] ERNST: ESCHSCHOLZIA 289 


SN 2 
Ee pe AE a | 
s\ SS fe LAY 2 


eae yy, 
Ve 


Wi a : 
i Ll lianal #14 & 
oun of 


“= Ny \) 


“aN 
ra lace? 1 hoe Ge \ 


; 
a o £ vez = YO” fe 


\\ 
YS 


UNITED STATES NATIONAL HERBARIUM. 2 
PLANTS OF CALIFORNIA. 


Each sellat i a Ya. Re re aCe: 
CAs Ss J 


belted fog eae One oo, Ve ae 
[BEF Ho Howes: 


Fic. 4. Drawing of E. lemmonii Greene (US). 


290 MADRONO [Vol. 17 


are found within a short distance and some flowers of the latter are un- 
usually large with orange rather than yellow petals but the difference in 
sizes of receptacles is maintained. 

The common occurrence of pubescence (plants infrequently glabrate), 
the nodding buds on shoots frequently with opposite leaves (or flowers 
sometimes on simple leafless scapes), the unusually large receptacles 
without conspicuous rim beneath the insertion of the calyx, and the 
darker petals will distinguish /. lemmonzu from all other species in the 
South Coast Ranges. The southeastern limit of distribution extends east- 
ward along the northern slopes of the Transverse Ranges in Kern Co. 
(Twisselmann 8037, 8060, 8079, US) and thus beyond the intended geo- 
graphical coverage of this paper. In the latter region a problem may arise 
in distinguishing depauperate and glabrate plants of L. lemmonu from 
depauperate plants of E. caespitosa subsp. kernensis Munz. 

Our knowledge of EL. rhombipetaia Greene rests on eleven collections, 
many without flowers. Neither the range nor the type materials were 
clearly set forth in the original description in 1885. Later, Greene (1905, 
p. 289) gave the distribution as “inconspicuous grain-field species, com- 
mon from Colusa Co. far southward along the foothills of the inner Coast 
Range and plains adjacent.”’ No specimens now are known from north of 
San Francisco Bay. The distribution (fig. 1), from north to south, seems 
to be Contra Costa Co.: Brandegee, April 1889 (DS); Curran, June 
1889 (CAS, DS); Greene, 25 March 1888 (UC, US); San Joaquin Co.: 
Hoover 1737 (UC); Alameda Co.: Eastwood & Howell 1949 (CAS); 
McClintock, 26 March 1950 (RSA); Stanislaus Co.: Hoover 4345 (DS, 
UC, US); and San Luis Obispo Co.: Hoover 7775 and 7861 (OBI); 
Lemmon Herbarium, in 1887-1888 (UC). Another specimen (CAS 2541) 
was collected by Greene from the locality “Lower San Joaquin Valley.” 
This one, not dated, was annotated ‘‘Part of my original material!” by 
Greene. The plants are annual and distinctive in this region for their 
small size, cespitose habit, erect buds, small petals, and rather conspicu- 
ous mature receptacles and fruits (table 1). The receptacles recall those 
of E. lemmonzu but the petals are smaller and the plants glabrous. 

The new combination £. caespitosa var. rhombipetala was made by 
Jepson but others have considered E. rhombipetala merely a synonym of 
E. caespitosa. It differs from E. caespitosa in the smallness of the flow- 
ers, the shortness of the peduncles (mostly not exceeding the leaves), 
and the texture of the herbage (not easily described). The chromosome 
number is unknown. The pollen appears to have five to eleven colpae 
(5 and 6 in Lemmon Herbarium, 1887-1888, and Hoover 1737, both 
UC; 8 and 9 in McClintock, 26 March 1950, RSA; 10 and 11 in Greene, 
25 March 1888, UC). The pollen of E. californica, E. caespitosa, E. hype- 
coides, and E. lemmonii usually have five or six colpae. 

The similarity of E. rhombipetala and some forms of E. minutiflora, 
a common and quite variable species in the desert regions of the South- 
west, occurring in Utah, Arizona, Nevada, California, and Baja Cali- 


1°64] ERNST: ESCHSCHOLZIA 291 


fornia, creates a problem. This species was described by Watson in 1876 
without citation of holotype or type locality; however, his material prob- 
ably was Watson 51 (US), collected at Truckee Pass, Nevada. Whether 
this species also occurs in the South Coast Ranges to the extent suggested 
by Munz (1959) and by Mosquin (1961, pp. 93-96) remains debatable. 
I have found only four collections from this region which might be refer- 
able to E. minutiflora (fig. 1). Two of these are from San Luis Obispo 
Co. (Armstrong 1112, alt. 1750 ft near Pozo, UC, and Axelrod 260, alt. 
2300 ft near Cuyama, RSA, UC) and two from Ventura Co. (Hoffmann, 
5 June 1930, Lockwood Valley, SBN, and Schretber 1045, road to Ozena 
UC). The nearest station of E. minutiflora to the east, toward the exten- 
sive desert distribution, perhaps is less than 40 miles distant. 

It is difficult to make a distinction between EL. rhombipetala and E. 
minutiflora in the South Coast Ranges because there is so little material 
available for comparison. Both species appear to have unusual disjunct 
localities in San Luis Obispo Co. (fig. 1). Normally, E. minutiflora 
(on the deserts) is annual, has upright, diffusely branched inflorescence 
shoots above a rosette of leaves, and the many flowers (except for the first 
few) are on relatively short pedicels. Eschscholzia rhombipetala also 
forms a rosette of leaves but there only are a few flowers, these on rela- 
tively long peduncles which do not seem to exceed the leaves. I find the 
pollen of the tentative E. minutiflora in the South Coast Ranges have 
nine to eleven colpae (9 to 11 in Armstrong 1112, UC, and 9 and 10 in 
Axelrod 260, UC). The mean numbers of colpae for 10 pollen grains from 
each of 27 plants of E. minutiflora (for which the chromosome number 
was determined) were given by Mosquin (1961, p. 95) as 8.2 to 10.4. 
The numbers of colpae alone seem insufficient to distinguish this species 
from E. rhombipetala. Since E. minutiflora of the California deserts is 
hexaploid with 18 pairs of chromosomes (Ernst, 1958, 1959; Mosquin, 
1961), its presence and distribution in the South Coast Ranges presum- 
ably could be verified on the basis of a few appropriate chromosome 
number determinations. If it does occur in this area, it would seem to be 
very local and rarely collected. 

The most common species of Eschscholzia in the South Coast Ranges, 
occurring in each county, is /. californica, distributed from the coastal 
strand through the mountains to the Central Valley of California and 
beyond. The original material, the type for the genus, was desribed from 
San Francisco by Chamisso and published with a plate in 1820. The hall- 
marks are the broad receptacle rim below the insertion of the calyx and 
the bifid cotyledons (only example in family). The receptacle rim, in 
this case, is a somewhat fleshy structure that is vasculated by a number 
of discreet, radiating, folded traces which differentiate in the receptacle, 
pass outward into the fleshy rim, then turn sharply back toward the re- 
ceptacle and finally enter the sepals (Ernst, 1962). The plants rarely are 
pubescent and sometimes persist for longer than a year. Their appear- 
ance during the hot and dry summer (with sparse foliage, small yellowish 


292 MADRONO [Vol. 17 


flowers) often contrasts markedly with the vernal appearance of the same 
plants (lush dense foliage, large orange flowers) (Jepson, 1922). A rather 
distinctive maritime phase is developed in some instances. 

Many of the biotypes of this species have been described as separate 
species, especially by Greene. Accepting the lack of experimental evi- 
dence, one readily could believe from the complex patterns of morpho- 
logical variation that this species hybridizes with others. If true, it would 
be helpful taxonomically to know how the distinctive receptacle rim and 
the bifid cotyledons are inherited in hybrids. Some plants much resembIl- 
ing E. californica but lacking well developed receptable rim present a 
taxonomic problem. Whether these are E. californica or E. caespitosa is 
diffcult to decide since the cotyledons seldom are available. 


Current practice has created of E. caespitosa a kind of taxonomic 
dumping ground. As popularly conceived this species equally is as vari- 
able as E. californica (taken in the broad sense) but somewhat less com- 
mon and less widely distributed. In the South Coast Ranges FE. caespitosa 
occurs at least in western Stanislaus, Monterey, San Luis Obispo, and 
Santa Barbara counties, usually at somewhat higher altitude than the 
lowest limits of EF. californica. The plants are annual, the receptacle 
moderate in size and without conspicuous rim beneath the calyx, and 
the cotyledons are undivided. The buds mostly are not clustered and 
nodding and the plants usually are glabrous (but + pubescent in other 
areas). Some of the plants seem obligately acaulescent while others pro- 
duce elongated inflorescence shoots. One of the more distinctive holotypes 
of the several binomials proposed for this complex is that of EZ. dolicho- 
carpa (Plaskett 84, CAS 2490), collected in Monterey Co. and described 
by Eastwood in 1903. 


Even after one has become accustomed to the spectrum of morpho- 
logical variation included in the popular conception of EF. caespitosa, the 
features of the holotype are surprising (Greene, 1905, p. 285). The holo- 
type (fig. 3), actually two similar superposed plants, to me is startlingly 
similar to E. californica and less like the plants usually identified as E. 
caespitosa. This especially is evident in the margins of the ultimate divi- 
sions of the foliage and the size and shape of the buds even though they 
lack a conspicuous rim. The cotyledons, of course, forever are missing. 
The peduncles of the earlier flowers are quite long but examination of the 
central portion of the plants suggests that later flowers probably would 
have been on shorter pedicels inserted on somewhat elongated shoots. 


An apparently similar biotype occasionally turns up in the South Coast 
Ranges (and infrequently from mountain systems further south) and 
although the match never seems to be quite perfect, it does seem possible 
that the holotype of E. caespitosa was collected along Douglas’s path 
in this region. Biologically, it may be just as sound to call these excep- 
tional plants (and the holotype of E. caespitosa) extremes of E. caltfor- 
nica. Since there seems no way to refute or to prove this hypothesis, the 


1564] ERNST: ESCHSCHOLZIA 293 


same specimens are just as likely extremes of the popular conception of 
E. caespitosa. 

The name of E. tenuifolza historically has been submerged under that 
of EL. caespitosa even though the holotype seems more distinctive. An 
apparently similar but frequently branching (sometimes pubescent) bio- 
type occurs along the western base of the Sierra Nevada and in the North 
Coast Ranges, especially in Mendocino and Lake counties, probably all 
beyond the limits of Douglas’s itinerary in California. In the South Coast 
Ranges, while none matches it very closely, here and there is a specimen 
that seems almost too similar to rule out the possibility that, as an excep- 
tional plant, the holotype of E. tenuzfolia also was collected in this region. 

More than a century of botanical collecting in the South Coast Ranges 
has not shown E. caespitosa and E. tenuifolia to be well defined species. 
Ironically, the two holotypes taken together provide a sort of range of 
morphological variation which seems to cover the popular conception of 
E. caespitosa. A biological understanding of this uneasy situation should 
follow a detailed comparison of E. californica and E. caespitosa through- 
out their respective geographical ranges. A taxonomic solution, however, 
which must be made in the South Coast Ranges, is not now in sight and 
only the following improvement in current practice is proposed. It seems 
sufficiently clear that neither E. rhombipetala nor E. hypecoides is con- 
specific with either E. caespitosa or E. tenuifolia. 


SUMMARY 


Six species of Eschscholzia occur in the South Coast Ranges, the type 
locality for all except E. minutiflora. The identity of E. kypecoides, ob- 
scurred since 1887, is clarified, an argument for its recognition as a species 
is advanced, and its similarity to E. lemmoni is pointed out. The simi- 
larity between E. rhombipetala and E. minutiflora in this region is men- 
tioned and the difficulties attending an interpretation of the holotypes 
of E. caespitosa and of E. tenuifolia are discussed. 


After this manuscript was accepted for publication I had the opportunity to visit 
the herbaria of the Royal Botanc Garden, Edinburgh (E), the British Museum 
(Natural History), London (BM), and the Royal Gardens, Kew (K). Now I am 
aware of additional specimens that probably are isotypes for Bentham’s E. caespitosa, 
E. hypecoides, and E. tenuifolia. Those shown in the accompanying illustrations are 
stamped “Herbarium Benthamianum” and bear Bentham’s own annotations. I pre- 
sumed these specimens to be holotypes but, in strict adherence to the Code (Art. 7), 
they should be lectotypes since they were not specifically singled out in the original 
descriptions. 


Department of Botany, Smithsonian Institution, Washington, D.C. 


LITERATURE CITED 


ApraMs, L.R. 1944. Illustrated flora of the Pacific States. Vol. 2. Stanford Univ. 
Press. 


294 MADRONO [Vol. 17 


BENTHAM, G. 1834. Report on some of the more remarkable hardy ornamental 
plants raised in the Horticultural Society’s garden from seeds received from 
Mr. David Douglas, in the years 1831, 1832, 1933. pp. 1-14. [Repaged reprint; 
see also Trans. Hort. Soc. London IT. 1:403-414. 1835]. 

Ernst, W.R. 1958. Chromosome numbers of some western Papaveraceae. Contr. 
Dudley Herb. 5:109-115. 

. 1959. Chromosome numbers of some Papaveraceae. Contr. Dudley Hedb. 
5:137-139. 

. 1962. A comparative morphology of the Papaveraceae. Unpublished Ph.D. 
dissertation, Stanford Univ. 

GREENE, E. L. 1905. Revision of Eschscholtzia. Pittonia 5:205-293. 

Jepson, W. L. 1922. A flora of California. Vol. 1. Assoc. Students Store, Univ. Calif., 
Berkeley. 

. 1933. David Douglas in California. Madrono 2:97-100. 

McKe vey, S. D. 1955. Botanical exploration of the Trans-Mississippi West, 1790- 
1850. Arnold Arboretum, Harvard Univ., Jamaica Plain. 

Mosguin, T. 1961. Eschscholzia covillei Greene, a tetraploid species from the Mojave 
Desert. Madrono 16:91-96. 

Muwz, P.A. 1959. A California flora. Univ. Calif. Press, Berkeley. 


NOTES AND NEWS 


ECHINOCHLOA ORYZICOLA IN CALIFORNIA.—For some years a Singular species of 
Echinochloa has been persisting in the rice fields at the Rice Experiment Station near 
Biggs, Butte Co., California. It closely resembles the rice plant in gross vegetative 
appearance but is readily distinguished by the densely hairy collar and sheath margins 
and in the absence of the ligule. The character and position of the hairs separate it 
from any forms of the common watergrass, Echinochloa crus-galli. The spikelets 
are 5-6 mm long, quite shiny and less hispid than those of watergrass. The lemma 
of the sterile floret is largely smooth and shiny, this smooth portion with a texture 
similar to that of the fertile one. It matures at the same time as rice. 

The grass was first collected at the Biggs station by the author on September 17, 
1957 (Crampton 4626, AHUC). A specimen of the first collection was identified by 
N. L. Bor, Royal Botanic Gardens, Kew, as E. oryzicola var. mutica. It is native 
in the Far East but more recently has been imported with rice into middle Asia 
and the Caucasus (Fl. USSR 2:33. 1934). It is regarded as a noxious rice weed by 
the Russians. Vasinger-Alektorova (Bull. Appl. Bot. Genet. Pl. Breeding 25:109-152. 
1931) clearly emphasjzed the weedy character of the grass and the problems of its 
management in his studies on rice weeds of the maritime Far East. 

It is not known how long the grass has been growing at the Biggs station though 
it is quite likely that it was introduced as an impurity in oriental rice varieties. 
P. B. Kennedy during his research on rice weeds in California and subsequent 
publication (Calif. Agric. Exper. Sta. Bull. 356:467-494. 1928), had collected this 
Echinochloa at a temporary rice station at Cortena, Colusa Co. (Kennedy, in 1922, 
1925, 1928, AHUC). Kennedy recognized the plants as “a new form of watergrass,” 
however, did not publish a specific or varietal name nor allude to it in his publication 
on rice weeds. 

The author visited Cortena in September, 1963, and collected in the rice there 
an Echinochloa (Crampton 6892, AHUC) which may be called E. oryzicola f. glabra 
(FI. USSR 2:33. 1934). It is similar in habit, inflorescence and spikelet characters 
to those previously collected by Kennedy and to those collected at the Biggs station. 
The sheaths and collar, however, lack the characteristic pubescence, though a band 
of short appressed hairs was present around the base of the lowermost sheaths. 
Considerable search did not reveal var. mutica as having persisted at Cortena. 


1964] NEWS AND NOTES 295 


As far as it is known, E. oryzicola is, and has been, confined to the above localities 
and certainly is not the weed problem presented by the common watergrass. A 
description of this grass new to California and possibly to North America follows: 

Echinochloa oryzicola (Vasing.) Vasing. var. mutica Vasing. Annual; culms 
40—75 cm tall, stiffly erect, rooting and sometimes branching at the lower nodes, 
bearing leaves to the base of the panicle; blades stiffish, folded, tapering to a 
sharp point, strongly scabrous on the margins and upper surface; sheaths keeled, 
with closely appressed non-pustulate based hairs around the base and conspicuous 
long, coarse hairs along the margins in the uppermost portion; collar of lower 
sheaths with a conspicuous ring of numerous, stiff, tawny hairs, these hairs and 
those of the sheath margin with swollen or pustulate bases, several hairs often arising 
from a common pustule; ligule absent; inflorescence green or light green, 8-13 cm 
long, axis scabrous bearing 15 or less 1—sided racemes 5 cm or less long; rachis 
scabrous with scattered, stiff pustulate-based hairs, the hairs more numerous about 
the point of insertion on the axis; spikelets 5—6 mm long, awnless, shiny, commonly 
in clusters of 2 or 3 or sometimes solitary along the rachis; first glume about 4% the 
length of the spikelet, ovate, prominently 5—nerved, ciliolate, scabrous over the 
back and on the nerves, second glume prominently 5—nerved, apiculate, ciliolate 
towards the apex, as long as the spikelet, scabrous over the back and on the nerves 
with some scattered, stout, pustulate-based hairs along the nerves in the upper half; 
sterile floret staminate, equalling the second glume, the lemma mostly smooth and 
shiny over the back resembling the texture of the enclosed fertile floret, 5—nerved, 
the central nerve (where back is smooth and shiny) indistinct, the lateral pairs 
marginal, prominent, scabrous with some scattered pustulate-based hairs, ciliate 
along the margin towards the apex, palea thin, about 7% the length of the lemma; 
fertile floret stramineous, 4—4.5 mm long, smooth and shiny except for the puberu- 
lent apex, indistinctly nerved. (Based on Crampton 4626 and 6887). 

Echinochloa oryzicola {. glabra Vasing. Vegetatively similar to the preceding 
except for absence of hairs on the collar and with only a few scattered hairs or none 
on the sheath margins. Inflorescence and spikelets are similar to var. mutica excepting 
the more obvious marginal pubescence at the apex of the sterile lemma. (Based on 
Crampton 6892.) 

The long-awned var. aristata Vasing. has not been observed in California thus 
far —BEECHER CRAMPTON, Department of Agromony, University of California, Davis. 


NEW COMBINATIONS IN WESTERN NorTH AMERICAN VIOLETS.—In a recent paper 
(Madrono 17:173-197. 1964) I treated Viola aurea and V. aurea ssp. mohavensis 
as subspecies of V. purpurea. This treatment was in accord with M. S. Baker’s 
original concepts. In a letter dated Feb. 2, 1939, he had convinced me that the two 
taxa, aurea and mohavensis, most naturally should be treated as subspecies of 
V. purpurea along with eight others. In a preliminary manuscript of his 1949 paper 
(Madrofio 10:110-128) Baker listed the 10 subspecies as they appeared in my 
recent paper. By the time the paper had appeared Baker had changed his mind and 
considered aurea and mohavensis as subspecies of V. aurea. In 1953 Baker (Ma- 
drofo 12:8-13) presented a review of V. aurea and described ssp. arizonensis as new 
without my knowledge. This latter subspecies is exceedingly rare and according to 
the description appears to be only an extreme variant of V. purpurea ssp. mohavensis. 
Since Baker did not publish these taxa as subspecies of V. purpurea the formal trans- 
fers are being made as follows: Viola purpurea Kell. ssp. aurea (Kell.) J. Clausen, 
comb. nov. (V. aurea Kell., Proc. Calif. Acad. 2:185. 1862. V. purpurea Kell. var. 
aurea (Kell) Baker ex Jepson, Flora Calif. 2:521. 1936.) Viola purpurea Kell. ssp. 
mohavensis (Baker & Clausen ex Baker) J. Clausen, comb. nov. (V. aurea Kell. ssp. 
mohavensis Baker & Clausen ex Baker, Madronfo 12:9. 1953. V. aurea Kell. ssp. 
arizonensis Baker & Clausen ex Baker, Madrono 12:11. 1953.)—J. CLAusEN, De- 
partment of Plant Biology, Carnegie Institution of Washington, Stanford, California. 


296 MADRONO 


[Vol. 17 


INDEX TO VOLUME XVII 


Classified entries: Chromosome numbers of plants, documented; Reviews. New 
Scientific names are in boldface type. Un-annotated taxa in floral lists are ommitted 


from the Index. 


Acacia: cornigera complex, Nomencla- 
tural problems in the, 198; sphaero- 
cephala, 199 

Adenocaulon: bicolor, 52; chilense, 52 
adhaerescens, 52; chilense, 52 

Agave: asperrima, 166; celsii, 164; Cy- 
tological observations on some genera 
of Agavaceae, 163; desertii, 166; fili- 
fera, 164; lechuguilla, 164; nexans, 
168; salmiana, 166; toumeyana, 165, 
victoria-reginae, 164. 

Ageratum: albidum, 134; cf. corymbo- 
sum, 134; salicifolium 134 

Agropyrum trachycaulum, A controlled 
hybrid between Sitanion hystrix and, 
10 

Anderson, D.: Notes on the leaf epi- 
dermis and chromosome number of 
Swallenia (Gramineae), 201 

Anderson, L. C.: Taxonomic notes on 
the Chrysothamnus viscidiflorum com- 
plex (Asterae, Compositae), 222 

Aphanostephus: cf. pachyrrhizus, 134; 
ramosus, 134 

Aporpogaster, sect. nov., 26 

Arceuthobium campylopodum, 254 

Arizona, Notes on the flora of, 236 

Arnica foliosa, 236 

Asclepias cryptoceras, 236 

Asplenium vesperitnum, A new locality 
for, 172 

Arctostaphylos uva-ursi, 97 


Baccharis sordescens, 134 

Bahia: glandulosa, 137; schaffneri, 137; 
xylopoda, 137 

Baileya multiradiata, 137 

Baker, W. H.: New distributional record 
for Heleochloa alopecuroides in Ore- 
gon, 197 

Bartlettia scaposa, 138 

Bebbia juncea, 137 

Beschorneria yuccoides, 168 

Besseya: and Synthyris, Natural and 
artificial hybrids of, 109; bullii, 114; 
plantaginea, 114; rubra, 109 

Bhatt, D. D.: Plant collecting in Nepal, 
145 

Bidens: cf. aurea, 136; pilosa, 136; cf. 
reptans, 136 

Blestogaster, sect. nov., 23 

Boyle, W. S.: A controlled hybrid be- 
tween Sitanion hystrix and Agropyrum 
trachycaulum, 10 


Calder, J. A. and R. L. Taylor: A new 
species of Isopyrum endemic to the 
Queen Charlotte Islands of British Co- 
lumbia, 69 


Calea: scabra, 137; urticifolia, 137 

California Botanical Society, 144 

Calypso bulobsa in the Santa Cruz 
Mountains, California, 92 

Cave, M.S.: Cytological observations on 
some genera of the Agavaceae, 163 

Ceanothus cf. griseus, 43 

Chaenactis: carphoclinia var. attenuata, 
137; stevioides, 137 

Chambers, K. L.: Saxifraga eschscholtzii 
Sternb., 203; and L. R. J. Dennis: 
New distributions for four grasses in 
Oregon, 91 

Chaparral shrubs during severe drought, 
Extended dormancy of, 161 

Chilean plants, Chromosome numbers of 
some phytogeographically interesting, 
52 

Chilopsis: linearis, Notes on Tetracoccus 
and, in Baja California, Mexico, 92 

Chromosome numbers in the Composi- 
tae, VII, 128 

Chromosome numbers of plants, docu- 
mented: Aquilegia nevadensis, 286; 
Arcytophyllum thymifolium, 116; Ar- 
temisia nana, 266; Baptisia calycosa, 
116, perfoliata, 116, simplicifolia, 116; 
Brachycome ciliaris var. ciliaris, 116, 
iberidifolia, 116; trachycarpa, 116; 
Bouteloua breviseta, 266, rothrockil, 
266; Bromus macrostachya, 266; Car- 
duus argyroa, 266, velebiticus, 266; 
Carex acutiformis, 266, alba, 266, ar- 
gyroglochin, 266, atrofusca, 266, au- 
stroalpina, 266, brizoides, 266, cam- 
posii, 266, cuprea, 266, curvata, 266, 
durieui, 266, ericetorum var. approxi- 
mata, 26, flavella, 266, frigida, 267, gri- 
oletii, 267, halleriana, 267, mucronata, 
267, ornithopoda var. elongata, 267, 
parviflora, 267, pauciflora, 267, prae- 
cox, 267, supina, 267, umbrosa, 267; 
Chaetopappa asteroides, 267; Clero- 
dendron aculeatum, 116; Crocidium 
multicaule, 267; Dalea citriodora, 116, 
schottii var. puberula, 116; Daphnop- 
sis americana ssp. caribaea, 116, philip- 
piana, 116; Desmanthus_ illinoensis, 
116; Digitaria patens, 267; Eleocharis 
acicularis, 267, arenicola, 267, atropur- 
purea, 267, compressa, 267, engelman- 
nii, 267, nervata, 267, obtusa, 267; par- 
vula, 267; Eragrostis lehmanniana, 
267; Eriochloa puncatata, 267; Frank- 
linia alatamaha, 267; Garrya elliptica, 
117; Hedyosmum arborescens, 117; 
Helianthus debilis var. cucumerifolium, 
267; Jaumea carnosa, 267; Leucothoe 
fontanesiana, 117; Lithospermum ci- 


1964] 


nereum, '267; Lolium temulentum, 
267; Lycurus phleoides, 267; Lythrum 
alatum, 117; Moehringia dasyphylla, 
267, markgrafil, 268, papulosa, 268, 
tommasinii, 268; Montia perfoliata, 
117, sibirica, 117; Mucizonia hispida, 
268; Omphalodes vera, 268; Onosma 
cinerascens, 268, frutescens, 268, tri- 
dentinum, 268; Orobanche grayana 
var. nelsonil, 117; Oxytropis ame- 
thystea, 268; Palafoxia callosa, 268; 
Panicum texanum, 268; Parietaria 
pennsylvanica, 117; Paspalum distich- 
um, 268; Petalostemum feayi, 117; 
Petasites frigidus var. palmatus, 117; 
Phytolacca decandra, 117; Polygonum 
scandens, 117; Quincula lobata, 268; 
Ribes laxiflorum, 117, sanguineum var. 
glutinosum, 117; Scleropoa rigida, 268; 
Sporobolus contractus, 268, virginicus, 
268; Trichachne patens, 267; Willkom- 
mia texana, 268 
Chrysactinia pinnata, 137 
Chrysanthellum mexicanum, 137 


Chrysothamnus: axillaris, 225; linifo- 
lius, 226; molestus, 222; spathulatus, 
226; viscidiflorus, 223; viscidiflorus 
complex (Astereae, (Compositae), 


Taxonomic notes on the, 22; viscidi- 
florus ssp. humilis, 223; viscidiflorus 
ssp. planifolius, 223; viscidiflorus ssp. 
stenophyllus, 225 

Cienfuegosia: argentina var. hasslerana 
f. escholtzioides, 84; drummondii, 84 

Clappia suaedifolia, 138 

Clarkia: springvillensis, 220; temblori- 
ensis, 220; unguiculata, Two new 
species related to, 219 

Clausen, J.: Cytotaxonomy and distribu- 
tional ecology of western North Amer- 
ican violets, 173; New combinations in 
western North American violets, 295 

Collomia: biflora, 206; cavanillesii, 206; 
debilis, 206; diversifolia, 208; grandi- 
flora, 208; heterophylla, 208; larsenii, 
208; linearis, 208; macrocalyx, 208; 
mazama, 210; The pollen grain mor- 
phology of, as a taxonomic tool, 205; 
rawsoniana, 212; tenella, 213; tinc- 
toria, 213; tracyi, 213 

Condalia mexicana Schlecht, var. peta- 


lifera M. C. Johnston, var. nov., 280 


Condit, I. J.: Cytological studies in the 
genus Ficus. III., 153 

Conoclinum greggii, 134 

Conyza: bonariensis, 134; coronopifolia, 
134; gnaphaloides, 134 

Copeland, E. B., 235 

Coreopsis cyclocarpa, 137 

Cosmos: bipinnata, 137; parviflorus, 137 

Crandall, T. A.: Calypso bulbosa in the 
Santa Cruz Mountains, California, 92 

Crampton, B.: Echinochloa oryzicola in 
California, 294 


INDEX 297 


Croptilon divaricatum, 134 

Crossosoma: bigelovii, 68; californica, 
68; Chromosome numbers in, 68 

Cupressus: and Pinus after thirteen years 
in Mendocino County, California, Sur- 
vival of transplanted, 250; abramsi- 
ana, 252; goveniana, 39, 252; pygmaea, 
252; sargentii, 252 


Draba nivalis var. elongata, 104 

Doryanthes palmeri, 169 

Douglas, David, and the digger pine, 
some questions, 227 

Dyssodia: pinnata, 137; porophylloides, 
137; setifolia, 137 


Echinochloa: cruz-galli, 294; oryzicola 
in California, 294 

Ectosperma alexandrae, 88 

Equisetum hiemale var. californicum, 44 

Erigeron: affinis, 134; delphinifolius, 134 

Ernst, W. R.: Review, Flora of our sier- 
ran national parks, 171; The genus 
Eschscholzia in the South Coast 
Ranges of California, 281 

Eschscholzia: alcicornis, 286; asprella, 
286; caespitosa, 281; californica, 283; 
crocea, 286; delitescens, 286; dolicho- 
carpa, 292; eximia, 286; hypecoides, 
281; in the South Coast Ranges of 
California, The genus, 281; lemmonii 
281; minutiflora 283; rhombipetala 
283; tenuifolia, 286 

Eupatorium prunellifolium, 134 


Ficus: acanthocarpa, 154; afzelii, 154; 
amplissiam, 154; aurantiaca var. par- 
vifolia, 154; aurea, 154; auriculata, 
154; avi-avi, 154; awkeotsang, 154; 
bengalensis, 154; burkei, 154; bussel, 
154; cabusana, 154; camarinensis, 154; 
capensis, 154; citrifolia, 154; cocculi- 
folia var. sakalavarum, 154; colum- 
naris, 154; congesta, 154; coronata, 
154; costaricana, 154; Cytological 
studies in the genus, 153; doliaria, 154; 
dusenii, 154; elastica, 154; eximina 
var. glabra, 154; garciniaefolia, 154; 
geniculata, 154; gnaphalocarpa, 154; 
goldmanii, 154; hillii, 154; hispida, 
154; hochstetteri, 154; insipida, 154; 
iteophylla, 154; krishnae, 154; lapa- 
thifolia, 154; macrosyce, 154; macro- 
phylla, 154; mallotocarpa, 154; mam- 
milifera, 154; minahassae, 154; monc- 
kii, 154; montana, 154; nekbudu, 154; 
nota, 154; nympheaefolia, 154; obtusi- 
folia, 154; palmeri, 154; perforata, 
154; pertusa, 154; petiolaris, 154; pi- 
losa, 154; preusii, 154; pretoriae, 154; 
procera var. crassiramea, 154; pumila, 
154; radulina, 154; retusa var. nitida, 
154; ribes, 154; rigo, 154; rumphii, 
154; satterthwaitei, 154; scabra, 154; 
soldanela, 154; sonderi, 154; stricta, 


298 MADRONO 


154; stuhlmanii, 154; subcordata, 154; 
thonningii, 154; tinctoria, 154; tsiela, 
154; umbellata, 154; urbaniana, 154; 
urceolaris, 154; variegata, 154; volk- 
ensii, 154; wildemaniana, 154; wilsonil, 
154 

Flaveria ramosissima, 137 

Fouquieria splendens, 230 

Franseria chamissonis, 52 

Fryxell, P. A.: Cleistogamy in Malvaceae, 
83 

Furcraea andina, 168 


Gaillardia pinnatifida, 137 

Garrya elliptica, 44 

Geraea canescens, 136 

Gnaphalium cf. leptophyllum, 135 

Gossypium; australe, 84; bickii, 85 

Griffin, J. R.: David Douglas and the 
digger pine: some questions, 227 

Grindelia: inuloides, 135; oxylepis var. 
eligulata, 135; robinsonii, 135 

Gutierrezia glutinosa, 135 


Haplopappus gracilis, 135 

Harvey, H. T.: A note on the type lo- 
cality of Tetracoccus ilicifolius, 268 

Harvey, R. A. and H. A. Mooney: Ex- 
tended dormancy of chaparral shrubs 
during severe droubt, 161 

Heiser, C. B.: Artificial intergeneric hy- 
birds of Helianthus and Viguiera, 118 

Heleochloa: alopecuroides, 92; alopecu- 
roides in Oregon, New distribution 
record for, 197 

Helianthus: agrestis, 121; angustifolius, 
120; annuus, 121; Artificial intergen- 
eric hybrids of, and Viguiera, 118; 
atrorubens, 121; canus, 121; carnosus, 
121; debilis, 120; divaricatus, 121; 
grosseserraus, 121; laciniatus, 121; mi- 
crocephalus, 121; neglectus, 121; niv- 
eus, 121; occidentalis, 121 

Heterotheca inuloides, 135 

Hibiscus denudatus, 83 

Hieracium cf. crepidispermum, 139 

Higgins, Ethel Bailey, 144 

Hooker oak, Note on damage to the, 143 

Hordeum jubatum — caespitosum — 
brachyantherum complex in Alaska, 
The, 269 

Hymenoxys: acaulis, 137; anthemoides, 
Cytophyletic analysis of, 27; cf. line- 
arifolia, 138; odorata, 138; odorata, 
Cytophyletic analysis of, a recapitula- 
tion, 77 

Hypericum anagalloides, 236 


Isocoma: heterophylla, 135; veneta, 135 

Isopyrum: A new species of, endemic to 
the Queen Charlotte Islands of British 
Columbia, 69; biternatum, 70; hallii. 
70; occidentale, 70; savilei, 70; stipi- 
tatum, 70 


[Vol. 17 


Jaeger, E. C.: Notes on Tetracoccus and 
Chilopsis in Baja California, 92 

Joe, B. A new locality for Asplenium 
vespertinum, 172 

Johnston, M. C.: Condalia mexicana 
Schlecht. var. petalifera, 280 

Juglans hindsii, the central California 
black walnut, native or introduced ?, 1 


Kimnach, M.: Review, Native orchids of 
Trinidad and Tobago, 30 

Kobresia myosuroides, 98 

Kruckeberg, A. R. and F. L. Hedglin: 
Natural and artificial hybrids of Bes- 
seya and Synthryris, 109 

Kuijt, J.: Review, The morphology of 
pteridophytes, 90; A peculiar case of 
hemlock mistletoe on larch, 254 


Langenheim, J. H. and J. W. Durham: 
Quaternary closed-cone pine flora from 
travertine near Little Sur, Calif., 33 

Laphamia lindheimeri, 138 

Larsen, E.: A new species of pine from 
Mexico, 217 

Lindsay, G. E.: Ethel Bailey Higgins, 
144 

Loeblich, A. R., III: The pollen grain 
morphology of Collomia as a taxo- 
nomic tool, 205 

Lolium: A note on taxonomic characters 
in, 79; multiflorum, 79; perenne, 79 

Lyonoxthamnoxylon: from the Lower 
Pliocene of western Nevada, 257; 
nevadensis, 258 

Lyonothamnus; floribundus, 260; mo- 
havensis, 264; parvifolius, 264 


Malacothrix fendleri, 139 

Mason, C. T.: Notes on the Flora of 
Arizona. III, 236 

McMillan, C.: Survival of transplanted 
Curpessus and Pinus after thirteen 
years in Mendocino County, 250 

McMinn, Howard E., 115 

Major, J. and S. A. Bamberg: Some 
cordilleran plant species new for the 
Sierra Nevada of California, 93 

Malvaceae, Cleistogamy in, 83 

Melanthera angustifolia, 136 

Mentzelia: chrysantha, 19; Cytotaxo- 
nomic observations on, 16; decapetala, 
18; densa, 19; laciniata, 20; laevicaulis, 
17; multicaulis, 21; multiflora, 20; 
nuda, 18; rusbyi, 19; speciosa, 19; 
strictissima, 18 

Meyer, F. G.: Review, Meet flora Mexi- 
cana, 143 

Mia, M. M., B. B. Mukherjee, and R. K. 
Vickery: Chromosome counts in the 
section Simiolus of the genus Mimulus 
(Scrophulariaceae). VI, 156 

Microsporogaster, sect. nov., 24 

Milleria quinqueflora, 135 

Mimulus: cardinalis, 54; Chromosome 


1964] 


counts in section Erythranthe of the 
genus, 53; Chromosome counts in the 
section Simiolus of the genus, 156; 
eastwoodiae, 54; glabratus, 159; gut- 
tatus, 159; lewisii, 54; luteus, 159; 
nasutus, 159; nelsonii, 54; tigrinus, 
159; verbenaceous, 54 

Mitchell, W. W. and A. C. Wilton: The 
Hordeum jubatum — caespitosum — 
brachyantherum complex in Alaska, 
269 

Mistletoe, parasitic on larch, A peculiar 
case of hemlock, 254 

Molinia caerulea, 91 

Mooney, H. A. and B. R. Strain: Bark 
photosynthesis in ocotillo, 230 

Moore, D. M.: Chromosome numbers of 
some phytogeographically interesting 
Chilean plants, 52 

Myrica californica, 42 


Nardus stricta, 91 

Nepal: climatic zones of, 146; floristic 
regions in, 157; Plant collection in, 145 

Nicolletia edwardsii, 138 

Nolina; beldingii, 168; bigelovii, 168; 
parviflora, 169 

Notes and News: 32, 68, 91, 115, 127, 
143, 172, 197, 203, 235, 268, 280, 294 

Nuphar polysepalum, 236 

Oregon, New distributions for four 
grasses in, 91 

Ornduff, R.: Review, Flora of New Zea- 
land, Vol. I, 66; Review, Aquatic plants 
of the Pacific Northwest with vege- 
tative keys, 30; Review, Vascular 
plants of the Pacific Northwest, Part 
3, 90 


Page, V. M.: Lyonothamnoxylon from 
the Lower Pliocene of western Ne- 
vada, 257 

Parthenium fruiticosum, 135 

Pectis: depressa, 138; latisquama, 138; 
saturejoides, 138; tenella, 138; cf. 
texanna, 138 

Pedicularis crenulata, 104 

Pelton, J.: Review, The Quiet Crisis, 
233 

Perymenium; flexuosum, 136; hypoleu- 
cum, 136 

Photosynthesis in ocotillo, Bark, 230 

Pinaropappus roseus, 139 

Pinus: A new species of, from Mexico, 
217; bolanderi, 252; contorta, 252; 
coulteri, 228; martinezii, 217; muri- 
cata, 38, 252; murrayana, 252; Quater- 
nary closed-cone pine flora from trav- 
ertine near Little Sur, California, 33; 
radiata, 38; sabiniana, 227; Survival 
of transplanted Curpessus and, after 
thirteen years in Mendocino County, 
California, 280 


INDEX 299 


Porophyllum: coloratum, 138; ervend- 
bergii, 138; scoparium, 138 

Powell, A. M. and B. L. Turner: Chro- 
mosome numbers in the Compositae. 
VII, 128 

Pseudoclappia arenaria, 138 

Pseudotsuga menziesii, 39 

Psilostrophe: cooperi, 138; villosa, 138 


Quercus: agrifolia, 40; lobata, 143 


Raven, P. H. and M. S. Cave: Chromo- 
some numbers is Crossosoma, 68 
Reviews: Allan, H. H., Flora of New 
Zealand, Vol. 1, 66; Enari, L., Orna- 
mental shrubs of California, 140; Fer- 
ris, R. S., Death Valley wild flowers, 
140; Jones, G. N., Flora of Illinois, 
170; Knobloch, I. W. and D. S. Cor- 
rell, Ferns and fern allies of Chihua- 
hua, 89; McKenny, M., The savory 
wild mushroom. A Pacific northwest 
guide, 32; Munz, P. A., California 
spring wildflowers, California desert 
wildflowers, and California mountain 
wildflowers, 140; Pesman, M. W., 
Meet flora Mexicana, 143; Pusateri, 
S. J., Flora of our sierran national 
parks, 171; Schultes, R. E., Native 
orchids of Trinidad and Tobago, 30; 
Shreve, F. and I. L. Wiggins, Vege- 
tation and flora of the Sonoran Desert, 
234; Sporne, K. R., The morphology 
of pteridophytes, 90; Steward, A. N., 
L. R. Dennis, and H. M. Gilkey, 
Aquatic plants of the Pacific North- 
west with vegetative keys, 30; Udal, 
S., The quiet crisis, 233; Wiggins, I. L. 
and J. H. Thomas, A flora of the 
Alaskan arctic slope, 31 
Ribes sanguineum var. glutinosum, 42 
Rollins, R. C.: Review, Vegetation and 
flora of the Sonoran Desert, 234 
Rubus, parviflorus, 42; vitifolius, 42 
Rudd, V. E.: Nomenclature problems in 
the Acacia cornigera complex, 198 
Russell, N. H. and F. S. Crosswhite: An 
analysis of variation in Viola nephro- 
phylla, 65 


Sabazia: cf. liebmannii, 137; cf. michoa- 
canna, 137 

Salix brachycarpa, 103 

Sanvitalia: ocymoides, 135; cf. procum- 
bens, 135 

Sartwellia mexicana, 138 

Saxifraga eschscholtzii, 203 

Scrabrogaster, sect. nov., 23 

Scirpus pumilus, 100 

Selliera radicans, 52 

Senecio: filifolium, 138; monoensis, 138 

Sharsmith, H.: Review, California spring 
wildflowers, California desert wild- 
flowers, California mountain’ wild- 
flowers, Death Valley wildflowers, 
Ornamental shrubs of California, 140 


300 


S:eglingia decumbens, 91 

Sierra Nevada of California, Some cor- 
dilleran plant species new for the, 93 

Simsia: foetida, 136; lagasciformis, 136; 
cf. megacephala, 136; triloba, 136 

Singer, R. and A. H. Smith: A revision 
of the genus Thaxterogaster, 22 

Sitanion hystrix, A controlled hybrid be- 
tween, and Agropyron trachycaulum, 
10 

Smith, S. G.: Review, A flora of the 
Alaskan arctic slope, 31 

Speese, B. M. and J. T. Baldwin, Jr.: 
Cytophyletic analysis of Hymenoxys 
anthemoides, 27 

Soderstrom, T. R. and H. F. Decker: 
Swallenia, a new name for the Cali- 
fornia genus Ectosperma, 88 

Stern, K. R.: Note on damage to the 
hooker oak, 143 

Stevia: elatior, 134; lucida, 134; pur- 
purea, 134; rhombifolia, 134;  salici- 
folia, 134; serrata, 134; viscida, 134 

Swallenia: a new name for the Califor- 
nia genus Ectosperma (Gramineae), 
88; alexandrae, 88, 201; Notes on the 
leaf epidermis and chromosome num- 
ber of, 201 

Synthyris: missurica, 109; Natural and 
a*tificial hybrids of Besseya and, 109; 
pinnatifida, 114; platycarpon, 112; 
reniformis, 112; schizantha, 112; stel- 
lata, 114 


Tagetes cf. elongata, 138 

Tavares, I.: Review, The savory wild 
mushroom. A Pacific Northwest guide, 
og 

Tetracoccus: hallii, Notes on, and Chi- 
lopsis in Baja California, Mexico, 92; 
ilicifolius, A note on the type locality 
of, 268 

Thaxterogaster: A revision of the genus, 
22; brevisporum, 23; conicum, 26; 
dombeyi, 24; fragile, 25; leucocepha- 
lum, 24; magellanicum, 24; pingue, 
24; porphyreum, 26; scabrosum, 23; 
subalbidum, 24; violaceum, 24 

Thiers, H. D.: The genus Xerocomus 
Quelet in northern California, 237 

Thompson, H. J.: Cytotaxonomic ob- 
servations on Mentzelia, 16 

Thomsen, H. H.: Juglans hinds, the 
central California black walnut, native 
or introduced ?, 1 

Thornber, John James, 32 

Townsendia mexicana, 135 

Trixis californica, 139 

Tryon, R. M.: Review, Ferns and fern 
allies of Chihuahua, Mexico, 89 

Turner, B. L.: Cytophyletic analysis of 
Hymenoxys odorata: A recapitulation, 
heh 


Varilla texana, 138 
Vasek, F. C.: Two new species related 


MADRONO 


[Vol. 17 


to Clarkia unguiculata, 219; and J. K. 
Ferguson: A note on taxonomic char- 
acters in Lolium, 79 

Vickery, R. K., Jr., B. B. Mukherjee, 
and D. Wiens: Chromosome counts in 
section Erythranthe of the genus Mim- 
ulus (Scrophulariaceae). II, 53 

Viguiera: Artificial hybrids of Helian- 
thus and, 118; adenophylla, 121; del- 
toidea var. parishii, 136; dentata, 121; 
longifolia, 121; multiflora, 121; ovalis, 
121; porteri, 121 

Viola: adunca ssp. adunca, 196; adunca 
ssp. ashtona, 196; adunca ssp. oxy- 
ceras, 196; adunca ssp. radicosa, 196; 
alliariifolia, 193; bakeri ssp. baker, 
193; bakeri ssp. shastensis, 194; bi- 
flora, 193; blanda, 195; brevistipulata, 
193; canadensis, 193; charlestonensis, 
194; clauseniana, 195; cognata, 195; 
crassa, 193; douglasii, 193; epipsila, 


195; eriocarpa, 193; glabella, 193; 
hallii, 193; howellii, 196; incognita, 
195; kiskidai, 193; lanceolata, 195; 


linguifolia, 193; lobata, 193; mcclos- 
keyi, 195; mocabeiana, 195; nephro- 
phyila, An analysis of variation in, 
56; nephrophylla, 195; nuttallii, 194; 
occidentalis, 195; ocellata, 193; orbi- 
culata, 193; pallens, 195; palustris ssp. 
brevipes, 196, 236; pedunculata ssp. 
pedunculata, 194; pedunculata ssp. 
tenuifolia, 194; praemorsa ssp. major, 
194; praemorsa ssp. oregona, 194; 
praemorsa ssp. praemorsa, 194; pri- 
muliiolia, 196; pubescens, 193; pur- 
purea ssp. atriplicifclia, 194; purpu- 
rea ssp. aurea, 194, 295; purpurea 
ssp. dimorpha, 194; purpurea ssp. 
geophyta, 194; purpurea ssp. integri- 
folia, 194; purpurea ssp. mesophyta, 
194; purpurea ssp. mohavensis, 194, 
295; purpurea ssp. purpurea, 195; pur- 
purea ssp. venosa, 195; purpurea ssp. 


xerophyta, 195; quercetorum, 195; 
rotundifolia, 193; rugulosa, 193; sco- 
pulorum, 193; sempervirens, 193; 


sheltonii, 193; shihokiana, 196; to- 
mentosa, 194; utahensis, 195; valli- 
cola, 194 

Violets: Cytotaxonomy and distribution- 
al ecology of western North American, 
173; New combinations in western 
North American, 295 

Webster, G. L.: Review, Flora of Illinois, 
170 


Xerocomus: chrysenteron, 246; illudens, 
241; subtomentosus, 243; The genus, 
in northern California, 237; truncatus, 
244; zelleri, 247 


Zaluzania: globosa, 136; montagnifolia, 
136; robinsonii, 136 

Zinnia: angustifolia, 135; haageana, 135; 
cf. leucoglossa, 135; tenella, 135 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree 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 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
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. Fourth Edition, 1959). 

Articles may be submitted to any members of the Editorial Board. 

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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


A quarterly journal devoted to the publication of botanical re- 
search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


Volume I, 1916-1929 . . . . = . $5.00 
Volume II, 1930-1934 . . . . . 5.00 
Volume ITI, 1935-1936 Pweg Ve ae EO 
Volume IV, 1937-1938. . . . . 5.00 
Volume V, 1939-1940 . . . . . 5.00 
Volume VI, 1941-1942 es eee tee OO 


Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 S ca waubeete te 0100 
Volume X, 1949-1950 . . . . . 7.00 
Volume XI, 1951-1952 Be ride co het e208 
Volume XII, 1953-1954 . . . = . + 8.00 
Volume XIII, 1955-1956 . . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 
Volume XV, 1959-1960 » a ee 1 O:00 
Volume XVI, 1961-1962 . . . . 12.00 


Single numbers: 


Volumescl: and Ties. Sea ss ae 50 
Volumes II through XIV. . ._.. 1.00 
VolumenccV te te Wao ete a ee ee 1.25 
Voltime XMiwes oes Or i ee 1.50 


The subscription price of MApRONo is $6.00 per year. 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 membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


Corresponding Secretary 
California Botanical Society 
Department of Botany 
University of California, Berkeley 4, California 


An © 


re gga eel 
eS ge ee 
ee eae a 7 


fi 


i | Lie ne ve 


' Mm 
i \ 
%, ts 


| y 


be 

Hobe eysiag OE. 
ie) 
rete 


gs 
aN 
Rc 


oom 


i i ee = ces 
Ppygaree La 
od Ce ae 
‘ IT gs ee 
Sedisorote? 


&, oy 
®. ae iis 
f 


f 


fF 


ul i e 
‘Mt Ps 


nS fe WabaAY A ae 
BB rey Te 
Seah cH? NE 


I 
! 


il 


: 


“3 
wa Ses, 
iy 


nAE- Eee pp 
Loran 


gs 
ae ait 


Hee Coen sei: 
eee oS A 
ik 2 aa #, 
a o i 
a 
nY 


witty, ae re 
neces 
Sictore”” 


i. 

* 
fete < ‘ 
PerCure bya “Aly ~ optgoat 
Soe ae see 
Fear by OR ee au =f 
Beacr SP UEeey all ete 


gor 
Le 


cia 
re 
fete Of 


[eee 


? 


is 
fy 4 a 
of 


| 


Aa) 
ae 
_ 


Jnl 


V 


| 


| 


iad 
%. 
i 


Tere, 
/ eal = 4 
Pp ’ ill wi 


| 


a lal 
a ee 
ee S 
Se ae Se ads | 


sf 
a 


RE 


v3 
TA 
HI: 


oy 

psa 
Fag! 

She 


Ws of 


{ 


SE, 
i 
oe 


ac 


0. 
ah 


| 


i iit 
t 1 
f 


,M 
s | 


i cen LAN oo ae a S ‘ 
. oe io in, 


\y ne 
‘umn had 


aE i: “yo a A 


ly % 3 
mE » 


cts 


” GES: 


ee 
A [e 


“ah E. Is » 


% | \ > j nS i 
oy, “ab 


‘an: 
"ff "i | 


eee Me ene 


wr ae 
oN 
med 
“t 
mero 


ce 
oer 
" Ane ee 


Ney 


g eas 
4 c rs cae 
Ve ae oe 
ia Sy 


ie 


f ti: re - os il a 

ng ihe uP iz ( ? i, er % +h : | i | 

ay Sy ys My 
oe < i Py - 1a 


om, 0