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MADRONO. 


A WEST AMERICAN JOURNAL OF BOTANY 


VOLUME XVI 
1961-1962 


BOARD OF EDITORS 


HERBERT L. Mason, University of California, Berkeley, Chairman 
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 
MiLpreD E. MATurtas, University of California, Los Angeles 
MARION OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 


Tra L. Wiccrns, Stanford University, Stanford, California 


SECRETARY, EDITORIAL BOARD 


ANNETTA CARTER, Department of Botany, University of California, Berkeley 


BUSINESS MANAGER AND TREASURER 
Joun H. Tuomas, Division of Systematic Biology, 


Stanford University, Stanford, California 


Published quarterly by the 


California Botanical Society, Inc. 
2016 Life Sciences Building, University of California, Berkeley 


Printed by Gillick Press, Berkeley, California 


Marion Cave photograph 


To Hersert Louis Mason. Upon the approaching retirement from 
your official duties, we, the California Botanical Society, wish to dedicate 
the sixteenth volume of Madrono to you. Our dedication is not only an 
expression of our appreciation for your wise counsel and devotion in your 
editorial capacity which, with the not inconsiderable help of two succes- 
sive secretaries of the Editorial Board, has helped to maintain our Journal 
at a consistently high degree of excellence, but also in recognition of your 
far-reaching and often provocative contributions to Western Botany... 
notably in the fields of paleobotany, ecology, taxonomy and in the basic 
philosophical approaches to botanical problems. 

During your nearly forty years of association with the University of 
California at Berkeley, an ever-widening circle of botanical students, who 
have since taken their places in leading universities throughout the world, 
has benefited by your wise and pioneering tutelage. Their number could 
be easily computed but the influence, through them, which you have 
exerted, is incalculable. On another plane, you have long been a staunch 
defender of our natural heritage. ““A Flora of the Marshes of California” 
is concrete evidence of your and your students’ support of the Wildlife 
Restoration Project. Your many other activities on behalf of conservation 
have furthered the preservation of our fast-disappearing natural areas. 
We hope you will persevere in the good work. 


CONTENTS 
PAGE 
Frontispiece: Herbert Louis Mason 
Edward Palmer’s visit to Guadalupe Island, Mexico, in 1875—S. F. Blake.......... 1 
Vegetation history of the Pacific Coast States and the “central” significance of 

thesKilamathe Region——K. Hi) WHttlQREr. -- efccccs..vccseio-ca nee ccedeecesceccccecananeelteaeeees 5 
Germination of Ceanothus Seeds—Clarence R. Quick and Alice S. Quick .......- eee 23 
ING@teSm andi ING W Sate =. sere eee. 52 Se ee eid an ess hee cee ches. 31, 108, 140, 236, 269 
Clathraceae in California—Wm. Bridge Cooke and George Nyland............--....------ 33 
Foliar xeromorphy of certain geophytic monocotyledons—Baki Kasapligil............ 43 
DER ERY TENGE set ot eh ne 70, 138, 171, 204, 268 
The genus Lepidium in Canada—Gerald A. Mulligan..................22...c22000000000000000000* rhe 
Eschscholzia covillei Greene, a tetraploid species from the Mojave Desert— 

ION SAD GUD TPS GING LTT eee EEE OEE ite OE REPT ARES SP ete OR eee, DORE Tee 91 
Abnormal fruits and seeds in Arceuthobium—Frank G. Hawksworth.................... 96 
MorAlibert WW. ©. Werre—ly a. Wig cin: 2.2 ccacieccxscecccesnse ose esetessescencde-cecesanegatastes 102 
Chromosome counts in the genus Mimulus (Scrophulariaceae)— 

Bare BeVukheriee and Robert K. VicROVy,, IP .a...c..cnc20c0-cincc<-cccteocceceeeacvncesese 104 
Sphenophyllum nymanensis sp. nov. from the Upper Pennsylvanian— 

2d CN OTR ATTSS 1 Mp ok aR aR RR PE 1 Ds Ee ES OE RE ee 106 
A new name in the algal genus Phormidium—Francis Drouet .............2.......--2000----- 108 


Evolution of the Galium multiflorum complex in western North America. 
I. Diploids and polyploids in this dioecious group—Friedrich Ehrendorfer..... 109 
A new species of Lycium in Nevada—Cornelius H. Muller .......2....22....10c20000-0000000++ 122 
Some recent observations on Ponderosa, Jeffrey and Washoe pines in North- 
eastern; Calmornlia—J ion Re aher: sc... 5. es ik fede aed ecm wh nates 126 


Influence of temperature and other factors on Ceanothus megacarpus seed 
Bermination——Hlmer Burton FLO ey cocoon cc ceccccsct ee occ ccce cesses ence ene ede Senne 132 


Chromosome counts in the section Simiolus of the genus Mimulus (Scrophu- 
lariaceae). V. The chromosomal homologies of M. guttatus and its allied 
species and varieties—Barid B. Mukherjee and Robert K. Vickery, J?............. 141 


Milo S. Baker (1868-1961)—Herbert L. MdSON .2..........21...211ccc2200cc00veeeeeseeeeceeeeeseeeeee 155 
Cytological observations on Adiantum xX tracyi C. C. Hall— 

YIP TEI TL VUES PVC I ao) Tena de eae gs re ONC cn | Oe ee 158 
Taxonomic and nomenclatural notes on Platydesma (Hawaii) and a new name 

for a Melicope (Solomon Islands)—Benjamin C. Stone........2-...22....c.000000000---- 161 
A new species of Galium in California—Lauramay T. DembPsStet............2...2-.0-------- 166 
A new species of Crytantha (section Circumscissae) from California and two 

recombinations (section Circumscissae and section Angustifoliae) — 

KIO MVC NER nd Peter Tl TRO UCI tase soc oee ee os es reve ee ect n es eeen ease eee eee 168 
The Santa Lucia Cupressus sargentii groves and their associated northern 

hydrophilous and endemic species—Clare B. Hardham...................2.0..000-000-0-+- 173 


California botanical explorers. XII. John Milton Bigelow— 
VALUE DS IE LI FOCI S ONE ea eA ore yak He cues Pi trae, i vepazwes g pieces Goteeesseceh ees te eek betas ak eee pet 179 


A subarborescent new Eriodictyon (Hydrophyllaceae) from San Luis Obispo 


County, ‘California 112 pb Vn Wielis oe 22 ae ee ee 184 
A new species of Quercus from Baja California, Mexico—Cornelius H. Muller.... 186 
Parasitism in. Pedicularis--Ekiizabeth F (Sprague... ee 192 
Nomenclature, life histories, and records of North American Uredinales— 

George B. Cumnmms and John W. Baster™ 2 eee 201 
An anatomical study of the secondary tissues in roots and stems of Umbellularia 

california Nutt. and Laurus nobilis L.—Baki Kasa pligtl.....0....2.......10cc-200c000000+ 205 
Rufus Davis Alderson (1858-1932 )—Reid M Of GN. ......-....---.0cc---0esseecceereeeesensecennsecceees 224 
The occurrence of new Arctic-Alpine species in the Beartooth Mountains, 

Wvoming-Montana—-Pizkp DL. JOnnSOn eee 229 
The unique morphology of the spines of an armed ragweed, Ambrosia bryantii 

(Gompositae)—=W tdlard W: -Payne. 22 eee ee ee 238 
Factors influencing survival and growth of a seedling population of Arbutus 

menziesii in California—-J ohn Peltom 22-28 cece ae 2377, 
A new species of Downingia—J ohn H.W ether... 12:22. -ncccc-cceceenendensenccnsnecansenseestecceesees 256 
Three new species related to Malacothrix clevelandii— 

Wiliam S.. Davis-and Peter Hs Ravens. 2 ee 258 
Documented chromosome numbers of plants...........0......0...222ccc22sceeeeeeeeeeeeeeeeeeeeeeeseeeee 266 
Title a2 cose scale cs wae eae 273 

ERRATA 


Page 96, line 14: for Madrofio 9 read Madrofio 11. 
Page 97: transfer figure to p. 99. 

Page 99: transfer figure to p. 97. 

Page 107, line 5: for Sphenopyllum read Sphenphyllum. 


Page 120, line 8: for GALIUM ROTHROCKII Gray subsp. ROTHROCKII read GALIUM 
WRIGHTII Gray subsp. ROTHROCKII (Gray) Ehrend. 


Page 121, line 38: for eu-ployploid read eu-polyploid. 

Page 149, next to last line: for n=25 read n—=24. 

Page 167, legend: after Fig. 1, add Galium hardhamae. 

Page 201, running head: for UREDINALIS read UREDINALES. 
Page 203, running head: for UREDINALIS read UREDINALES. 


Bot. 


\e) 
at 
VOLUME 16, NUMBER | JANUARY, 1961 
Contents 
PAGE 


EDWARD PALMER’S VISIT TO GUADALUPE ISLAND, 
Mexico, IN 1875, S. F. Blake 1 


VEGETATION HISTORY OF THE PACIFIC COAST STATES 
AND THE “‘CENTRAL” SIGNIFICANCE OF THE 
KLAMATH REGION, R. H. Whittaker 5 


GERMINATION OF CEANOTHUS SEEDS, Clarence R. 
Quick and Alice S. Quick 23 


NoTES AND NEws: THE DISCOVERY OF THE LICHEN 
PARMELIOPSIS PLACORODIA IN WESTERN NORTH 
AmERICcA, William L. Culberson; OBSERVATIONS ON 
ARCEUTHOBIUM VAGINATUM IN Mexico, Frank G. 
Hawksworth, ADDITIONS TO THE AQUATIC FLORA 
oF Arizona, Charles T. Mason, Jr. and Richard H. 
Hevly; NoTE TO MEMBERS 31 


A WEST AMERICAN JOURNAL OF BOTANY 


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 


HERBERT L. MAson, University of California, Berkeley, Chairman 
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 
MItprepD E. MaArtuias, University of California, Los Angeles 24 
MaArIon OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
TrA L. Wiccrns, Stanford University, Stanford, California 


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


Business Manager and Treasurer—JoHN H. THomaAs 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Baki Kasapligil, Department of Biology, Mills College, California. 
First Vice-president: Lawrence R. Heckard, Department of Botany, University of 
California, Berkeley, California, Second Vice-president: Kenton L. Chambers, De- 
partment of Botany, Oregon State College, Corvallis, Oregon. Recording Secretary, 
Mary L. Bowerman, Department of Botany, University of California, Berkeley, Cali- 
fornia. Corresponding Secretary: Wallace R. Ernst (January-June), Lauramay 
Dempster (July-December), Department of Botany, University of California, Berke- 
ley, California. Treasurer: John H. Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


EDWARD PALMER’S VISIT TO GUADALUPE ISLAND, 
MEXICO, IN 1875 


S. F. BLake! 


From the standpoint of the discovery of new forms of birds, one of the 
most important short expeditions in the history of North American orni- 
thology (in the sense of the American Ornithologists’ Union Checklist) 
was that made by Dr. Edward Palmer to Guadalupe Island off the coast 
of Baja California in the spring of 1875. In addition, his plant collections 
formed the first scientific botanical records known from the island and 
they give the best approximation of the vegetation there before intro- 
duced goats had done extensive damage. 

Palmer (18312-1911), at the time a man of about 44, had already been 
active off and on for a score of years, beginning in 1853, making collec- 
tions in most branches of biology and ethnology in various parts of the 
United States, northern Mexico, and Paraguay, principally for the Smith- 
sonian Institution. Further details of Palmer’s life and work are discussed 
by McVaugh (1956) and in a paper read by Safford (1911) at the meet- 
ing of the Botanical Society of Washington on 10 January 1911 to cele- 
brate Dr. Palmer’s (supposed) 80th birthday, only a few months before 
his death. 

Among the archives of the New Crops Research Branch (formerly 
Division of the Plant Exploration and Introduction) of the Agricultural 
Research Service, United States Department of Agriculture, are fourteen 
envelopes of manuscript material relating to Palmer’s work from 1853 to 
1911, twelve envelopes containing field books and copied data covering 
the years 1902-1910, and an unpublished manuscript in twelve envelopes 
by the late William E. Safford (1859-1926), a former botanist in the 
Department, dealing especially with Palmer’s work as a plant collector. 
All of this material was drawn upon by Dr. McVaugh in writing the book 
on Palmer, but unfortunately space limitations prevented him from in- 
cluding many quotations. Two items relating to Palmer’s first collecting 
trip to Guadalupe Island have so much human interest in connection with 
this first scientific expedition to that island that they deserve to be put on 
permanent record. One is an eight page manuscript by Palmer; the other 
a letter from Robert Ridgway to Palmer. The manuscript tells of Palmer’s 
disagreeable experiences after his collections had been made, owing to 
shortness of food and the failure of the promised boat to come and take 


1 EpiTor’s NOTE: Dr. S. F. Blake passed away on December 31, 1959. His manu- 
script had been prepared in the form of a note; its posthumous publication in another 
form necessitated transferring parenthetical citations to “Literature Cited” and mak- 
ing other minor alterations in the introductory material. In addition, it seemed per- 
tinent to incorporate information about Harry Bye Stewart and the items from the 
San Diego Union, all of which were contributed by Dr. Reid Moran of the San Diego 
Natural History Museum. 


Maprono, Vol. 16, No. 1, pp. 1-32, January 20, 1961. 


SMIHSUNIA 
werntiinion JAN 3 8 9G 


2 MADRONO [Vol. 16 


them off. The eight pages, written on only one side and measuring 22.3 
14.5 cm., appear to have been copied from Palmer’s notes after his return 
to San Diego and not entered from day to day upon the island. Safford 
evidently planned to use the manuscript in his projected publication (he 
refers to it in his published biographical sketch) and had corrected in 
blue crayon Palmer’s frequently faulty spelling and grammar, but the 
journal is here printed in the original spelling, punctuation, and capital- 
ization, with the addition in brackets of a few words that are needed for 
clarity, and the indication by ‘‘(sic)” of the principal misspellings. 

Palmer sailed from San Diego on 30 January 1875 aboard the ‘‘San 
Diego” (San Diego Union, 30 January 1875) and may well have arrived 
at Guadalupe Island by the first of February. With him was his assistant 
for the trip, Harry Bye Stewart (1862-1922), the twelve year old son of 
Wm. W. Stewart, San Diego shipping agent. As related in the journal fol- 
lowing, they were not taken from the island until nearly four months had 
passed. Palmer’s second trip to Guadalupe Island in 1889 was briefer 
and less harrowing. 


Palmer’s journal follows. 


Guadalupe Island. Lower Cal. Dissapointment (sic). I miss the Alaska Expe- 
dition. 1875. As arrangements had been made to send over a schooner after me 
in 6 weeks and no boat came, I became anxious & I went daily to Pt. Lookout 
to watch for it. All provisions had given out but goat meat & coffee & beens 
(sic) that had been on the island for years. 

The young man [Safford’s note: Harry Stewart] who had accompanied me, 
bore up well until the bread gave out, when he said he wanted to go home. 
I was kept busy carrying collections on my back a foot (sic), to the beach in a 
cave. These journeys were very tiresome, & all but one load was stored when 
I became sick. 


April 30—it was [thought that] a schooner was [sighted] in the distance 
but, it was not,—a cruel dissapointment (sic) as I had expected to join a party 
for Alaska May 1st.—I had hurried my collections so as to [be] ready in/[p. 2 | 
6 weeks & now to be kept back, together with physical weakness made me so 
sick that for some days [I] could with difficulty move about. 


May 11—with much difficulty reached the Point, but no schooner. Coffee all 
gone & nothing to eat but old beans & goat meat. I ate mustard leaves which 
toned up the stomach somewhat. 


May 14—all hands sick, including myself in bed for 3 days, with violent dis- 
order of the bowels—& the fleas nearly ate me up & the flies by day, nearly 
worried the life out of me. As I could no longer [go] to point disappointment, 
asked Mr. Sanford [Safford’s note: an old sailor, who was in charge of the 
island], to go. he was just able to be about. he said if Jack [the burro] came 
up he would go, so the yard gate was opened, for this faithful old /[p. 3] Jack 
was very fond of bones, they where [were] always thrown in a pile for him, 
he could crunch them lik [like] a dog, this Jackass had carried not only myself, 
but several others over the Island, was a great favorite, he being worthy of a 
title named him Saint John, which name he went by. but he soon came along 
and while devouring his bones, the saddle was put on, and to [1.e., the] journey 


1961] BLAKE: PALMER 3 


to the point made but no boat. May the 15th—the gate left open Saint John 
entered [and] while at the usual bone pile was saddled by Mr. Sanford, who 
rode to the look out, he returned and reported seeing an object like a boat 
approaching shore,/| p. 4| hopes revived, and the old name of the point restored, 
he went to the landing riding St. John, after much effort [I] rolled out of bed, 
and dressed, was sitting by the bed putting the remainder of my specimens the 
best I could together in bundles to carry them down to the landing, when Mr. 
Sanford returned with a man from the boat. he said he had come for me and 
my companion, the agents son Mr. W. Stewart, He said seeing the bundles, 
but you are not going to carry all that are you, yes I must try was the reply, 
they are birds, and valuable. no he said you are not able, and I doubt if/[p.5 | 
you can walk to the beach. yes I must try, for the mules must be left for to- 
morrow. the sick men must go also. they are no use hear (sic), no medicine or 
food. The man took my choicest bundles on his back, and little by little, with 
his help started for the beach, leaving the rest with the baggage to come next 
morning, nearing the beach, and the boat was by a sudden puff of wind carried 
out to sea, my feeling at this sight was indiscribable (sic) it was near dark 
before she came again to anchor. It was a hard task to make the Journey to 
the beach, owing to my feebleness,/|p.6| after getting on board, my first re- 
quest was after bread and tea, a small slice of bread was toasted a little, butter 
spread over it with a cup of tea, was given me and I fell over a sleep (sic) to 
awaken next morning late. The party from the Island came, things and men 
were put on board, they opened my Cave on shore, its choice contents brought 
on board. The Superintendent and one of the ablest men was left on the Island, 
all the provisions that could be spared from the boat, was given them, with 
the promise that more should spedily (sic) be sent them, when we /[p.7] bid 
them, the Island, and the faithful St. John, farwel (sic), and pushed off, food 
and drink was given to us sparingly, Reached San Diego, Thursday night 
may 20, 1875, weighing 125 pounds, going on the Island, weighing one hun- 
dred and sixty. 

The cause of the long detention on the Island, was owing to the inability of 
W. W. Steward [Stewart ],—with whom arrangements had been made to send 
a boat in six weeks, to dispatch one from San Diego before, as no boat smaller 
than mail steamer, entered the harbor of San Diego, during that time, if a boat 
was kept at the/[p.8] Island, the hands might escape, and when the want of 
food, was made known by Telegraph to the President of the goat company, it 
turned out he had unintentianally (sic) kept the memorandum of Provisions 
in his pocket, instead of sending them on the Island at the time of my visit,— 
The distressed condition of those from the Island,—awoke the simpathy (sic) 
of the company, A new president was elected, and six months provisions with 
some new men was promptly sent on the Island. 


In the San Diego Union for May 21, 1875, is the following: “The 
schooner Coso arrived last evening from Guadalupe Island to W. W. 
Stewart & Co., bringing up as passengers Dr. Palmer, of the Smithsonian 
Institution, Master Harry Stewart, and two of the men employed on the 
island. Dr. Palmer has been engaged in collecting specimens of natural 
history for several weeks past. There has been some sickness among the 
men on the island, the supply of flour having given out some weeks ago, 
and an exclusively meat diet having proved unwholesome. The schooner 


4 MADRONO [Vol. 16 


left the men all the flour she could spare, and more will be forwarded 
immediately.” 

The second item of interest is a holograph letter from Robert Ridgway, 
then twenty-five years old, requesting Palmer to use his influence with 
S. F. Baird, Secretary of the Smithsonian Institution, to have Palmer’s 
birds turned over to him rather than to Dr. Elliott Coues for identifica- 
tion. History shows that this was done. 


Ridgway’s letter to Palmer runs as follows: 


Smithsonian Instn. 
Washington, D.C. 
Nov. 14, 1875 
Dear Doctor: 


I have just returned from a months absence at my old home in Illinois, and 
embrace my first opportunity to answer your favor of the 16th of October. 
On visiting the Smithsonian yesterday I found your birds in Dr. Coues’ hands, 
but informed Professor Baird that you had requested me to work them up, and 
strongly urged my claim to the first right. Please write to him yourself at the 
earliest moment, regarding this matter, as I would like to work up all your 
collections in the bird line—will do it with pleasure, and in a manner which I 
am sure will meet your approval in all respects. 

In looking over your collection I was astonished to find apparently every 
species an entirely new one [last two words not italicized |—most of them very 
distinct from any previously known, while Dr. C., was not aware of any differ- 
ence whatever until I informed him. I will be glad to have all the information 
you can possibly give me regarding each species of these birds, and also full 
notes upon the geographical location, geological formation, natural productions 
of all kinds (particularly the flora and sylva, since these influence so much the 
distribution of the birds) so that I will have material for an elaborate paper 
—in which you shall have full, and entire credit. 

I am now making a hobby of big trees; and if you can supply me with any 
newspaper scraps, or original notes, on large trees of any part of the world, 
I will accept them most gratefully; the smallest items will be thankfully re- 
ceived. During the course of your explorations and ramblings you must have 
come across many ‘‘monarchs of the forest”—particularly in the tropics. And 
you probably have stored up much information, both general and detailed, in 
this branch. 

Let me hear from you, and believe me, in haste Yours truly Robert Ridgway 


Dr. Edward Palmer 


St. George, Utah Crops Research Division, 
United States Deparment of Agriculture, 
Beltsville, Maryland. 


LITERATURE CITED 


McVaucH, R. 1956. Edward Palmer, plant explorer of the American West. Univ. 
Oklahoma Press, Norman. 430 p. 

Ripcway, R. 1876. Ornithology of Guadeloupe Island, based on notes and collec- 
tions by Dr. Edward Palmer. Bull. U.S. Geological and Geographical Survey 
of the Territories 2:183-195. 

SAFFoRD, W. E. 1911. Edward Palmer. Popular Sci. Monthly 78:341-354. 


Ut 


1961] WHITTAKER: KLAMATH REGION 


VEGETATION HISTORY OF THE PACIFIC COAST STATES 
AND THE “CENTRAL” SIGNIFICANCE OF THE 
KLAMATH REGION? 


R. H. WHITTAKER 


INTRODUCTION 


Among the major developments in the biogeography of the eastern 
United States were three interrelated ideas: (1) the close floristic relation 
between the eastern United States and eastern Asia (Gray 1846, 1873, 
Li 1952), (2) the central relation of the southern Appalachians to eastern 
vegetation (Adams 1902), and (3) the significance of the mixed meso- 
phytic forests in eastern forest history (Braun 1947, 1950). It is through 
vegetation history that these points take on their meanings in relation to 
one another. During the Tertiary, in Oligocene and Miocene time, Arcto- 
Tertiary forests occupied much of the present temperate latitudes of 
Eurasia and North America. Although these forests differed from one 
area to another, there was floristic exchange between the two continents 
across the Bering land bridge, and the forests showed general floristic 
similarity around the Northern Hemisphere. With the increasingly dry 
climates and glaciation of later time, these forests were increasingly frag- 
mented, restricted, and modified, while other, more dry-adapted types of 
vegetation spread through the interiors of Eurasia and North America. 
Remnants of the Arcto-Tertiary forests exist now on opposite sides of the 
two continents—in the eastern United States and eastern Asia, and (with 
fewer surviving genera) in the western United States and western Europe. 
In the southern Appalachians especially, mixed mesophytic forests occur 
which are suggestive of the Arcto-Tertiary forests, have a ‘“‘central”’ rela- 
tion to the forest floras of other parts of the eastern United States, and 
have strong floristic affinities with forests of eastern Asia. 

It is natural to seek a comparable center for western forests. An exten- 
sive area of old and geologically complex mountains, the Klamath Ranges, 
extends from the southern end of the Cascade Range and the northern 
end of the Great Valley of California, west to the Pacific Ocean. One may 
observe of these mountains that: (1) The area has, like the Southern 
Appalachians, one of the most highly complex vegetation patterns in 
North America (Whittaker 1956, 1960). Into this area extend and meet 
in a complexly interdigitating pattern, various types of vegetation which 
form the prevailing climaxes of other areas. All western plant formations 
dominated by trees occur in the Klamath Region, as in no other area. 
(2) Those forest formations which are of most highly mixed tree-stratum 
composition and are thought most to resemble Arcto-Tertiary forests in 


1 A contribution from the Department of Zoology, Washington State University, 
and the Department of Biology, Brooklyn College. The author’s studies in the Klam- 
ath Mountains were supported in part by the funds for medical and biological research 
of the State of Washington Initiative Measure No. 171. The author is indebted to 
R. W. Chaney, D. I. Axelrod, and H. D. MacGinitie for comments on the manuscript. 


6 MADRONO [Vol. 16 


the West, occur in this region—the redwood forests and mixed evergreen 
forests. Of these the mixed evergreen forest is the link between two major 
fractions of western forest vegetation—the coniferous forests, and the 
sclerophyll and oak-pine woodland grouping. (3) The Klamath Region 
has also an exceedingly rich flora for its latitude; it is a center of floristic 
diversity and narrow endemism (Jepson 1923-25, 1935, Mason 1927, 
Peck 1941, Detling 1948b), and many plant genera have maximum 
numbers of species in the West, including endemics, occuring there. One 
may, with certain qualifications to be observed, regard the Klamath Re- 
gion as a “center” for the western forests. 

The prevailing climax at low elevations over much of the Klamath 
Region is the Mixed Evergreen Forest (Munz & Keck 1949, 1950, 1959, 
cf. Cooper 1922, Clark 1937)—mixed forests with two-level canopies of 
larger evergreen-needleleaf or coniferous trees (Pseudotsuga menziesii, 
Pinus lambertiana, Chamaecvparis lawsoniana, etc.) and smaller ever- 
green-broadleaf or sclerophyllous trees (Lithocarpus densiflora, Arbutus 
menziesu, Castanopsis chrysophylla, Quercus chrysolepis, Umbellularia 
californica, etc.), with deciduous trees (Acer macrophyllum and A. circt- 
natum, Cornus nuttallu, Corylus californica, Quercus kelloggu, etc.) , usu- 
ally present also. In relation to moisture the canopy changes, from mesic 
stands in which the coniferous stratum is dense and deciduous trees may 
outnumber sclerophylls, through stands in which the conifers are scat- 
tered in open growth above a dense sclerophyll stratum, to more xeric 
stands in which both strata are open and pines (P. lambertiana, P. pon- 
derosa) rather than Pseudotsuga are principal conifers. 


The complex vegetation of the Klamath Region may be conceived in 
terms of these mixed evergreen forests as the central, prevailing climax 
or vegetational matrix for the region, giving way to other types of com- 
munities in various ways (Whittaker 1960). (1) Within the main area 
of the mixed evergeen forests, distinctive communities of different compo- 
sition and structure occur on serpentine and other special parent materi- 
als. (2) Toward more humid environments nearer the Coast the density 
of the conifers increases while that of the sclerophylls decreases, and the 
mixed evergreen forests gradate into Pseudotsuga forests. These in turn 
gradate into coastal Sequoia forests in which sclerophylls are represented 
by small numbers of stems. (3) Toward the north and higher elevations 
the sclerophylls decline, and the mixed evergreen forests gradate into 
montane forests dominated by Pseudotsuga, Abies concolor, and Pinus 
ponderosa. Toward still higher elevations these gradate into subalpine for- 
ests dominated by Abies procera, Tsuga mertensiana, and (locally) Pinus 
monticola and Picea breweriana. (4) Toward the drier interior, Pseudot- 
suga declines and the sclerophyll strata become more open, and the mixed 
evergreen forests gradate into northern oak woodland (Quercus kelloggit, 
QO. garryana) in Oregon, pine-oak foothill woodland (Pinus sabiniana, 
Quercus douglasi, O. agrifolia, etc.) in California. (5) Toward the south, 
the mixed evergreen forests narrow toward the coast (to become part of 


1961] WHITTAKER: KLAMATH REGION 7 


the ‘redwood border” vegetation of the California Coast Ranges), and in 
drier climates gradate into broad-sclerophyll forests and these into 
chaparral. 

It will be the object of this paper to consider the vegetation history of 
the Pacific Coast states with special reference to the Klamath Region 
and two questions—the origin of this vegetation pattern and the central 
relation of this region to the western forests. 


VEGETATION HISTORY 


Some aspects of vegetation history bearing on the Klamath Region 
have been summarized by Chaney (1936, 1938a, 1938c, 1940, 1947, 
1948a) and Axelrod (1940a, 1950c, 1952, 1958, 1959). Geological history 
bearing on the story has been summarized by Diller (1894, 1902), Her- 
shey (1903), Smith & Packard (1919), Clark (1921), Willis (1925), 
Fenneman (1931), Smith (1933), Reed (1933), Weaver (1937), and 
Williams (1948). 

Pre-Cenozoic (Jurassic and Cretaceous) floras of the Klamath Region 
and Oregon Coast Ranges are described by Fontaine (1905a, 1905b, 
1905c) and Chaney (1948a). Forest trees of more modern types became 
widespread in Cretaceous time; Cretaceous floras include almost all the 
families of the subtropical Eocene floras of the West, as represented by 
the Goshen flora (Chaney & Sanborn 1933). Much of the area of Cali- 
fornia and Oregon, inland to the Sierra Nevada and Blue Mountains, was 
submerged in the Cretaceous; the Klamath Region itself formed an exten- 
sive, mountainous island which later was probably reduced by erosion 
and subsidence to an archipelago of scattered islands (Diller 1894, Con- 
don 1902, 1910, Smith & Packard 1919, Smith 1933, Reed 1933). To- 
ward the close of the Cretaceous the Klamath Region was raised above 
the sea again. 

In Eocene time the full land surface of the Klamath Region was occu- 
pied by vegetation as it has been (except for local alpine glaciation) 
through the whole of Cenozoic time since. With continued evolution of 
modern plant types, extinction of archaic ones, and probable climatic 
warming accompanied by migrations toward the north, subtropical for- 
ests of essentially modern types appeared in the United States in the 
Eocene, as the Wilcox and other floras of the East (Berry 1916, 1930, 
1937), the Goshen and other floras of the West (Chaney & Sanborn 1933, 
Chaney 1936, 1938c, 1947). During the Eocene epoch, the Oregon coast 
north from the Klamath Mountains was submerged (Weaver 1937), as 
was the Great Valley and much of California west of the Sierra Nevada 
(Clark 1921, Reed 1933). The Oregon and California Coast Ranges were 
not yet formed, although submarine volcanic activity on an immense scale 
was producing the lavas later to become the core of the Coast Ranges 
of Oregon and Washington (Williams 1948). The Cascade Mountains 
were not yet elevated to intercept the moisture of maritime air masses. 
Most of Oregon was a broad plain, across which mesophytic forests ex- 


8 MADRONO [Vol. 16 


tended from the coast to the John Day Basin area of eastern Oregon 
(Clarno flora, Knowlton 1902, Chaney 1938c, 1948a) and beyond. The 
high temperatures of the Eocene permitted subtropical floras to extend 
northward to about 50° north latitude on the coast (Chaney 1947), and 
some elements of these forests to extend as far as 56—57° in Alaska (Hol- 
lick 1936, Chaney 1949). 


Mesophytic subtropical forests, representing the Neotropical-Tertiary 
Geoflora, appear in fossil floras from widely separated points in the Pacific 
Coast states—from the California Sierra Nevada (Chalk Bluff and La 
Porte floras, MacGinitie 1941, Potbury 1935), through western Oregon 
(Comstock and Goshen floras, Sanborn 1935, Chaney & Sanborn 1933, 
Chaney 1936, 1948a) to the Puget floras of Washington (Newberry 1898, 
Chaney 1947). Physiognomically, such forests were dominated by trees 
with leaves of subtropical types—of moderate size, thick and probably 
evergreen texture, mostly entire margins, and in many cases elongate tips; 
floristically the Lauraceae (Cinnamomum, Persea, Ocotea, Neolitsea, 
Cryptocarya, Lindera, Nectandra) predominated along with Ficus, Ano- 
na, Meliosma, Magnolia, and other subtropical or tropical forms. Such 
subtropical forests doubtless prevailed in the lowlands of the Klamath 
Region, There is little indication of the upland forests of that time; but 
it may be presumed that temperate forests, probably including such gen- 
era as Sequoia, Pseudotsuga, and Abies, Alnus, Lithocarpus, and Ulmus 
occurred there (Chaney 1936, 1938a, 1938c) and were related to the tem- 
perate forests which then existed far to the north in Alaska (Hollick 1936, 
Chaney 1938a, 1947). 

Much of western California, Oregon, and Washington was submerged 
during the Oligocene, but the submergence was less extensive in the lands 
adjacent to the Klamath Region (Clark 1921, Reed 1933, Weaver 1937). 
Volcanic activity in the area of the Cascade Mountains, which had begun 
in the Eocene, continued in the Oligocene to form a belt of scattered 
mountains which were still not effective as a climatic barrier (Williams 
1948). In the Klamath Region itself a major uplift believed to have 
occurred at the close of the Eocene (Diller 1902) initiated the long cycle 
of erosion which was to produce the Klamath peneplain. With lower tem- 
peratures in the Oligocene, subtropical forests were displaced to the south, 
while the temperate forests were shifted southward and downward. In the 
western states Metasequoia and other temperate forms which had oc- 
curred in Alaska entered lowland forests along with Sequoia and other 
forms which had occurred on the uplands of the West during the Eocene 
(Chaney 1936, 1947, 1951). Through a wide area of the West there oc- 
curred forests which may be broadly characterized as redwood-mixed, 
dominated by either evergreen or deciduous redwood (Sequozta or Meta- 
sequoia, see Chaney 1948b, 1951) mixed with a variety of deciduous and 
some evergreen broad-leaved trees. These temperate forests of the Eocene 
in the Far North and the Oligocene and Miocene in the United States, 
taken in the broad sense and with allowance for the regional and topo- 


1961] WHITTAKER: KLAMATH REGION 9 


graphic differentiation within them, represent the Arcto-Tertiary Geo- 
flora (Chaney 1947, 1959). The transition between the Arcto-Tertiary 
and subtropical forests of the West was apparently represented in moist 
lowlands by warm-temperate forests in which Taxodium was dominant 
rather than the redwoods, with Nyssa as a major broad-leaved form 
among a mixture of subtropical and temperate forms—forests suggestive 
of the swamp forests in warm-temperate eastern North America of the 
present. 

Lowland forests of this sort, dominated by Taxodium and Nyssa and 
including forms of both temperate (Metasequoia, Juglans, Salix, Quercus, 
Platanus, Tilia, Ulmus) and tropical (Ocotea, Lindera, Persea, Ficus) 
affinities are represented in the Klamath Region by the Oligocene Weaver- 
ville flora (MacGinitie 1937). Although these do not represent the up- 
land forests, they imply the prevalence of temperate forests over most 
of the land surface of the Klamath Region from that time on. Far east 
from this, the Florissant flora (MacGinitie 1953) occurred in the area of 
the Colorado Front Range; this flora also included forms of subtropical 
affinities but was predominantly temperate in character. Sequoia, Cham- 
aecvparis, Fagopsis, and Zelkova are believed to have occurred along 
streams and on moist bottom-lands, broadleaf forests with many forms 
now represented in forest-grassland transitions of the eastern and south- 
western states in sites of intermediate moisture conditions, and pine wood- 
land with evergreen oak and chaparral forms on drier uplands. Species of 
Picea, Abies, and Acer in the flora are believed to represent mountain 
forests of higher elevations. As observed by MacGinitie (1953, p.52), the 
low-elevation pattern from mesophytic streamside forest to pine-oak 
woodland is suggestive of vegetation patterns now existing in parts of the 
Klamath Region. A related complex pattern ranging from mesic forest 
with Zelkova, Cercidiphyllum, and Fagopsis through prevailing deciduous 
forest to dry-slope communities with pines, sclerophyll oaks and xeric 
shrubs, with coniferous mountain forests also represented, is described by 
Becker (1956) from the Ruby River Basin of Montana. 


Temperate forests of the upper Oligocene are represented in the Bridge 
Creek flora of the John Day and Crooked River basins (Knowlton 1902, 
Chaney 1924, 1925a, 1927, 1938c), forests of redwoods (Metasequoia) 
mixed with many other species, the living relatives of which occur in the 
West (Tsuga, Abies, Taxus, Lithocarpus, Quercus, Acer, Alnus, Cornus, 
Fraxinus, Philadelphus, Rhamnus), and in forests of eastern North Amer- 
ica and eastern Asia (Carpinus, Castanea, Fagus, Liquidambar, Nyssa, 
Ostrya, Platanus, Tilia, Ulmus, Cercidiphyllum). Forests of this type, but 
with Sequoia rather than Metasequoia, are represented in the Klamath 
Mountains by the Ashland flora (Chaney 1938c). Although no Oligocene 
fossils of upland forests are available for the Klamath Region, the Ash- 
land, Florissant, and Bridge Creek floras together suggest a probable veg- 
etation pattern: mesophytic forests of mixed needle-leaved evergreen 
(Sequoia, Chamaecyparis, etc.), broad-leaved evergreen (Quercus, Litho- 


10 MADRONO [Vol. 16 


carpus, etc.), and deciduous trees, giving way toward higher elevations 
to cool-temperate forests including A dies and Picea. In the coastal climate 
the pattern would be more strongly mesophytic, with less contrast of the 
extremes of the moisture gradient, than the Florissant pattern. Allowing 
for a warmer and more humid climate than at present, and the extinction 
of some early-Cenozoic forms, especially among deciduous trees, this pat- 
tern would be not unlike that now occurring in the more humid Klamath 
Mountains near the coast. 

With continued cooling of climates from Oligocene through and beyond 
Miocene time, the Neotropical-Tertiary flora almost wholly disappeared 
from most of the United States, though certain members of predominant- 
ly tropical and subtropical families became adapted to life in temperate 
forests and remain as remnants of the Eocene forests (Chaney 1944b, 
1947). In the earlier Miocene, the belt of Oregon now occupied by the 
Coast Range, and additional lands to the west of it, were above sea level 
(Weaver 1937). Warm-temperate forests including forms of subtropical 
affinities extended north on this coastal plain through and beyond the 
Klamath Region, in a manner comparable to that of the vegetation of the 
coastal plain of the eastern United States today. In floras from Rujada 
and Cascadia, in west-central Oregon (Chaney 1938c, 1948a), forms of 
the redwood forests (Sequoia, Lithocarpus, Alnus, Berberis) and decidu- 
ous trees now extinct in the West (Tilia, Castanea, Ulmus, Carya) occur 
together with subtropical Persea, Ocotea, and Sabalites. In the Klamath 
Region itself, the long-continued Klamath erosion cycle (Diller 1902) 
reduced much of the land to a peneplain of gentle or moderate relief. 
Scattered, low mountain ranges, which later became the monadnock sum- 
mits of the major mountain groups of the region, rose locally 1000 meters 
or more above the peneplain. It may be presumed that inland from the 
coastal plain the Klamath lowlands continued to be occupied by red- 
wood-mixed forests, while mountain forests occurred at higher elevations. 


Through later Miocene time, the widespread Arcto-Tertiary forests 
were affected by increasingly dry climates. Great lava flows successively 
destroyed existing vegetation in the interior of Oregon and Washington 
in Miocene and later time (Williams 1948), and formed land surfaces 
which were occupied by new and more dry-adapted vegetation. In the 
Mascall flora of the John Day Basin, and related floras widely distributed 
from California to Washington and Oregon (Knowlton 1902, Chaney 
1925b, 1948a, 1959), mixed forests with Taxodium and redwoods ap- 
peared. The reduction of the redwoods and other mesophytic forms in 
these suggests, however, a climate drier than that of the Bridge Creek flora 
(Knowlton 1902, Chaney 1925b, 1938c, 1948a, Axelrod 1940a). Resem- 
blance of these forests to the redwood-border forests was emphasized in 
earlier accounts (Chaney 1925b, 1938c, Oliver 1934). The redwood in 
question was the deciduous Metasequoia, however; and the oaks were 
predominantly species with larger, dissected leaves resembling many of 
those now in the deciduous forests of the eastern states, O. kelloggi and 


1961] WHITTAKER: KLAMATH REGION 11 


other deciduous western oaks (Knowlton 1902). The Mascall flora of the 
John Day Basin was thus a predominantly deciduous forest adapted to 
still relatively humid, but increasingly continental climates, of eastern 
Oregon (Chaney 1948a). Forms of subtropical affinities in the Mascall 
flora and the Latah flora of eastern Washington (Chaney 1938c, 1938a, 
Knowlton 1926) suggest continued warmth of climate. Mixed forests in- 
cluding conifers (Sequoia, Abies, Libocedrus, Pseudotsuga, Picea, Thuja), 
sclerophylls (Lithocarpus, Quercus), and deciduous trees occurred at 
Weiser, southwestern Idaho (Dorf 1936). A vegetation pattern including 
mixed sclerophyllous and deciduous trees as the prevailing climax, and a 
montane forest with Abies, Pinus, Pseudotsuga, and Chamaecyparis, is 
suggested by La Motte (1936) for the upper Cedarville flora of north- 
western Nevada and northeastern California. Farther south, vegetation 
more distinctly adapted to drier climates appeared in the sclerophyll for- 
ests of the Tehachapi and Mint Canyon floras (Axelrod 1939, 1940b). 


Changing climates of the later Miocene were thus reflected in geo- 
graphic and topographic shrinkage of the mesophytic, Arcto-Tertiary 
forests. The complement to this process was the spread of dry-adapted 
vegetation types and floras, many forms of which expanded northward 
from centers of origin probably in scattered areas of the Southwest where 
Neotropical-Tertiary plants became adapted to aridity in Cretaceous 
and Paleocene time (Axelrod 1958), other forms of which probably 
evolved from species of temperate forests to occupy cooler dry environ- 
ments as these became increasingly available, some forms of which en- 
tered the North American flora from the dry-climate flora of eastern Asia 
(Babcock & Stebbins 1938). Because of the importance of the spread out 
of the Southwest, and of the Mexican mountains as a center, Axelrod 
(1940a, 1950a, 1950c, 1958) has termed the dry-adapted floras of south- 
western derivation, an even broader grouping than the Arcto-Tertiary, the 
Madro-Tertiary Geoflora. 

At the end of the Miocene, the whole Cascade belt was upheaved by 
folding and tilting (Williams 1948), further desiccating the interior of 
Oregon and Washington. Uplift occurred in the Klamath Region (Diller 
1902, Williams 1948), the Olympic Mountains (Weaver 1937) and the 
Sierra Nevada (Diller 1894, Fenneman 1931), drying the interior farther 
south; and further, major uplift occurred at the end of the Pliocene. In 
Pliocene time most of the coastal belt of Oregon was above the sea, but 
lobes of the sea extended into some areas of California and Washington 
(Clark 1921, Reed 1933, Weaver 1937). Deformations producing the 
California and Oregon Coast Ranges occurred at the beginning and end of 
the Pliocene. Islands off the California coast, the history of which may be 
traced backward through earlier Cenozoic time (Reed 1933), supported 
and permitted the differentiation of the California closed-cone pine flora 
(Mason 1934, Cain 1944). The trend of increasing dryness of climate 
continued through the Pliocene, though with fluctuations toward more 
humid climates during part of the epoch. Axelrod (1944c, 1944d, 1948) 


iy MADRONO [Vol. 16 


suggests climates which were more humid and warmer than at present in 
the lower, drier and warmer than at present in the middle, and cooler and 
moister than at present in the upper Pliocene. 

Some of the floras of lower Pliocene (or upper Miocene) age are meso- 
phytic and warm-temperate in character. Coastal plain vegetation of 
warm climate and moist situations is represented in central California 
(San Pablo or Neroly flora, Condit 1938, Axelrod 1944d), with forests 
including Taxodium, Nyssa, Persea, and Magnolia. The Remington Hill 
flora of the Sierra Nevada (Condit 1944a) and the Troutdale flora of the 
Columbia River Gorge (Chaney 1944a) include Sequoia and Chamae- 
cyparis, together with deciduous and sclerophyllous broad-leaved trees. 
These are the last samples of forests of Arcto-Tertiary type in which 
Sequoia, Chamaecyparis, and other conifers, Umbellularia and other 
sclerophylls, are mixed with a diverse deciduous component including 
many genera now restricted to the eastern United States or eastern Asia. 
Metasequoia had apparently become extinct by the end of the Miocene 
(Chaney 1951). At lower elevations in the area of the Remington Hill 
flora, the Table Mountain flora (Condit 1944b) included more xero- 
phytic woodland and chaparral forms. Eastward from these areas, forests 
of the interior are represented in floras of west-central Nevada (Axelrod 
1956, 1957), and the Alvord Creek flora of southeastern Oregon (Axelrod 
1944e). In the Nevada floras Sierra redwoods (Sequotadendron) occurred 
with other conifers with modern equivalents in the Sierra Nevada and 
Klamath Region on cooler slopes, and chaparral on exposed slopes, in 
vegetation patterns dominated by oak woodlands (Axelrod 1956). At Al- 
vord Creek montane forests of Pseudotsuga, Abies and Pinus on more 
mesic slopes gave way to woodland and chaparral forms on drier slopes. 
Vegetation patterns most nearly resembling this contact of an interior, 
montane derivative of the Arcto-Tertiary forest with Madro-Tertiary 
woodland occur now in the drier, eastern portion of the Klamath Region. 
The Alvord Creek flora suggests the increasing importance of conifers 
other than redwoods (Pseudotsuga, Abies, Picea, Pinus) which were to 
dominate the later forests of the interior. These and other lower Pliocene 
floras represent the latest occurrence over extensive areas of the West 
of forest types and vegetation patterns similar to those now existing in 
the Klamath Region and California Coast Ranges. 


With increasing dryness of middle Pliocene and later time, the more 
strictly mesophytic forms of these forests were eliminated from most or 
all of the western states. Middle Pliocene Mulholland and Petaluma 
floras of west-central California, and the Oakdale from the central Sierra 
(Axelrod 1944a, Dorf 1933, Axelrod 1944b, 1944d) represent oak-wood- 
land communities and reflect the expansion of Madro-Tertiary vegetation. 
The Deschutes and Alturas floras (Chaney 1938b, Axelrod 1944f) of 
northeastern California and eastern Oregon, with Populus, Salix, and 
other riparian forms of semi-arid climates, indicate the elimination of the 
mesophytic forests from the lowland interior east of the Cascades. Wood- 


1961] WHITTAKER: KLAMATH REGION 13 


land, chaparral, grassland, and desert were spreading over much of the 
area formerly occupied by forest (Axelrod 1948, 1950c, 1958). Cooler 
and more humid climates of later Pliocene time are indicated by extension 
into west-central California of more mesophytic forests—the Sonoma, 
Wildcat, and Santa Clara floras (Dorf 1933, Axelrod 1944c, 1944d), in 
which Sequoia was present or dominant, together with sclerophyllous and 
deciduous trees. 

The cooler and drier climates of the Pliocene, accompanied by and in 
part produced by rising mountain ranges along the Pacific Coast, effected 
the replacement of widespread Arcto-Tertiary forests by essentially mod- 
ern vegetation patterns. The mesophytic redwood-mixed forests shrank 
from wide occurrence into a limited area of coastal California and south- 
ern Oregon. Sequoia sempervirens and Chamaecyparis lawsoniana have 
become wholly restricted to this area; the evergreen-broadleaf or sclero- 
phyll component (Umbellularia, Lithocarpus, Castanopsis, Arbutus, and 
Quercus spp.) has become largely restricted to this same coastal belt and 
somewhat less humid climates inland from it. Deciduous components 
were even more strongly affected by increasingly dry summer climates. 
Metasequoia and many broad-leaved deciduous forms are extinct in the 
West; those that have survived have done so by restriction to the same 
areas of humid forests or to mountain forests, by restriction to valleys 
and the vicinity of water-courses in more arid regions, or by such adapta- 
tions to aridity as are indicated by smaller and thicker leaves (Chaney 
1944b). As the range of the mesophytic forests decreased, that of the 
diverse Madro-Tertiary forms increased; and woodlands and other types 
of the less humid West spread as regional climaxes (Axelrod 1948, 1958). 
Floristic differentiation separated the vegetation of southern California 
from that of northern California (Axelrod 1937, 1950b). Floristic differ- 
entiation also separated the forests of the North Pacific Coast, and those 
of the Rocky Mountains and interior ranges, from those of the California 
coastal belt (Axelrod 1940a, 1948, 1950c, Mason 1947), although forms 
now of the North Coast and Rocky Mountains lived with the redwood- 
mixed forests in the California Coast Ranges into middle or upper Plio- 
cene time (Axelrod 1944a, 1944c, 1948), and many of these forms are 
represented in the Klamath Region and Cascade Mountains today. 
Climatic and topographic changes combined to convert mesophytic for- 
est patterns, which changed slowly across great distances of the West, 
into complex and strongly zoned patterns of many plant communities 
closely juxtaposed along steep climatic gradients of the mountains and 
valleys of the Pacific Coast states. 


It is thought that during the Pliocene epoch the vegetation of the 
Klamath Mountains took on essentially its present character. Forests 
related to the present Sequoia and mixed evergreen forests have probably 
existed in this region, with changing distributional relations to elevation 
and topography, through most of Cenozoic time. But it is probably in 
middle Pliocene time that the Sequoia forests, which had occurred at 


14 MADRONO [Vol. 16 


Ashland in the eastern Siskiyou Mountains in Oligocene time, became 
restricted to the coastal belt, while Madro-Tertiary woodland forms en- 
tered the Klamath Region from the south. Thus would result the major 
features of the modern pattern—coastal redwood forests, mixed ever- 
green forest in the central portion of the region, and oak woodland and 
other more xeric types toward its eastern limits. 

California Pleistocene floras (Chaney & Mason 1930, 1933, Potbury 
1932, Mason 1934) represent essentially modern vegetation types. Cli- 
mates distinctly cooler than those of the present are indicated, however, 
by the Willow Creek and Carpinteria floras (Chaney & Mason 1930, 
1933); forests corresponding to the former now occur 600 km. or more 
north along the coast from Santa Cruz Island. The extent to which Pleis- 
tocene climates were cooled and vegetation displaced south of the ice sheet 
in the eastern United States has been debated (Braun 1947, 1950, 1955, 
Potzger & Tharp 1947, Deevey 1949). The combination of fossil forest 
types well south of their present occurrence with glacial topography in 
the higher Klamath Mountains (Hershey 1900, Flint 1957) suggests sub- 
stantial climatic effects accompanying glaciation in this area. Displace- 
ment of the northern limit of the Sequoia forest southward, and expan- 
sion of the montane forests into lower elevations at the expense of the 
mixed evergreen forests, are likely. Cooler climates would also displace 
species which had previously occurred farther north, southward into the 
Klamath Region. In warmer, post-glacial climates, these species could per- 
sist in the area by movement upward in the mountains, as well as north- 
ward. Detling (1954) has observed that the flora of Saddle Mountain in 
the Oregon Coast Range includes a number of boreal relicts resulting 
from this kind of displacement; a number of these have the present south- 
ern limits of their distributions at higher elevations in the Klamath Moun- 
tains. Other Klamath species have distributions suggesting that they are 
relicts from glacial time—notably Chamaecyparis nootkatensis, the 
known southern limit of which is represented by three isolated patches at 
high elevations in the Siskiyou Mountains, two reported by Mason (1941) 
and one found by the author on Preston Peak. 


Retreat of the glaciers was followed by warmer and drier climates until 
the xerothermic period, about 4000-8000 years ago, which was drier and 
warmer than the present (Hansen 1947, Flint 1957). Effects of the drier 
climate were less evident near the coast than in the interior (Hansen 
1947); but vegetational displacements the reverse of those during the 
glacial periods must have occurred in the Klamath area—with movement 
northward of the Sequoia forests and expansion of the mixed evergreen 
forests and still more xeric types relative to forests of more mesic situa- 
tions and higher elevations. Expansion of chaparral over an extensive 
area of California and northward into the Klamath Region probably oc- 
curred also during drier climates of Pleistocene time (Axelrod 1937). 
Relicts of xerothermic vegetation occur on some peaks west of the Cas- 
cades in Oregon, in areas now dominated by mesophytic forests (Detling 


1961] WHITTAKER: KLAMATH REGION TS 


1953). Comparable relicts, including some of the species listed by Det- 
ling, occur in the Klamath Region, especially on serpentine and other 
special parent materials and on drier mountain slopes in the eastern part 
of the region. 


THE CENTRAL RELATION OF THE KLAMATH REGION 


Major points on the origin of the Klamath vegetation pattern are in- 
dicated in the preceding account; they may be summarized: (1) The 
history of the western forests from Miocene time to the present has been 
one of progressive shrinkage toward the coast and higher elevations, ac- 
companied by progressive differentiation in the different areas of the 
West. Within the Klamath Region, mixed forests of Arcto-Tertiary deri- 
vation were modified by extinction of the greater share of their tree spe- 
cies, especially among deciduous forms, and became restricted to more 
humid climates near the coast. (2) From the Arcto-Tertiary forests 
evolved also, with even more severe depletion of tree species, montane and 
subalpine forests adapted to environments which were cooler, or drier, 
or both than those in which the redwood and mixed evergreen forests 
occur. Higher elevations of the Klamath Region are occupied by montane 
and subalpine forests which are in large part similar in derivation and 
character to those of other western mountains, in part distinctive in occur- 
rence of species (Picea brewertana, Quercus sadleriana, Ribes marshalh, 
etc.) endemic to the Klamath Region. (3) An extensive transition of com- 
munities belonging neither simply to Arcto- nor Madro-Tertiary floras 
has probably existed at least since early Tertiary time and has, with the 
evolution and differentiation of its own species and evolution of Arcto- 
and Madro-Tertiary species into and through it, differentiated into vari- 
ous communities occurring between forests and more xeric non-forest 
communities. The mixed evergreen forests and woodlands of the Klamath 
Region have probably such mixed derivation from Arcto- and Madro- 
Tertiary and intermediate floras. Among these communities there is a 
range of climatic variations and probable derivation, from more mesic 
types of mixed evergreen forests which are primarily of Arcto-Tertiary 
derivation, to more xeric pine-oak woodlands primarily of Madro-Tertiary 
derivation. (4) In reciprocal relation to the forest history, dry-adapted 
communities progressively expanded and differentiated in the Southwest 
and interior lowlands, communities predominantly of Madro-Tertiary 
derivation in the south but with increasing representation of forms of 
Arcto-Tertiary or other cooler-climate derivation toward the north. As 
part of this development, woodland, chaparral, and grassland communi- 
ties spread in the drier inland environments of the Klamath Region. 

The thesis has been developed by Braun (1935, 1938, 1947, 1950, 
p. 39, 1955) that the Mixed Mesophytic Association of the Appalachian 
Plateaux is the central, the oldest, and the most complex association of 
the Deciduous Forest Formation, that from the Mixed Mesophytic, or its 
ancestral progenitor, the mixed Tertiary forest, all other climaxes of the 


16 MADRONO [Vol. 16 


deciduous forest have arisen. The corresponding relation of the coastal 
redwood and mixed evergreen forests of the Klamath Region to the west- 
ern forests is suggested, but a number of qualifications on too literal an 
interpretation of this relation should be observed. The Arcto-Tertiary 
forests were not “a community,” but a vegetation pattern with marked 
regional differentiation in dominance and floristic composition, with dif- 
ferentiation also in relation to moisture gradients and presumably other 
local factors. Their species were variously distributed, and widespread 
species probably showed marked ecotypic differentiation then, as today 
(Axelrod 1941). Much of the West has been occupied by vegetation at 
all times; and the effect of climatic change was not to segregate different 
forest types from a single ancestral type, but to cause increasing local dif- 
ferentiation of forests that were already regionally differentiated in the 
Oligocene and Miocene. Many tree species and species-groups have been 
in existence through the whole of the Cenozoic (Stebbins 1950); and 
many of the trees of the West have some history of association with red- 
woods and the Arcto-Tertiary forests (Mason 1947). But the species and 
ecotypic populations that have evolved into the present have been vari- 
ously associated with one another and total community floras variously 
derived from different sources, resulting from diverse patterns of evolu- 
tion and migration in different species and species-complexes. In evolu- 
tionary time species change their patterns of ecotypic differentiation and 
association with other species; and the evolution of communities is reticu- 
late, not simply divaricate (Mason 1936, 1947, Whittaker 1957). If the 
forests of the West are in part derived by differentiation from the red- 
wood-mixed forests, they may also be derived in part from other conifer- 
ous forest communities whose history in the West—in higher elevations 
and drier situations in the mountains—may go back as far as that of the 
redwood-mixed forests. Resemblance of the coastal redwood and mixed 
evergreen forests to the earlier redwood-mixed forests does not imply that 
the former are in any very real sense ancestral to western forests in gen- 
eral. It implies only that, in the progressive shrinkage, species-extinction, 
and regional differentiation of the western forests from the Miocene to 
the present, the largest fraction of Arcto-Tertiary forms, representing all 
three tree growth-forms, survived in the most favorable climate—that 
of the Klamath Region and northern California Coast Ranges. 


One reason for the ‘“‘central” relation of the Klamath Region thus lies 
in geographic and climatic circumstance. It is in this region, as in the 
Southern Appalachians in the East, that a combination of sufficient hu- 
midity and warmth of climate occurs to support mesophytic, mixed for- 
ests which are most like Arcto-Tertiary forests among existing vegetation 
types. Location and climate of the Klamath Region, and the steep climatic 
gradient from the coast inland, are responsible also for much of the vege- 
tational diversity of the region, and for the meeting there of plant com- 
munities of diverse climatic and geographic relations. 

The notable floristic diversity of the region is also in part a conse- 


1961] WHITTAKER: KLAMATH REGION 17 


quence of edaphic diversity. Geological history has resulted in an un- 
usually complex mosaic of parent materials, often with striking effects on 
vegetation and flora (Whittaker 1954, 1960). Parent material contrasts 
also contribute to the meeting of community-types and species of widely 
different geographic relations. Thus at low elevations in the central Sis- 
kiyou Mountains, Chamaecyparis-Pseudotsuga forests with deciduous 
and sclerophyll trees and Northwestern floristic affinities, and Jeffrey pine 
woodlands with Libocedrus decurrens and Arctostaphylos viscida and 
floristic affinities with the montane forests of the Sierra Nevada, occur in 
close proximity—but the former on diorite, the latter on serpentine. Many 
of the numerous species which reach their distributional limits in the 
Klamath Region occur there as localized, “relict” populations on serpen- 
tine, gabbro, or other special parent materials. Greatest numbers of nar- 
rowly endemic species occur on these same parent materials; other narrow 
endemics appear on more “normal” parent materials at high elevations 
and in other special situations. Concentrations of narrowly endemic 
species in the area are thus related to edaphic factors (cf. Mason 1946a, 
1946b) and other environmental extremes (cf. Detling 1948a). 


The Klamath Region thus shares characteristics with other centers of 
floristic diversity and narrow endemism—topographic complexity, edaph- 
ic diversity, and age of land surfaces. The diversity of habitats has been 
characteristic of the area throughout its long history, although climatic 
gradients were probably less steep before middle Pliocene time. Even at 
the maximum development of the Klamath peneplain, mountains of di- 
verse parent materials existed in the area. The region has at all times 
offered a complex mosaic of habitats in which species of diverse environ- 
mental requirements might survive, while submergence, glaciation, cli- 
matic desiccation, and lava flows have affected surrounding areas. From 
the unlimited diversity of the present geographic and probable historic 
relations of species represented in the area, one may recognize such 
major groupings as: (1) widespread western species, and Sierra-Cascade 
species, which extend through the region; (2) formerly more widespread 
species which are now relict endemics or epibiotics in the region; (3) 
species of diverse present distributional relations which extend, from the 
south, the north and the interior, into communities in appropriate cli- 
mates in the region, many of these species being at or near their limits 
of distribution there; (4) species of diverse distributional relations rep- 
resented in the region by localized, “relict” populations on special parent 
materials, at higher elevations, or both; (5) narrowly endemic species of 
diverse origin, many of which may have evolved within the region to 
occupy some part of its complex mosaic of habitats. 

The central relation of the Klamath Region is regarded primarily not 
as one of a center of origin for forests of other parts of the West, but as 
a center toward which mesophytic forests of the past have shrunk, and as 
a center of accumulation of species of varied evolutionary history in the 
diverse habitats of ancient land surfaces. This does not mean, however, 


18 MADRONO [Vol. 16 


that the area has not also been a center of origin of major significance for 
some groups of plants—a reservoir of species populations of diverse envi- 
ronmental adaptations and of genetic diversity within some species and 
species-complexes, from which populations have evolved and migrated 
into other areas. The genus Crepis provides an example (Babcock & Steb- 
bins 1938), with a number of diploid species now relict in the Klamath 
area, while genetic material from these has been used in apomictic poly- 
ploids which have spread over semi-arid environments of the interior. 
The cytogenetics of Crepis further suggest that endemic species shared by 
the Klamath Region and the Tehaman area of the northern Sierra Nevada 
(Jepson 1923-25) have reached the latter from the Klamath Mountains 
(Babcock and Stebbins 1938). 

It may be noted that the two aspects of the central relation of the 
Klamath Region discussed are to some extent separate phenomena. The 
central vegetational relation is a consequence of location and climate, 
primarily because of adjacency to the Coast; the concentration of species 
diversity and endemism is a consequence of climatic and edaphic diver- 
sity and age, primarily because of the mountains inland from the coastal 
belt. A series of criteria for centers were suggested by Adams (1902, 
1909); but these are each subject to limitations and are to some extent 
independent of one another (Cain 1944). “Centers” are conceptual pro- 
ducts of interpretation according to chosen criteria (Whittaker 1956). 
The Klamath Region is by no means the only center for forest vegeta- 
tion and flora in the West. The Sierra Nevada is of comparable antiquity; 
this and other California ranges are rich in species, including narrow 
endemics. A center of maximum development of conifereous forests may 
be located in the Puget Sound area, and the center for the sclerophyll 
complex is well to the south of the Klamath Region. In the complex 
vegetational and floristic pattern of the West there may be no single area 
which has the same degree of ‘‘central”’ significance as the Southern Appa- 
lachians in the East. Yet, when these various allowances are made, it 
remains true that the Klamath Region possesses a central relation to other 
forest areas which is one of the significant features of the biogeography 
of the western United States. 


SUMMARY 


1. The Klamath Region of northwestern California and southwestern 
Oregon is an area of old and geologically complex mountains, supporting 
a complex vegetation pattern and a diverse flora rich in narrowly endemic 
species. The region is a floristic and vegetational ‘‘center” for the forests 
of the western United States. 

2. Vegetation history of the Pacific Coast states since Miocene time 
has involved progressive shrinkage of Arcto-Tertiary forests and progres- 
sive expansion and differentiation of Madro-Tertiary communities. Mixed 
forests (coastal Sequoia and mixed evergreen forests) most nearly related 
to the Arcto-Tertiary forests in the West are now limited to the Klamath 


1961] WHITTAKER: KLAMATH REGION 19 


Region and northern California Coast Ranges, while woodland, chaparral, 
and grassland communities primarily of Madro-Tertiary derivation have 
entered the Klamath Region from the south to form the more xeric part 
of its vegetation pattern. 

3. Floristic diversity of the Klamath Region has resulted from climatic 
and parent-material diversity, together with age of the mountains which 
has permitted species of diverse histories and environmental relations to 
survive there, often as relicts restricted to special parent materials or 


situations. Biology Department, Brooklyn College 


Brooklyn 10, New York 


LITURATURE CITED 


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. 1909. The Coleoptera of Isle Royale, Lake Superior, and their relation to 
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Lake Superior. Mich. Geol. Rept. 1908:157-215. 

AxELrop, D. I. 1937. A Pliocene flora from the Mount Eden beds, southern Cali- 
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. 1939. A Miocene flora from the western border of the Mohave Desert. 
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—_—_——. 1944a. Pliocene floras of California and Oregon. 5. The Mulholland flora. 

Publ. Carnegie Inst. Wash. 553:103-146. 
——. 1944b. Pliocene floras of California and Oregon. 6. The Oakdale flora. 

Publ. Carnegie Inst. Wash. 553:147-165. 
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. 1944d. Pliocene floras of California and Oregon. 8. The Pliocene sequence 
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. 1944f. Pliocene floras of California and Oregon. 10. The Alturas flora. 
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. 1948. Climate and evolution in western North America during Middle 
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. 1950c. Studies in late Tertiary paleobotany. VI. Evolution of desert vege- 
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20 MADRONO [Vol. 16 


—. 1959. Geological history. In A California flora, by P. A. Munz & D. D. 
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Bascock, E. B. anp G. L. STEBBINS, JR. 1938. The American species of Crepis; their 
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Cain, S.A. 1944. Foundations of plant geography. Harper, New York & London. 
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. 1938b. The Deschutes flora of eastern Oregon. Publ. Carnegie Inst. Wash. 

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—. 1938c. Ancient forests of Oregon; a study of earth history in western 

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. 1940. Tertiary forests and continental history. Bull. Geol. Soc. Am. 51: 

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. 1944a. Pliocene floras of California and Oregon. 12. The Troutdale flora. 

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. 1944b. Pliocene floras of Califoria and Oregon. 13. Summary and conclu- 

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. 1948b. The bearing of the living Metasequoia on problems of Tertiary 
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. 1949. Early Tertiary ecotones in western North America. Proc. Natl. Acad. 
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—-———. 1951. A revision of fossil Sequoia and Taxodium in western North America 
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1961] WHITTAKER: KLAMATH REGION 21 


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CuaneEy, R. W. and H.L. Mason. 1930. A Pleistocene flora from Santa Cruz Island, 
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. 1933. A Pleistocene flora from the asphalt deposits at Carpinteria, Cali- 
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CuaneEy, R.W. and Etruer I. SANBorN. 1933. The Goshen flora of west-central 
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Crark, B. L. 1921. The marine Tertiary of the West Coast of the United States; its 
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Criark, H.W. 1937. Association types in the north coast ranges of California. Ecol- 
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Conpit, C. 1938. The San Pablo flora of west-central California. Publ. Carnegie 
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. 1944a. Pliocene floras of California and Oregon. 2. The Remington Hill 
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DeEEVEY, E. S., Jr. 1949. Biogeography of the Pleistocene. Bull. Geol. Soc. Am. 60: 
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—. 1953. Relict islands of xeric flora west of the Cascade Mountains in Oregon. 

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22 MADRONO [Vol. 16 


Hansen, H. P. 1947. Postglacial forest succession, climate, and chronology in the 
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1961] QUICK: CEANOTHUS 23 


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36:641-678. 


GERMINATION OF CEANOTHUS SEEDS! 


CLARENCE R. QUICK AND ALICE S. QUICK 


The many species of ceanothus in California are ecologically diverse, 
but they occur most frequently and most abundantly on relatively arid 
sites and where repeated wildfire has been a determinant of vegetative 
composition. Because of prompt and abundant seedling regeneration after 
fire (Quick, 1959) and because many or all ceanothus species have nitro- 
gen-fixing nodules on their roots (Quick, 1944), the genus is an important 
factor in development and conservation of high-quality soil profiles under 
wildland vegetation. In order to survive fire, seeds “stored” in duff and 
topsoil must lie in relatively well-insulated positions and be quite obdu- 
rate to heat (Quick, 1956). 

Most ceanothus seeds will not imbibe water and ‘‘plump”’ in the labora- 
tory unless they are first subjected to some type of heat treatment (Quick, 
1935), or to seed-coat scarification. Unplumped seeds cannot germinate 
because they are dry. Seeds of montane species of ceanothus commonly 
will not germinate, even if thoroughly plumped, unless an embryo dor- 
mancy has been obviated by appropriate stratification treatment; i.e., by 
continuously-moist aerated storage for some weeks at temperatures slight- 
ly above freezing. The present paper reports data from experiments aimed 


Facilities for seed storage and culture, stratification and germination were made 
available by the California Forest and Range Experiment Station, United States 
Forest Service, in cooperation with the University of California at Berkeley. 


24 MADRONO [Vol. 16 


TABLE |. EFFECTS OF "STEEP" TYPE HOT-WATER TREATMENT ON GERMINATION 
OF DEERBRUSH (CEANOTHUS INTEGERRIMUS) SEED 


Water Seed sample ,weeks stratified at 2.2° C., and age of seeds (years) 
temperature when cultured. 
degrees C|! Percent of 40 seeds germinating 
Q-Ol5 Q-Ol8 Q-Ol8 Q-087 
14 12 5 14 
Z "%m 2 
70 Sei led GCRS Soe 
75 SaS= AS 50.0 eae 
80 G229 47.5 75.0 100.0 
82.5 == S545 60.0 seme 
85 85.0 87.5 95.0 92.5 
87.5 Fess SS 87.5 aaa 
90 70.0 70.0 67.5 87.5 
95 Snes 70.0 85.0 ose 
Mean 75.8 TAL te 74.1 S)SES) 


'Temperature of one liter of water in container at start of "steep" treatment. 


at defining optimum laboratory methods for inducing germination of seeds 
of snowbrush (Ceanothus cordulatus Kell.) and deerbrush (C. integer- 
rimus H. & A.)” Records of seed longevity of these and other ceanothus 
species are reported also. 

The two separate requirements for germination of ceanothus seed of 
montane species, plumping followed by stratification, are equally impor- 
tant in that either treatment alone is ineffective. Stratification as a pre- 
requisite to germination of seeds of wildland plants is much more fre- 
quently encountered than need for special treatment to obviate seed-coat 
impermeability. However, because seeds must be plumped before they 
can be conditioned for germination by stratification, methods of plump- 
ing will be considered first. 


2 SEED SAMPLES. Seeds used in the experiments were collected from vigorous plants 
when fruits were fully mature. Collections were thoroughly air dried. Seeds were 
then extracted from pods by rubbing as gently as possible between two pieces of 
board. Seeds were sieved, winnowed, recleaned, desiccated over calcium chloride, 
placed in air tight containers, and stored at 2.2°C. (36°F.). 

Collection data for seed lots of Ceanothus cordulatus Kell. follow: Q-155, August 
1937, South Fork Stanislaus River, at ca. 4800 ft. altitude. Q-156, August 1937, 
southwest of Cow Creek Guard Station, Stanislaus National Forest, at ca. 5800 feet 
altitude. Q—239, September 1940, Stinchfield Place, west of Pinecrest, Stanislaus 
National Forest, at ca. 5500 feet altitude. 

Collection data for seed lots of Ceanothus integerrimus H. & A. follow: Q-015, 
collected September 1934, same locality as Q-155. Q-018, July 1935, roadside, state 
highway 49, south of Grass Valley, Nevada County, at ca. 2500 feet altitude. Q-026, 
August 1931, same locality as Q-155. Q-087, September 1934, sub-sample of Q-015 
which passed 12-mesh sieve. Q-113, September 1936, same locality as Q-239. Q-158, 
October 1937, southeast of Cow Creek Guard Station, Stanislaus National Forest, at 
ca. 6300 feet altitude. 


1961 | 


3 


PERCENT GERMINATION 


Q-/55. 4800! 4h a. 
E=53.65-0.37)¢ xX 


40 50 €0 7O 
SECONDS BOILED 


Oo 10 20 30 80 90 


Fic. 1. Boiling water and germination 
of snowbrush (Ceanothus cordulatus) 
seeds. (Seed collection number, altitude 
of collection, and age of seed in months.) 


QUICK: CEANOTHUS 25 


3 


PERCENT GERMINATION 
Be € e 8 os 


10 70 «$0 


20 30 40 50 60 
DAYS STRATIFIED 

Fic. 2. Temperature of stratification 
and germination of deerbrush (Ceano- 
thus integerrimus) seeds, sample Q-018. 
(Seeds were placed in 1 liter of water at 


85°C. [185°F.] and allowed to cool to 
room temperature before culturing.) 


Hot WATER TREATMENT 


Two convenient methods of treating seeds with hot water to increase 
seed-coat permeability have frequently been used. In the ‘“‘steep”’ method, 
seeds are tossed into a measured volume of water at a given temperature 
and left in the water until cooled to room temperature. In the boiling- 
water treatment, seeds are vigorously boiled in water (212°F.) for a given 
length of time. After the allotted period of boiling, the seeds are soused 
in an excess of cold water and then cultured. 

Table 1 reports results of treating four lots of deerbrush seeds in hot 
water by the steep method. The volume of water in all tests of this series 
was one liter (1.06 quarts). Temperature at start of treatment varied 
from 70°C. (158°F.) to 95°C. (203°F.). All cultures were of 40 seeds, 
and all were stratified at 2.2°C. (36°F.) after the hot-water treatment. 
The steep treatment obviously is satisfactory for removing impermeabil- 
ity of deerbrush seed-coats. 

Boiling water also will condition ceanothus seeds for successful strati- 
fication and germination (Quick, 1935). A treatment somewhere between 
a few seconds and perhaps 10 minutes might be expected to be the opti- 
mum period in boiling water. This optimum treatment could be expected 
to vary with different species, and with differences in maturity, age, and 
condition of seed of a single species. 

Figure 1 presents data resulting from treatment of three samples of 
snowbrush seed (C. cordulatus) in boiling water. The objective was to 
determine if optimal treatment with boiling water lay between 5 and 90 


26 MADRONO [Vol. 16 


z a 

35° 

% 

= 

= 

o 

Stok C.C. 
Q-155 

= 4800' 

® 41 Mos. 

io 


X=SECONDS Bowed 
BE ACTUVAL DATA, PLOTTED Points isfaae 
Lo E2S359 = 0.37 K TT Eee a 


--e- FE 279.76 - 28.18 Loa X es 
—e—E=62.35- 1.025 X% 40.0072 xe Dee 


OQ Te) 20 30 4-0 50 60 7oO 80 - Go 
SECONDS IN BOILING WATER 


Fic. 3. Statistical generalization of germination data from snowbrush (Ceanothus 
cordulatus) seeds boiled in water, showing curves for straight-line regression, log- 
arithmic regression, and multiple regression. (Seed collection Q—155, 4800 feet alti- 
tude, age 41 months.) 


seconds. Each culture consisted of 100 seeds. Seeds were treated in Berke- 
ley at an altitude of about 125 feet by tossing them into vigorously boiling 
tapwater and by pouring the boiling water and the seeds into excess cold 
water at the end of treatment. Seeds were planted in autoclaved river 
sand, left at room temperature a few days to plump, then stratified at 
2.2°C. for 94 days, and finally germinated in the greenhouse for 5 weeks. 

No obvious optima appear on the graphs of figure 1. The best period 
of treatment in boiling water for sample Q—239 at time of testing may 
have been more than 90 seconds. In contrast, the other two lots appeared 
to have very short optimal periods of boiling. 

Some tests on deerbrush seeds (C. integerrimus) indicate that very 
short periods of boiling will condition only part of the seeds of a sample 
for germination. For example, seeds of collection Q—087 (age 26 months) 
were boiled for various short periods. Final germination was as follows: 
boiled 4 seconds, 20% germination; 8 seconds, 78%; 16 seconds, 63%; 
32 seconds, 68%; and 64 seconds, 73% germination. The four-second 
treatment obviously was too short to be effective on the majority of seeds 
of this two-year-old collection. Likewise seeds of sample Q—026 at age 
of 28 months were boiled for short periods, stratified at 2.2°C. for 102 
days, and germinated in the greenhouse. Germination follows: boiled 30 
seconds, 60% germination; 1 minute, 76%; and 2 minutes, 64% germi- 
nation. Most lots of snowbrush seed seem to be adequately treated by 
shorter periods of boiling than deerbrush seeds. No very short and obvi- 


1961] QUICK: CEANOTHUS pa 


Go 
80 
z 
Zz 2 
£7 t-7o 
< 
g Se 
260 
= = 
1 50 os 
J 
40 B40 
wy 8) 
2 fe 
ee rm 
e 


3 


3 


20 25 30 
BowED 


3 + 6 ‘87 wn oO 5 i) IS 
WEEKS eT AT eee (LOGS.) MINUTES 


Fic. 4. Length of stratification and ger- 
mination of deerbrush (Ceanothus inte- 
gerrimus) seeds. (Seed collection number, 
altitude of collection, and seed age in 
months.) 


Fic. 5. Germination of ceanothus seeds 
after long boiling in water. (Species of 
ceanothus, seed collection number, alti- 
tude of collection, and age of seed in 
months.) 


ously inadequate treatments of snowbrush seeds with boiling water, such 
as described above for deerbrush seeds, have been observed. 


STATISTICAL GENERALIZATION 


When small lots of ceanothus seeds of a single sample are boiled for 
various short periods, then stratified and germinated, a graph of results 
commonly appears to fit a curve rather than a straight line (fig. 1). From 
theoretical considerations of the effects of boiling water on horny seed 
coats, an exponential scale on the time axis of a graph would be expected 
to fit the data better than an arithmetic scale. This generalization can be 
conveniently handled in linear regression analysis by using logarithms of 
time units rather than time units as such (Snedecor, 1938, pp. 308-312). 
Another common method of “fitting” a curve is to add, as a second vari- 
able, the square of the independent variable—in this case the square of 
the time units (Snedecor, 1938, pp. 313-316). 

Figure 3 graphs data resulting from eight cultures of boiled snowbrush 
seed. Plotted first in this figure is the straight-line regression in which 
time of boiling is handled arithmetically as number of seconds. Also on 
the graph is a logarithmic regression, computed by linear regression meth- 
ods, in which time (the independent variable) was the common logarithm 
of seconds boiled. The third line on the graph is the multiple regression 
curve in which two time variables, (1) seconds boiled, and (2) square of 
seconds boiled, were used. 


28 MADRONO [Vol. 16 


The straight-line equation appears to be an oversimplification of the 
data involved. The multiple regression equation in which the square of 
seconds boiled was added as a separate variable is not a valid general- 
ization because it predicts a minimum germination percentage at about 
70 seconds of boiling and a steadily rising germination percentage after 
70 seconds. The logarithmic transformation appears to be the best gen- 
eralization of the three presented and will be used hereafter whenever a 
straight-line relationship is not considered adequate. 


a 


9 


PERCENT STON 


a LS 2 25 3 4+ 5S 6 T8BFH 12 IS 1820 25 
MINUTES BOILED. (LOGARITHMIC SCALE) 


Fic. 6. Additional tests on long-boiled ceanothus seeds. (Species of ceanothus, 
seed collection number, and age of seed in months.) 


STRATIFICATION TIME AND TEMPERATURE 


A reasonably effective temperature and period of stratification must be 
known before conclusions about effects of other variables in germination 
of ceanothus seed can be considered precise. Work reported by Quick 
(1935) indicates one treatment (2.2°C. for 3 months) that seems gen- 
erally effective, but offers no comparison with other time-and-temperature 
combinations. Figure 2 reports results of a series of stratification tests on 
deerbrush seed of sample Q—018. Obviously any one of three stratification 
temperatures will satisfactorily condition water-permeable deerbrush seed 
of this lot for germination. Other experiments have shown that snowbrush 
seeds react similarly, but commonly are best stratified at 2.2°C. or O°C. 
rather than at 5°C. 

Differing severities of hot-water treatment might conceivably change 
the time-and-temperature reactions of ceanothus seeds to subsequent 


1961] QUICK: CEANOTHUS 29 


TABLE 2. GERMINATION OF OLD CEANOTHUS SEEDS 


> <Ww (ep) (ep) ep) Q 
28 =935 $3 23 esas a 83 
Pa e=n ao =a oo o> x3 
5 a8 Z ae ot <2 82 
29 S22 32 Sn $m 38 88 
oa Sa EP 2 9-2. = o 
~~ r oO canes 3 oO 3 = 

= Species of Ceanothus o © 3 S ‘ 
= > 
n 

014 C. arboreus Greene'’® 225 20-5 50 6 none 38 
370 C. arcuatus McMinn 6000 9-5 50. 20 90 66 
155 C. cordulatus Kell. 4800 15-4 100 10 108 Te 
Oe. M 5 800 15-4 100 10 108 87 
7 )2) ae u i 5500 12-3 100 10 108 90 
Sl" : u 5800 13-4 100 20 90 8| 
SS o " y 6600 13-3 100 20 90 86 
056 C. cuneatus (Hook.) Nutt. 2700 iS Keye) 10 98 98 
024 C. divaricatus Nutt. ——- 19-6 100 5 108 46 
183 C. impressus Trel.' 400 ine 25 5 none 88 
O26 C. integerrimus H.8A. 4800 24-4 100 20 930 90 
On 7" Y ss 4800 235 lOO 20 90 100 
Osi" u " 4800 ZN = eye) 20 90 93 
Oley " : 2500 20-5 100 20 90 25 
okey i H 6300 15-3 100 10 108 98 
ZOC ml” a 5500 12-3 100 10 108 on 
O57 C. /Jemmon Parry 2(00 =) 100 10 108 96 
247 C. prostratus Benth. 5800 12-5 80 10 108 100 
028 C. sorediatus H.&A.° 190 22 -| lOO 10 15 85 
Oly. : 800 17-5 100 10 86 96 
025 C. spinosus Nutt.? 200 19-4 66 5 108 iS) 
246 «C. velutinus Dougl. 4300 12-4 100 10 108 be 1s 


' C.arboreus and C. impressus not stratified prior to germination. 
2 Seeds from landscape planting. 
> C. sorediatus ‘stratified at 5°C. (4I°F.), all other species at 2.2°C. (36°F). 


stratification. However, two series of cultures of deerbrush seed, sample 
Q-018, one of seeds boiled for 20 seconds and the other for 70 seconds, 
reacted the same to stratification, insofar as could be told from inspec- 
tion of the data. 

Quick (1935) found that requirements for optimal stratification of 
ceanothus seed apparently varied among the species in relation to the alti- 
tude at which the species commonly grew. Results from series cultures of 
three collections of deerbrush seeds from different altitudes are summar- 
ized 'in figure 4. Deerbrush seeds from lower altitudes appear to respond 
progressively to shorter periods of stratification. 


TOLERANCE TO BOILING WATER 


Immersion in boiling water for 10 to 20, or perhaps 30 seconds will 
satisfactorily condition most ceanothus seeds for germination, if subse- 
quently the seeds are adequately stratified. The limit of tolerance of both 
deerbrush and snowbrush seeds to boiling water was tested in Berkeley 
by individually boiling subsamples from 1 to 20 or 30 minutes. Figure 5 
graphs the results from long-boiling treatments on one lot of deerbrush 
seed, Q-026, and one of snowbrush seed, Q—239. 


30 MADRONO [Vol. 16 


The regression equation for the deerbrush series of treatments is E = 
91.64 — 4.214X% (r = 0.982), where X is simply minutes boiled. This 
equation predicts that on the average 4.2 percent of germination is lost 
for each minute the seeds of lot Q-026 are boiled. The corresponding 
equation for the snowbrush series is E = 108.67 — 3.571X (r=0.989). 
The regression line for deerbrush seeds crosses the time axis—zero ger- 
mination—at 21.75 minutes, and for snowbrush seeds at 30.43 minutes! 
These two tests indicate that snowbrush seeds may be more resistant to 
boiling water than deerbrush seeds. Additional tests of resistance to boil- 
ing water were made. Results are presented in figure 6. 

It is amazing that ceanothus seeds can stand such prolonged periods of 
boiling. The seed coats presumably exclude water from the seed proper; 
the embryo and endosperm are in effect subjected to dry heat for the 
period of the boiling. It is unknown whether death of over-boiled seeds 
is due to the effects of dry heat on embryo or endosperm or to the final 
penetration of boiling water or steam through the seed coats. Effects of 
dry heat on seeds from which coats have been removed have not been 
determined. 

SEED LONGEVITY 


In an ecological sense many ceanothus species are pioneer plants and 
therefore might be suspected of having durable, long-lived seeds. Seeds 
of some species are known to be generally distributed in the duff and soil 
of Sierra Nevada forests (Quick, 1956). Actual germination tests of old 
ceanothus seeds would be of some ecologic interest. Table 2 presents a 
few records of longevity for seeds of known age. 

Many factors may condition results of germination tests on old seeds, 
and high levels of consistency between species and collections, ages and 
individual tests are not necessarily expected. Additional tests will be re- 
quired to define maximum seed life under the pertinent conditions of seed 
collection, handling and storage. The reported data, however, confirm 
the fact that seeds of many ceanothus species are long-lived. 


Berkeley, California 


LITERATURE CITED 


Quick, C. R. 1935. Notes on the germination of ceanothus seeds. Madronio 3:135—140. 
. 1944. Effects of snowbrush on the growth of Sierra gooseberry. Jour. 
Forest. 42:827-832. 
. 1956. Viable seed from the duff and soil of sugar pine forests. Forest Sci. 
2:36—42. 
. 1959. Ceanothus seeds and seedlings on burns. Madrono 15:79-81. 
SNEDECOR, GEORGE W. 1938. Statistical methods. Rev. Ed., xvi + 388 pp., Collegiate 
Press Inc., Ames, Iowa. 


1961] 31 


NOTES AND NEWS 


THE DISCOVERY OF THE LICHEN PARMELIOPSIS PLACORODIA IN WESTERN NorTH 
America.—The foliose lichen Parmeliopsis placorodia (Ach.) Nyl. is a locally common 
epiphyte of pine in the eastern United States. Six years ago I presented a map of 
its distribution from a study of materials from many herbaria. The species was 
found in twelve states from Maine to North Carolina and northwestward to Michi- 
gan and Wisconsin (Culberson, Revue Bryol. Lichénol. 24:334-337. 1955). Many 
new localities in the eastern states have since been found, most of them by Dr. Mason 
E. Hale. These new localities, including those in West Virginia and Kentucky (speci- 
mens at US) where the species had not before been recorded, fall within the pre- 
viously delimited range. 

In a current study of some Parmelia specimens from various herbaria, I found 
a misidentified specimen of Parmeliopsis placorodia from Arizona. It was collected 
in 1946 by Dr. R. A. Darrow, but it was not determined by him. Dr. Hale then sent 
me a 1957 collection from Arizona by Dr. W. A. Weber and Dr. S. Shushan. In the 
spring of 1959, in correspondence about these western specimens, Dr. Weber wrote 
that he and Dr. Shushan had just found the species in Colorado and sent a sample; 
later his student, Mr. R. A. Anderson, also sent me material from South Dakota. The 
known western localities for P. placorodia are then: 

ARIZONA. Santa Cruz County: Santa Rita Mountains, 8,600 feet elevation, Darrow 
4351 (Darrow Herbarium, College Station, Texas; WIS). Cochise County: Chiri- 
cahua Mountains west of Portal, 8,500-10,000 feet elevation, Weber & Shushan 
S8980 (US). Cotorapo. Boulder County: Boulder Canyon, north slope, 8,000 feet 
elevation, Weber & Shushan S17,954 (COLO). SoutH Dakota. Lawrence County: 
Black Hills, vicinity of Roubaix Lake, 5,450 feet elevation, Anderson $20,941 
(DUKE). Pennington County: Black Hills, Rockerville Camp Ground, 4,000 feet 
elevation, Anderson $20,893 (DUKE). 

The habit of the western specimens, all with apothecia, is identical to that of 
specimens from the eastern states. The western specimens also contain the depside 
thamnolic acid identified in microchemical analysis by the presence of typical crystals 
of the aniline condensation product. 

In the five new localities, the species was collected on the bark of Pinus pon- 
derosa Laws. sensu lat. (including var. arizonica). All known epiphytic specimens 
from the eastern United States are likewise from pines, but the species also occurs 
on old fence rails in some places in New England. In the East, Parmeliopsis placorodia 
habitually grows with Cetraria fendleri (Nyl.) Tuck., another North American lichen 
of ecologic amplitude very similar to that of Parmeliopsis placorodia. Cetraria fend- 
leri, however, has been known for some seventy years from pine and “dead wood” in 
New Mexico and Colorado. Although in the eastern states C. fendleri may be some- 
what more broadly distributed than Parmeliopsis placorodia, the high ecologic simi- 
larity and doubtless the similar distributional history of the two species seem to be 
borne out by the western finds reported here. WiL1t1AM L. CULBERSON, Department 
of Botany, Duke University, Durham, North Carolina. 


OBSERVATIONS ON ARCEUTHOBIUM VAGINATUM IN MeExico.—The dwarfmistletoe 
Arceuthobium vaginatum (Willd.) Presl is a common parasite of pines in Guatemala, 
Mexico, and the southwestern and central Rocky Mountain areas of the United 
States. The taxonomic status of the parasite, particularly in the southern parts of its 
range, is unsettled. Gill (Conn. Acad. Arts & Sci. Trans. 32:111-245. 1935) desig- 
nated a northern form on Pinus ponderosa var. scopulorum Engelm. as Arceuthobium 
vaginatum forma cryptopodum (Engelm.) Gill. Gill did not subdivide A. vaginatum 
as it occurs in Mexico, but listed it on Abies religiosa Schl. and Cham., Pinus leiophylla 
Schl. and Cham., and P. hartwegit Lindl. Sosa (Bol. Dept. Forest. y Caza y Pesca 
[Mexico] 4:123-156. 1939) recorded this parasite on Pinus montezumae Lamb. as 
did Kuijt (Bot. Rev. 21:569-626. 1955) for P. tenuifolia Benth. Gill (loc. cit.) also 


32 MADRONO [Vol. 16 


noted the predominantly Mexican form of A. vaginatum on Pinus engelmannii Carr., 
P. leiophylla var. chihuahuana (Engelm.) Shaw, and P. ponderosa var. arizonica 
(Engelm.) Shaw in southern Arizona and southern New Mexico. To this list of hosts 
may be added P. pseudostrobus Lindl., which was parasitized by A. vaginatum 
(Hawksworth 51; March 10, 1956) in Atzimba National Park between Zitacuaro 
and Morelia in the State of Michoacan, Mexico. This parasite probably occurs on 
other Mexican pines as there are several reports of it in the literature that do not 
classify the host species of Pinus. 

Arceuthobium vaginatum is common on Pinus montezumae (Hawksworth 49; 
March 10, 1956) along Highway 15 between Toluca and Zitacuaro in the State of 
Mexico. No infection was seen on P. leiophylla in the stands examined, although this 
tree was closely intermixed with infected P. montezumae. Pinus leiophylla is attacked 
by A. vaginatum elsewhere in Mexico so this suggests the possibility of rather specific 
host preferences among races of this parasite, as was reported by Gill (loc. cit.) for 
A. campylopodum in Western North America. 

In central Mexico Arceuthobium vaginatum is a robust plant with shoots fre- 
quently more than thirty centimeters high. In Arizona and New Mexico, shoots of 
A. vaginaum f. cryptopodum rarely exceed twenty certimeters in length. An addi- 
tional differences is that witches’ brooms caused by the dwarfmistletoe are not nearly 
as conspicuous in Mexican pines as in Pinus ponderosa var. scopulorum in the south- 
western United States. 

The biology and taxonomy of the dwarfmistletoes of Mexican conifers are poorly 
known and present a challenging opportunity for critical study —Franxk G. Hawks- 
worTH, Rocky Mountain Forest and Range Experiment Station, U.S. Forest Service, 
Fort Collins, Colorado. 


ADDITIONS TO THE AQUATIC FLORA OF ARIZONA.—The aquatic flowering plants 
of Arizona have not been adequately studied, consequently it is not surprising to find 
species previously unreported for the state in such habitats. The following three new 
records were obtained during the summer of 1958 while the junior author assisted 
with collecting for the herbarium of the Museum of Northern Arizona (MNA) ; the 
fourth was sent by the collector to the University of Arizona for identification. 

ELATINE CALIFORNICA Gray (Hevly s.n., 17 August 1958, MNA) and Limosella 
aquatica Sesse & Mocino (Hevly s.n., 17 August 1958, ARIZ, MNA) occur in White 
Horse Lake south of Williams, Coconino County, altitude 6500 feet. This lake was 
created by the relatively recent damming of a tributary to Sycamore Creek; it is sug- 
gested that migratory birds making use of this new environmental area may have 
introduced these species. 

POTAMOGETON RICHARDSONII (Benn.) Rydb. was collected at Wheatfields Lake, 
Apache County, on the Arizona-New Mexico boundary (Hevly, Haskell and Deaver 
sn., 23 July 1958, ARIZ, MNA). The introduction of this species might also be at- 
tributed to migratory birds. 

TYPHA ANGUSTIFOLIA L. was collected south of Yuma in the marshes along the 
Colorado River, Yuma County (D. Tuttle s.n., 14 September 1959, ARIZ). Both 
T. latifolia L. and T. domingensis Pers. also occur in the Arizona flora—CuHar_Les T. 
Mason, Jr. and RicuHarp H. HeEvty, University of Arizona, Tucson, Arizona. 


Note TO MemBeErRS.—AIl back numbers of Madrofno are currently available. How- 
ever, some issues are in very short supply. It is therefore suggested that members 
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Secretary so that they may be used for the benefit of the Society. To facilitate 
record-keeping, it is recommended that those members complying with this request 
return issues by book rate not oftener than once a year-—EDITOR. 


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MADRONO 


VOLUME 16, NUMBER 2 APRIL, 1961 


Contents 


PAGE 


CLATHRACEAE IN CALIFORNIA, Wm. Bridge Cooke and 
George Nyland 33 


FOLIAR XEROMORPHY OF CERTAIN GEOPHYTIC MONO- 
COTYLEDONS, Baki Kasapligi 43 


Reviews: Armen Takhtajan, Die Evolution der Angio- 
spermen (Herbert F. Copeland); Amos G. Avery, 
Sophie Satina, and Jacob Rietsema, Blakeslee: The 
Genus Datura (Alton H. Gustafson); C. Leo Hitch- 
cock, Arthur Cronquist, Marion Ownbey and J. W. 
Thompson, Vascular Plants of the Pacific Northwest 
(Robert Ornduff) 70 


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 
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 
MivtprepD E. MATHIAS, University of California, Los Angeles 24 
MARION OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
Tra L. Wiccrns, Stanford University, Stanford, California 


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


Business Manager and Treasurer—JoHN H. THoMAS 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Baki Kasapligil, Department of Biology, Mills College, California. 
First Vice-president: Lawrence R. Heckard, Department of Botany, University of 
California, Berkeley, California. Second Vice-president: Kenton L. Chambers, De- 
partment of Botany, Oregon State College, Corvallis, Oregon. Recording Secretary, 
Mary L. Bowerman, Department of Botany, University of California, Berkeley, Cali- 
fornia. Corresponding Secretary: Wallace R. Ernst (January-June), Lauramay 
Dempster (July-December), Department of Botany, University of California, Berke- 
ley, California. Treasurer: John H. Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1961] COOKE & NYLAND: CLATHRACEAE 33 


CLATHRACEAE IN CALIFORNIA 


Wo. BRIDGE COOKE AND GEORGE NYLAND 


Although the Phallales are found mainly in tropical regions there are 
a few representatives known from temperate areas. Of the two families 
comprising this order of fungi, the Phallaceae are better represented in 
the north temperate zone than are the Clathraceae. Nevertheless, many 
collections have been reported in the United States of some members of 
the latter family. 

The literature on those clathraceous fungi which have been collected 
in North America is very scattered. In eastern North America, Burt 
(1894) described Anthurus borealis from material collected in New 
England, and Murrill (1912) added a report, illustrated with a colored 
plate, of a collection of this species from Blackwell’s Island, New York. 
It is possible that Burt’s species and Murrill’s record were based on 
imported material since it has been shown that Anthurus borealis Burt 
is a synonym of Lysurus sulcatus (Cooke & Massee) G. H. Cunningham, 
a species usually found in Australia. Lysurus texensis Ellis appeared as a 
nomen nudum in a casual collector’s report (Gerard, 1880), and Long 
(1917) later published a description of another collection from Texas 
which he provisionally assigned to L. texensis Ellis. Lloyd’s Synopsis 
(1909) has not been very useful for North American workers. It is a 
compilation in which several species are described briefly and illustrated, 
sometimes poorly; some of these descriptions may prove to represent 
the same taxon. In “The Genera of Fungi” by Clements and Shear (1931), 
both Anthurus and Lysurus are illustrated, the classic illustration of 
Lysurus mokusin credited to Cibot being copied there. Likewise, this 
illustration is used by Gaumann and Dodge (1928) but credited to 
“Cibot and Fischer” rather than to “Cibot in Fischer.’’ Coker and Couch 
(1928) present a description of Anthurus and cite the above-mentioned 
illustrations. 

The structure and development of the fruiting body of Anthurus bore- 
alis were described in considerable detail both by Burt in 1894 and by 
White in 1944. White referred the material that he studied to Lysurus 
sulcatus. A summary of these two studies was made by Rea and Heiden- 
hain (1955) and a comparison of the characteristics of this species with 
those of Lysurus mokusin was made. This publication, based on the exten- 
sive observations made by Rea on specimens from Santa Barbara, Cali- 
fornia, of Lysurus mokusin (Cibot ex Pers.) Fries, represents the first 
detailed study of material collected in the western part of the United 
States. 

Following his retirement from the directorship of the Santa Barbara 
Natural History Museum, Paul Marshall Rea devoted much time to 


Maprono, Vol. 16, No. 2, pp. 33-76. April 18, 1961. 


34 MADRONO [Vol. 16 


mycological studies in the Santa Barbara area. His collections and notes 
are deposited in the Herbarium of the University of Michigan, Ann 
Arbor, where, through the courtesy of Dr. A. H. Smith, the senior author 
had the opportunity of studying Dr. Rea’s collections of phalloid fungi. 
Dr. Rea noted the first specimens of Lyvsurus mokusin in Santa Barbara 
in March, 1941. From that time until the end of his collecting activities 
he acquired, through his own efforts and those of his neighbors, no less 
than 130 collections of this species from the Santa Barbara region. 

The present paper has grown out of a study of recent collections of 
clathraceous fungi which appeared in Bermuda grass |Cynodon dactylon 
(L.) Pers.| lawns in Fresno, Yolo, and Sacramento counties, California, 
particularly after the areas had received an excess of irrigation water. 
For making available to us the specimens collected in their area, we wish 
to thank John Bartels, Agricultural Commissioner’s Office, Woodland, 
California, and K. A. Kimble and H. A. McCain, Department of Plant 
Pathology, University of California at Davis. 

The family Clathraceae includes those phalloid fungi (stinkhorns) 
whose receptacles are stipitate or sessile, clathrate, columnar, or divided 
into several arms, and whose mucilaginous spore mass is borne on the 
interior or exterior of the arms or between the arms. In 1931 Cunningham 
divided the family into three tribes containing eleven genera. The tribe 
Stellateae, the only one with which we are concerned in California, 
included the following four genera as keyed out by him: 


Arms apically organically united or united by a membrane. 
Glebiferous layer composed of irregular pseudoparenchymatous processes 
M ycopharus 
Glebiferous layer consisting of walls of the chambers of the arms . . Anthurus 
Arms apically free, connivent or expanded. 
Arms connivent (usually), attached to the apex of a simple cylindrical 


Cor diuted) stems. 5 -. ee ee SUITES 
Arms attached laterally to a horizontal discoid expansion of the apex of 
the cylindrical stem 9.2: 2 2.04 Se ee Se ee sero” 


The other seven genera of Clathraceae were placed in the tribes Col- 
umnateae and Clathrateae. In a later treatment, Cunningham (1942) 
placed Mycopharus in synonymy with Lysurus; the same organization 
into tribes was retained. Fischer (1933) recognized fifteen genera in the 
Clathraceae, but he did not arrange them into tribes. He recognized the 
four genera listed in the key above, as well as Pseudocolus, which Cun- 
ningham considered to be a synonym of Anthurus. 

Mycopharus was established by Petch as a segregate from Lysurus in 
which the glebiferous surface of the receptacular arms is formed of a 
series of minute shingle-like plates in contrast with that of Lysurus in 
which this surface is merely strongly wrinkled longitudinally, possibly 
as a result of the compact arrangement of the walls of the stipe chambers 
in this portion of the receptacle. In 1931 Cunningham accepted this basis 
for differentiation, but in 1942 he reverted to the earlier position that 


1961] COOKE & NYLAND: CLATHRACEAE 30 


the two genera were synonymous. In this report Cunningham’s 1931 
interpretation is being followed and the two genera are considered distinct. 
With the exception of some of the Santa Barbara material, which 
exhibits characters similar in many respects to those of Mycopharus, the 
California specimens studied to date appear to belong in Lysurus. 


Lysurus Fr. Syst. Myc. 2: 286. 1822. 

Phallus Pers., Syn. Meth. Fung. 245. 1801. pro min. parte. 

Aseroephallus Lepr. & Mont., Ann. Sci. Nat., Bot. III, 4: 360. 1845. 

Anthurus sensu Burt, Mem. Bost. Soc. Nat. Hist. 3: 504. 1894. 

Clathraceae; with an egg-like membranous volva; receptacle com- 
posed of a glebiferous surface raised on a hollow cylindrical or fluted 
stipe; stipe white or tinted near the receptacular arms; receptacular arms 
seated at the top of the stipe, of a more compact tissue, divided into 
4—7 arms, arms separate or organically united, grooved on the outside, 
rounded toward the inside, longitudinally wrinkled, not continuous in 
the center with the hollow portion of the stipe, covered on the rounded 
inner glebiferous surface with the ill-smelling gleba. 

Type. Lysurus mokusin (Cibot ex Pers.) Fr.—only species described 
in original publication of genus. 


Key TO CALIFORNIA SPECIES OF LYSURUS 


Stipe weakly to strongly angular-fluted, upper area red . . . . . L.mokusin 
Stipe cylindrical, upper area brown . . . 2 Se ee BE sulcatus 


LYSURUS MOKUSIN (Cibot ex Pers.) Fries, Syst. Myc. 2:286. 1822. 
Figs. 1, 2. Phallus mokusin Cibot, Nov. Comm. Petrop. 19:373-378, t. 5. 
1775. P. mokusin Cibot ex Pers., Syn. Meth. Fung. 245. 1801. Mutinus 
pentagonus Bailey, Queensland Bot. Bull. 10:35. 1895. Lysurus beau- 
vaisii MOll., Rev. Gen. Bot. 12:61. 1900. Mutinus pentagonus var. hardyi 
Bailey, Queensland Agr. Jour. 16:494. 1906. M. hardy: Bailey, Comp. 
Cat. Queensland Pl. 747. 1910. Lysurus sinensis Lloyd, Myc. Notes 5: 
718. 1917. 

Peridium white, 3-6 (—11) cm. long, with white rhizomorphs at base; 
receptacle 6.5—7.5 cm. long to apex of usually connivent arms; stipe 5 cm. 
long, 4-7 mm. in diameter, narrow at base and expanding upward, white 
to orange-pink below, dark orange-pink to reddish above, internally di- 
vided into hollow chambers, fluted, the flutings 2-3 mm. deep, continuing 
above into midribs of receptacular arms; receptacular arms 4—6, 1.5—2 
cm. long, red, the texture similar to that of stipe, but wrinkled and more 
compact, the tips of arms pointed; gleba brownish in color, becoming 
purplish-black when dry; spores hyaline to pale yellowish, rod-shaped, 
somewhat rounded at ends, (3.8—) 4.5-5 & (1.2—) 1.5—2.0uy. 

California collections examined. 

In Bermuda grass [C yvnodon dactylon (L.) Pers.| lawn, Fresno, Fresno 
County, summer, 1952 (DAV). The stipe in this specimen is hollow, 
5 cm. long, 4 mm. in diameter at the base and expanding upward, and pen- 


36 MADRONO [Vol. 16 


tagonal in cross-section, with the surface flutings 1 cm. from tip to tip of 
the adjoining pairs. The arms are five in number. 

In loam in flowerbed at edge of lawn, Bakersfield, Kern County, April 
11 and 22, 1936, Mrs. A. Ashley and Mrs. F. Hamlin (UC 553876), The 
collectors thought the fungus might have been introduced from Maine 
with some cultivated plants. When first picked, the stipe in the later col- 
lection was orange above, cream below, and the volva was white; a few 
days later, the stipe had become entirely orange, the arms red, and the 
glebal mass madder. The odor was described as that of “‘acetum squill.” 
Too rapid development because of artificial watering caused cracking and 
abnormalities. Receptacles with both five and six arms are represented 
in the collection. In more robust specimens, the receptacular arms may be 
capped by a portion of the volva. The stipes are made up of at least two, 
possibly three, layers of chambers. 


In lawn, Cajon Street, Redlands, San Bernardino County, May, 1944, 
G.J. Hollenberg (UC 695849). When fresh its color was described as pale 
pinkish. The specimen is evidently immature; the spores measure 3.8 
1.2u, and the stipe (the specimen was collected without the peridium) 
measures 4 cm. in length and 6 mm. in diameter at the top in the dry 
condition. 


The collection of a specimen in Sacramento late in 1956 is represented 
only by a colored illustration (fig. 1) made by Norma O’Neil of Sacra- 
mento. This illustration, which was sent by the Agricultural Extension 
Office in Sacramento to the University of California at Davis for deter- 
mination, clearly represents a sporophore of this species. No herbarium 
specimen is available. 


In garden, Santa Barbara, Santa Barbara County, November 18, 1936, 
Miss Caroline Hazard (UC 568835). Upon the arrival of the specimens at 
Berkeley, Mrs. Vera M. Miller observed that the stipe was not white any- 
where, but pinkish throughout, shading down from a color somewhat 
lighter than that of the arms to an Ibis pink (Maerz and Paul, 1950, Pl. 1, 
B-10) where the stipe went into the volva, to a very delicate pink at the 
lower end of the stipe. The stipe is coarsely chambered above, acuminate 
below, bearing apically the receptacular arms which were united at their 
tips. One of the receptacles in the collection bears four arms, while the 
other has six arms. The glebiferous layer is wrinkled and continuous over 
the unfused area of the arms and the base of the arms above the stipe. 
The collector reported that when a hot day was followed by a cool night, 
the texture of the specimens was crisp, while a cool day followed by a cool 
night resulted in limp receptacles. 


In gardens, Santa Barbara, Santa Barbara County, at least 130 collec- 
tions made between 1941 and 1952 or later by Dr. P. M. Rea or Mrs. Rea 
or by residents who gave the specimens to Dr. and Mrs. Rea, the collec- 
tions all deposited at the Herbarium, University of Michigan, Ann Arbor, 
and dated as follows: March, 1941; June, 1943; April through Novem- 


1961] COOKE & NYLAND: CLATHRACEAE Sit) 


[DEEP RED- PINK RIDGES 

UINTH BLACK ALONG SIDES AT 
TOP OF RIDGES- o siDen 
COVERED WITH WET SUBSTANCE 


PINK TO ALMOST WHITE 
POROUS STEM —-6 SIDED 


+RARK ON TREE ROOT 
(moDESTO ASH) 


* ATTRACTS FLIES- 
UNPLEASANT ODOR GIMILAR TO DECAYED FLESH 


Fic. 1. Lysurus mokusin showing habit and mature receptacle. The gleba is shown 
as occurring between the receptacular arms, not covering their outer surface. Draw- 
ing by Norma O’Neil, Sacramento. 


ber, 1944; May through September, 1945; May, June, October through 
December, 1946; January through March, 1947; October, 1952. A wide 
range of morphological variation is found among these collections as is 
evidenced in the notes Dr. Rea made from fresh material. On the large 


38 MADRONO [Vol. 16 


mass of earlier material, he took voluminous notes, among which are three 
different versions of a paper he was preparing on this species. This ma- 
terial was assembled and developed into a paper for publication by Berta 
Heidenhain (as co-author) at the University of Michigan (1955). The 
report gives a complete description of Lysurus mokusin throughout its 
development from the very young buttons or eggs to mature and senescent 
receptacles. 

The senior author has made the following general observations from 
several representative collections of the above series. Dr. Rea sectioned 
nine peridia and mounted them on black paper. These sections show that 
in the unopened receptacle the gleba appears to lie on the outer surface 
of the receptacular arms and not on the inner surface. The gleba is sepa- 
rated, by plates arising from the center of the backs of the arms, into as 
many units as there are arms. As was pointed out by Rea and Heidenhain 
(1955), the glebal masses actually lie between the arms and are attached 
to their sides, but do not invade the central chamber. The backs, or outer 
surfaces, of the arms are thus free from glebal material. These surfaces 
are concave or flat in mature receptacles. However, it has been observed 
by the present authors that as the arms mature and are raised from the 
volva, the gleba appears to shift in position and surround the receptacular 
arms on all surfaces except the backs of the arms. 

The receptacular arms in the specimens accumulated by Dr. Rea vary 
from those in specimens in which they are completely free through those 
which are connivent, those which are held together by fragments or caps 
of volva material, and those which are organically united by a very small 
bit of tissue to those which are united in such a way that the receptacular 
arms form an expanded structure similar to a Chinese lantern and join 
above the glebiferous surface into an apical spire at least 1.5 cm. long. 
The longest dry receptacle observed in the Rea collections was 11 cm. long. 

Some of the specimens of Lysurus mokusin from Santa Barbara, both 
those of the Reas deposited at the University of Michigan and that of 
Hazard at the University of California (UC 568835), have characteristics 


EXPLANATION OF FIGURES 2-4. 


Fics. 2-4. Development of receptacles in Lysurus. Fic. 2. L. mokusin. Each of the 
four fresh specimens shows a different amount of sterile tissue projecting beyond the 
glebiferous area. Flutings on stipe are apparent and continuous with the outer edge 
of the receptacular arms between which lies the gleba. Previously unpublished photo 
by Paul Marshall Rea, Santa Barbara, California. Courtesy of the Herbarium, Uni- 
versity of Michigan. Fic. 3. L. sulcatus. Cluster of four receptacles in various stages 
of development. Lower left: young receptacle just breaking through the peridium, 
the receptacular arms hold the gleba between them; lower right: receptacle with stipe 
nearly completely elongated, gleba still intact; center: gleba nearly completely re- 
moved; upper left: mature receptacle collapsed after removal of gleba. Note cylin- 
drical stipe. Collected at Woodland, and photographed at University of California, 
Davis, October 1, 1958. Fic 4. L. sulcatus. Single receptacle collected at Woodland, 


September 23, 1958, showing receptacular arms with wrinkled glebiferous surface. 
=_> 


1961] COOKE & NYLAND: CLATHRACEAE 39 


Be i a 


a 


F INGHES 3 


Fics. 2-4. Development of receptacles in Lysurus. 


49 MADRONO [Vol. 16 


which in many respects are similar to those of Mycopharus gardneri 
(Berk.) Petch of Ceylon. This latter species was first placed in Lysurus 
by Berkeley, then in Colus by Fischer. Petch (1919) established for it the 
new genus Pharus, and later, finding the name Pharus to be preoccupied, 
he (1926) renamed the genus Mycopharus, distinguishing it from Lysurus 
on the basis of the type of glebiferous surface on the receptacular arms. 
He illustrated the genus as having the arms slightly separated from one 
another below the glebiferous surface. Although, as stated above, some of 
the characteristics of Mycopharus gardneri, the type and at first the only 
species of this genus, are to be found in some of the Santa Barbara collec- 
tions of Lysurus mokusin, the present authors consider that there are 
sufficient differences between the two species to justify the separation of 
the two genera. They differ primarily in the nature of the glebiferous sur- 
face, which is composed of tightly packed scales of “pseudoparenchyma- 
tous processes” in Mycopharus and of tightly packed wrinkles in Lysurus ; 
in addition, the stipe is weakly fluted in Mycophkarus and strongly fluted 
in Lysurus. Among the many specimens of Lysurus mokusin from the Rea 
collection of Santa Barbara, however, there is a wide variety of stipe sur- 
faces, the stipes varying from deeply fluted or winged to shallowly or 
weakly angular. 


Lysurus suLcatus (Cooke & Massee) G. H. Cunningham, Proc. Linn. 
Soc. N.S. W. 56 (3):189, pl. viil, figs. 3,4. 1931. L. texensis Ellis in W. R. 
Gerard, Bull. Torrey Club 7:30. 1880, nomen nudum. Mutinus sulcatus 
Cke. & Mass., Grev. 17:69. 1889. Lysurus australiensis Cke. & Mass., 
Grev. 18:6. 1889. Anthurus australiensis (Cke & Mass.) Fisch., Denksch. 
Schweiz. nat. Gesell. 33:27. 1893. A. borealis Burt, Mem. Bost. Soc. Nat. 
Hist. 3:504. 1894. Lysurus borealis (Burt) P. Henn., Hedw. 41:167. 1902. 
L. borealis var. klitzingi P. Henn., Hedw. 41:173. 1902. L. tenuis Bailey, 
Comp. Cat. Queensland PI. 745. 1910. L. texensis Ellis (?) in Long, Myco- 
logia 9:271-274. 1917. 

Peridium at dehiscence 2—2.3 cm. tall, 2 cm. in diameter at the widest 
point, rhizomorphic; receptacle 7.5—10.5 cm. long; stipe 6.5—9.0 cm. long, 
tapering downward, 5—6 mm. in diameter below, 10-13 mm. in diameter 
at apex just beneath receptacular arms, cream-colored below to yellowish 
above, composed of one layer of chambers, hollow, apex open, at top of 
stipe a collar on which are produced 5—7 hollow receptacular arms on the 
rounded inner surfaces of which is the glebiferous layer; glebiferous layer 
longitudinally strongly wrinkled, dark olive to blackish; receptacular 
arms orange-buff to buff to tan in color, composed of very compact tissue 
with a different appearance from that of stipe, even in length, 6-15 mm. 
long, 3.5--4 mm. in diameter, tapering to a rather sharp point, more or less 
flattened on outer grooved surface, the outer surface 1-2 mm. wide, 
smooth; spores rod-shaped, with rounded ends, slightly yellowish, 3.8— 
4.3 1.5-1.8u. 

In 1880, Gerard noted that Ellis had received a species of Lysurus from 


1961] COOKE & NYLAND: CLATHRACEAE Al 


Texas which he considered to be new and thus had provisionally named it 
L.texensis. No description was given. In 1917, Long described a Lysurus 
collection from Texas, designating it as Lysurus texensis Ellis (?), since 
this nomen nudum had been applied to an undescribed Lysurus from 
Texas. He stated that his material might be only a red form of Anthurus 
borealis. Lysurus texensis Ellis in Long is placed in synonymy with L. sul- 
catus because Long’s description appears to be based on material of the 
L. sulcatus type rather than of the L. mokusin type. In spite of the inter- 
pretation by Rea and Heidenhain (1955), Long’s description of the stipe 
is interpreted here as indicating that the walls of the chambers in the stipe 
wall are polygonal rather than that the stipe itself is polygonal, fluted or 
winged. Cunningham (1931, 1942) places L. texensis in synonymy with 
L. sulcatum, but he lists only Ellis’ nomen nudum and makes no mention 
of Long’s description. 

California collections examined. 

In Bermuda grass lawn, Woodland, Yolo County, September 23, 1958, 
John Bartels; October 1, 1958, K. A. Kimble (DAV). The specimens 
usually appeared after occasional heavy spray irrigation. They were either 
solitary in occurrence, grouped in small clusters, or disposed in large num- 
bers in the lawn. In one specimen, the receptacle, instead of bearing six 
arms of equal length, bore three arms 1.7 cm. long which alternated with 
three shorter arms, 1.4—1.5 cm. long. 

West Sacramento, Yolo County, October, 1958, H. A. McCain (DAV). 

In lawn, Fresno, Fresno County, September 16, 1941, George W. Graves 
(UC 660274). Each of the three somewhat immature receptacles bears 
six arms. The length of the receptacles in the dried condition is 3 cm., 
while that of the arms is 5 mm.; the stipe is white, with no indication of 
the original color, had it been other than white when fresh. 

Santa Barbara, Santa Barbara County (P. M. Rea collection, Univer- 
sity of Michigan, Ann Arbor). Two small specimens are present in this 
collection. 

In flower garden, ‘“‘The Flower Shop,” San Diego, San Diego County, 
September, 1913, A. M. Rainford (specimen in alcohol, deposited by Pro- 
fessor W. A. Setchell in the collection of class demonstration material, 
Botany Department, University of California, Berkeley). According to 
Dr. Lee Bonar, who reported the existence of this specimen to the senior 
author (letter, December 11, 1958, the stipe of this specimen is 6.5 cm. 
long, 2 cm. in diameter at the apex, and 1 cm. in diameter at the base. 
There are five receptacular arms, which are free at their tips. Of the five 
arms, one is shorter than the other four; the short arm is 1.7 cm. in length, 
while the others are 2.1 cm. in length. 


SUMMARY 


Even though members of the Clathraceae are not commonly found in 
California, a large number of specimens of Lysurus have been found in a 


42 MADRONO [Vol. 16 


few isolated localities. The two species of Lysurus that have been reported 
from California, Lysurus mokusin and L. sulcatus, are distinguished pri- 
marily on the basis of color and the nature of the surface of the stipe. Since 
basidia and spores tend to be similar in appearance and size, gross morpho- 
logical features such as color, the shape of the stipe in cross-section, the 
type of glebiferous surface, and the extent of the connection of the apices 
of the receptacular arms serve as the basis for determining the species in 
the genus Lysurus. On the whole, these characters seem to the writers to 
be of importance only at the species level at our present stage of knowl- 
edge of the Phallales. However, the nature of the glebiferous surface of 
the receptacular arms is considered to be of sufficient importance to sepa- 
rate the two genera, Lysurus and Mycopharus. 
Robert A. Taft Sanitary Engineering Center 
Bureau of State Services 
Public Health Service 
U.S. Department of Health, Education, and Welfare 
Cincinnati 26, Ohio 
and 
University of California, Davis 


LITERATURE CITED 


Burt, E. A. 1894. A North American Anthurus—its structure and development. 
Mem. Boston Soc. Nat. Hist. 3:487-505. 

CLEMENTS, F. E., and C. L. SHear. 1931. The genera of fungi. H. W. Wilson Co., 
New York. 

CoKEr, W. C., and J. N. Coucu. 1928. The Gasteromycetes of the eastern United 
States and Canada. University of North Carolina Press, Chapel Hill, North 
Carolina. 

CUNNINGHAM, G. H. 1931. The Gasteromycetes of Australia. XI. The Phallales, 
Part II. Proc. Linn. Soc. N.S.W. 56: 182-200. 

. 1942. The Gasteromycetes of Australia and New Zealand. John McIndoe, 
Dunedin, New Zealand. 

FiscHer, E. 1933. Gasteromyceteae. In Engler, A., and K. Prantl, Die Natiirlichen 
Pflanzenfamilien 7a:1—122. 

GAUMANN, E. A., and C. W. Doncr. 1928. Comparative morphology of fungi. 
McGraw-Hill Co., New York. 

GeRARD, W. R. 1880. Additions to the U. S. Phalloidei. Bull. Torrey Club 7:29-30. 

Lioyp, C. G. 1909. Synopsis of the known phalloids. Myc. Writ. 3: Phal. 11-96. 

Lonc, W. H. 1917. Notes on new or rare species of Gasteromycetes. Mycologia 
9:271-274. 

Maerz, A. J.. and M. R. Pau. 1950. A dictionary of color. McGraw-Hill Co., 
New York. Second Edition. 

Murriz, W. A. 1912. Illustrations of fungi—XI. Mycologia 4:163-169. 

Petcu, T. 1919. Further notes on Colus gardneri (Berk.) Fischer. Trans. Brit. Mycol. 
Soc. 6:121-132. 

. 1926. Mutinus bambusinus (Zoll.) Ed. Fischer. Trans. Brit. Mycol. Soc. 
10:272-282. 

Rea, P. M., and Berta HEmENHAIN. 1955. The genus Lysurus. Pap. Mich. Acad. 
I. 40:49-66. 

Waiter, N. H. 1944. The development of Lysurus sulcatus (Cooke and Massee). 
Trans. Brit. Mycol. Soc. 27:29-34. 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 43 


FOLIAR XEROMORPHY OF CERTAIN 
GEOPHYTIC MONOCOTYLEDONS! 


BAKI KASAPLIGIL 
INTRODUCTION 


Xeromorphy in the vegetative organs of plants and especially in foliage 
leaves has long attracted the interest of morphologists and ecologists. 
There is now considerable information about foliar xeromorphy and its 
relation to factors of the environment (Clements 1905, Harshberger 1908, 
Starr 1912, Hanson 1917, Hayden 1919, Pool 1923, Mothes 1932, Evenari 
1938, Shields 1951a). Most of these studies deal mainly with dicotyledons, 
while the monocotyledons, with the exception of xeric grasses, have 
received less attention. As a matter of fact, dicotyledons are more abund- 
ant than monocotyledons in arid and semi-arid regions; however, certain 
“drought resistant”? monocotyledons do occur widely in xeric habitats. 

Geophytic monocotyledons, i.e., those having subterranean bulbs, 
corms, rhizomes, or tuberous roots, are quite common in the steppic flora 
of central Anatolia. In the vicinity of Ankara, there are over fifty such 
species in the Gramineae, Araceae, Liliaceae, Iridaceae, and Orchidaceae; 
they form about thirty-seven per cent of the total number of known 
species of monocotyledons in the local flora (cf. Krause 1937). In the 
present study the foliar xeromorphy of eighteen of these species, rep- 
resenting five families, has been investigated anatomically. 

The geophytic monocotyledons in central Anatolia produce aerial shoots 
as early as February and complete their flowering and fruiting stages 
during March and April. After forming seeds they usually return to sub- 
terranean dormant stages before the drought period starts. In their habit 
of resting during the dry season, they resemble ephemeral annuals. Some 
botanists may not consider the ephemerals and geophytes as true xero- 
phytes, since both of these groups escape instead of endure the critical 
drought period. This argument may be answered by considering the 
dormant seeds or subterranean fleshy organs as adaptational drought- 
enduring stages in the continuous life cycles of these plants. The geo- 
phytes thus fit the ‘drought evading” class of Shanz (1927). Further- 
more, the geophytes can also be considered as succulent xerophytes since 
they are well adapted to withstand prolonged drought by means of their 
fleshy subterranean organs in which water, carbohydrates, and mucilag- 
inous substances are stored (cf. Warming 1909). 

In this paper, the term “‘xerophyte” is used as a convenient qualitative 
term to designate the plants of xeric habitats where the available soil 
water is limited. The classical concept of this term refers to plants living 
in places with limited water supply and which are equipped with various 
arrangements to reduce transpiration (cf. Willis 1957, p. 692). Although 


1 This paper was supported in part by a faculty research grant from Mills College. 


44 MADRONO [Vol. 16 


Bakee (1914) concluded that xerophytes have higher indices of tran- 
spiring power than mesophytes, recent experimental studies have dem- 
onstrated that there is no direct correlation between structural modifica- 
tions and transpiring powers of xerophytes. The degree of drought resist- 
ance cannot be judged simply according to the xeromorphic features 
(Maximov 1931, Evenari 1938, Weaver and Clements 1938, Shields 1950, 
Daubenmire 1959). 


There is general agreement that xerophytes can by morphological and 
physiological means either escape or resist permanent injuries caused by 
excessive loss of water (Maximov and Maximov 1924), and leaf xero- 
morphy is, in addition to geophytic xeromorphy, another such means. 
Studies on leaf xeromorphy and its relation to physiological and struc- 
tural influences are discussed comprehensively in a recent review article 
by Shields (1950); therefore the present paper will not attempt to give 
a detailed account of the environmental and genetic causes of the foliar 
xeromorphy. However, I should like to point out briefly the factors of 
the steppic environment in central Anatolia that contribute to the struc- 
tural modifications of xeromorphic leaves in the eighteen geophytic mono- 
cotyledons chosen for investigation. In Ankara the atmosphere is very 
dry and induces strong transpiration. The wind velocity is high during 
the winter months when foliage leaves of many geophytic monocotyledons 
develop. The wind increases the rate of transpiration. The light intensity, 
one of the factors contributing to foliar xeromorphy (Shirley 1929, Tur- 
rell 1940, Shields 1951a) is very strong due to high altitude (from 830 m. 
at the railroad station to 1855 m. on the Elmadag mountain), cloudless 
skies, with the exception of a few rainy and snowy days distributed 
irregularly through fall and winter, and to the reflection of solar radia- 
tion from gray steppic soils and calcareous rocks of the mountains which 
are poorly covered by vegetation. The precipitation is low, and the 
drought period usually extends from May to October (Colasan 1946, 
Walter 1955 a and b). The limited water supply of the steppic soils is 
further decreased by direct evaporation from the soil surface, and the 
soil solution acquires the high osmotic pressure which is so typical of 
arid and semi-arid regions. Also, the steppic soils in this habitat are 
extremely deficient in nitrates, which is another important factor con- 
tributing to foliar xeromorphy (cf. Mothes 1932). 


The monocotyledons in this study exhibit certain uniformities in addi- 
tion to their geophytism, reduced leaf surface, and phenological periodic- 
ity. Their root systems are fibrous and very shallow and are limited to 
the uppermost soil layers (Birand 1938, Karamanoglu 1955). The latter 
author points out the anatomic similarities in regard to epidermis, exo- 
dermis, cortical parenchyma and endodermis in the roots of Iris, Orni- 
thogalum and Muscari species, although these tissues show minor dif- 
ferences from one species to another in regard to thickness of tissues. 
Furthermore the geophytic monocotyledons studied by Birand (1938) 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 45 


exhibit low osmotic pressures since they complete their activities above 
ground during the moist season. The present study was initiated pri- 
marily to see if there is any parallelism in the structural modifications 
and in the adaptational degree of foliar tissues among unrelated species 
of geophytic monocotyledons. 

In this study the following widely accepted xeromorphic characters 
have been considered: 


A. General habit of the plant and gross morphology of leaves 
1. Reduction of the surface 
2. Revolute margins 
3. Folding and fusion of the blade 
4. Involution upon wilting 
5. Increased thickness of the blade 
B. Epidermis 
6. Strong cutinization 
7. Increased thickness of the epidermal cell walls 
8. Water storage in depression of stomata 
9. Cutinization and ledge formation on the guard cells 
10. The occurrence of trichomes | 
11. Water storage in the epidermal cells 
C. Mesophyll 
12. Isolateral organization of the palisade 
13. Strongly developed palisade parenchyma at the expense of spongy 
parenchyma 
14. Compactness of the tissues 
15. Decreased size of cells 
16. Development of a water storing tissue 
17. Abundance of excretory idioblastic cells 
D. Vascular system 
18. Strong development of vascular bundles 
19. Compactness of vascular bundles, i.e., lateral proximity of veins 
E. Sclerenchyma 
20. Strongly developed fibers, tracheoid and sclerenchymatous idio- 
blasts (cf. Foster 1956 for these idioblasts). 


MATERIALS AND METHODS 


The leaves were collected from plants growing in different localities 
of the steppe of Ankara. Portions of leaves were fixed in a solution of 
formaldehyde-propionic acid—acetic acid, then transferred into 70 per 
cent ethyl alcohol. Transverse leaf sections were made by free hand and 
stained with safranin for general histological study and mounted in 
gelatin-glycerin. Additional sections were stained with phloroglucinol 
and hydrochloric acid for lignin test. Sudan IV was used for the cuticle 
(Foster 1949). Leaf portions, cleared in 3 per cent sodium hydroxide, 
stained with safranin, dehydrated with xylene and mounted permanently 


46 MADRONO LVol. 16 


in Canada balsam (cf. Kasapligil 1951 for method), were entirely satis- 
factory to observe the orientation of various tissues and their cellular 
morphology. Drawings of leaf sections were prepared by camera lucida. 
Herbarium specimens of the plants were also prepared and distributed 
among the herbaria of the University of Ankara, University of Califor- 
nia, Berkeley, and the Royal Botanic Garden, Edinburgh. 


DESCRIPTIONS OF PLANTS AND THEIR COMPARATIVE FOLIAR ANATOMY 


GRAMINEAE 

HoRDEUM BULBOSUM L. (fig. 1) is a tall perennial grass of cultivated 
lands as well as of the mountains of the steppe. The flat stiff blades are 
characteristically involute upon wilting. The blades are 6-10 mm. wide, 
230-300 microns thick along the median bundle and 150-160 microns 
thick in the lateral portions. The cormous basal nodes are aggregated in 
large clumps. 

Epidermal cells are elongated with undulate cell walls, and alternate 
with short silica cells. Outer cell walls of the epidermal cells are thicker 
than their inner tangential walls. Cutinization of the cell walls is not 
prominent, but the epidermal cells adjacent to sclerenchyma strands on 
both adaxial and abaxial surfaces of the blade have lignified cell walls. 
The stomata are not sunken, and they occur on both surfaces. The guard 
cells are dumb-bell shaped and have lignified cell walls. Each stoma is 
accompanied by a pair of subsidiary cells. Bulliform cells are situated 
along the grooves of the adaxial surface in 4-6 rows. They have straight 
cell walls. Hinge cells underlying the bulliform cells are filled with water, 
contain very few chloroplasts and form one or two layers. Two different 
types of unicellular trichomes were observed: short and hooked tri- 
chomes with thick silicified walls along ridges on the abaxial surface and 
along margins of the blade; long slender trichomes with lignified cell 
walls near the abaxial base of the blades. 

Leaves are dorsiventral and a single layer of palisade parenchyma is 
situated on the abaxial side. Cells of the mesophyll are small and very 
compactly arranged. Prismatic crystals are abundant in the spongy 
parenchyma cells along the bundles. Collateral bundles are surrounded 
by mestome sheath (Schwendener 1890) as well as by bundle sheath. 
Bundle sheath extensions (Esau 1953) are well developed, especially 
along the median bundle of the blade, and connect the vascular bundles 
to the adaxial fibre strands as seen in transverse sections. Vascular 
bundles are close together and are interconnected by small bundles. 
Fibres are more strongly developed on the abaxial sides of the bundles 
than on the adaxial ones. Abaxial fibre strands are connected directly 
with the lignified epidermis and the mestome sheath of the bundles, 
while the adaxial fibre strands are “separated” from the bundles either 
by bundle sheath extensions along large bundles or by spongy parenchyma 
cells along small bundles. Strands of 8-15 fibres extend along both 
margins of the blade, beneath the epidermis. 


Bulliform cells Adaxial epidermis-. ay) 


Hinge cells Spongy parenchyma Aca 
Sclerenchyma Stoma~ ESOS > 
rolensa een Ue ene, 
OBR ES ON COR TEED SAIS es 
Os see ED BossocAs eae 
COS kes \\ ema cok) ae 
Bees eet pea we ive 
Ah) LA OY AEN ASR 
Ye |) AYA ARES Sut Ce Gps 8? mace 
RRIF sow “ ES Ait ai 
g ee are Sees s ee eee 
Sey ees Outer bundle sheath 
Abax. epidermis SOB reece, TH) . Mestome sheath 
Palisade par. Mylene PAR ORO Sclerenchyma 
Phloem SOOO ‘Lignified epidermis 
Adax.epidermis — | 


Spongy par. 
Mestome sheath {= 


Sclerenchyma 
Abax. epidermis 


Median bundle 


lOOy Spongy parenchyma 


—.Adax. epidermis 
ae 
hy / Palisade par. 


Spongy par. 
Laticifer 


Subsidiary cells 


Idioblast 


Bundle sheath 


Laticiferous tube DEO ‘ 
. ; ; Cpig) POT KD 
Adaxial epidermis ae eae fa i, 
xl pidermi ence ags! oes a A 
Spongy parenchyma ( je 3 % 


Bulliform cells 
Median bundle 


Small bundles 
Stomata 


a oy 
fe Nes Gaol 
see 
4 by yy \ 
OK 


Stomata 
Cuticle 


fA 
: 


aN 
‘ 


Or 
Intercell. spaces 


a 


Water stor. par. 


Abaxial epidermis 


Palisade parenchyma 


eee 


O222O 


Fics. 1-4. Transverse sections of leaves: 1, Hordeum bulbosum; 2, Poa bulbosa; 
3, Arum orientale var. elongatum; 4, Allium rotundum. 


48 MADRONO [Vol. 16 


PoA BULBOSA L. (fig. 2) is a small perennial grass which is very com- 
mon in the steppe as well as in the fields and fallow lands. It has short 
and narrow leaves, involute upon wilting. The blades are 2-4 mm. wide 
and 4—7 cm. long. Thickness of the blades ranges from 180—220 microns 
along median bundles and from 60-125 microns in the lateral portions. 
The basal portions of the shoots are swollen and enveloped by thickened 
bases of leaves forming clumps of small bulbs. 

Epidermal cells are long tabular cells with straight walls. Outer walls 
are thicker than the inner ones and are moderately cutinized. The bulli- 
form cells are 5—6 cells wide, situated along two grooves of the adaxial 
surface on two sides of the median bundle. Bulliform cells are not accom- 
panied by hinge cells. The stomata are slightly sunken and occur in both 
surfaces of the blade above assimilatory tissue. Guard cells are dumb- 
bell shaped, and each stoma has two very small subsidiary cells which 
are also sunken. Unicellular hooked trichomes are situated along both 
margins and dorsal ridges of the blade. 

Mesophyll is not differentiated into palisade and spongy parenchyma. 
It consists of very small isodiametric parenchyma cells closely packed 
together. Mesophyll is only two cell layers thick beneath the bulliform 
cells, while its thickness in other portions of the blade is 5—6 cell layers. 
Prismatic crystals are abundant throughout the mesophyll parenchyma. 

The collateral vascular bundles are surrounded by mestome sheaths, 
but differentiated border parenchyma or bundle sheath extensions are 
not present. The median bundle is “imbedded”’ directly within assimila- 
tory tissue and is not connected with the dorsal strand of fibres. On the 
other hand, most of the small bundles are connected to upper and lower 
epidermis by strongly developed fibre strands. Groups of 3-6 fibres 
extend along leaf margins beneath the epidermis. Comparatively speak- 
ing, the vascular bundles in the Poa leaf are more compactly arranged 
than the vascular bundles of the Hordeum leaf, and likewise anasto- 
mosing veins are more frequent than those in the previous species. 


ARACEAE 
As far as this author knows, this family is represented only by one 
species in the steppic flora. 


ARUM ORIENTALE M. Bieb. var. ELONGATUM Engl. (fig. 3) is a herba- 
ceous perennial plant which grows in little soil pockets on eroded and 
exposed limestone or within narrow rock crevices at the highest altitudes 
of the steppic region. Broadly-hastate leaves are 15-25 cm. long. The 
lamina is tender and 250-270 microns thick. The adaxial surface of the 
lamina is darker green than the abaxial one. The plants have rounded 
tubers, slightly adpressed. 

The epidermal cells are polygonal and appear isodiametric in face 
view. They are about 50—70 microns long and 40-50 microns deep in 
transectional view. The epidermal cells along the vascular bundles are 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 49 


narrow tabular in form. A few chloroplasts were observed within the 
epidermal cells. Outer tangential walls of the epidermal cells are cutinized 
moderately and these walls are thicker than the inner tangential and 
radial ones. Stomata are not sunken and appear on both surfaces of the 
lamina, more frequently on the dorsal side. As an average, the guard 
cells are 45 microns long and 15 microns wide, although their sizes are 
variable. The inner and outer ledges of the guard cells are distinctly 
developed. Each stoma is subtended by a pair of lateral subsidiary cells 
which are thick walled. Each subsidiary cell is equipped with a curved 
ledge on its inner tangential wall adjacent to the guard cell. Possibly 
these ledges produce complete closure of stomatal openings and form an 
additional air chamber between the guard cells and the substomatal air 
cavity. Trichomes and idioblasts are absent in the epidermal tissue. 

Mesophyll is dorsiventral and provided with a biseriate palisade par- 
enchyma on the adaxial side. Palisade parenchyma cells are large and 
occupy the upper half of the mesophyll. The spongy parenchyma con- 
sists of 5—6 layers of large irregular and loosely arranged cells. As a 
whole the mesophyll has very conspicuous intercellular spaces presenting 
a mesomorphic rather than a xeromorphic structure. Idioblasts con- 
taining raphides are distributed within the spongy parenchyma. Individ- 
ual raphides are 180-200 microns long and form crowded bundles within 
the idioblasts. Druses are also present, but are not as common as raphides. 
Another feature of the spongy mesophyll is the presence of articulated 
laticifers which were noted also by Solereder and Meyer (1928). Scleren- 
chyma is not present in any form. 

Vascular bundles are reticulate and the leaf resembles a dicotyledo- 
nous leaf in this respect. The veins of first, second, third, and fourth 
orders are closed and form vein islets, while the veins of fifth and sixth 
orders frequently end freely in mesophyll. Marginal bundles run very 
close to the leaf edge. The major veins are surrounded by a vaguely dif- 
ferentiated bundle sheath. 


LILIACEAE 


This family is represented by eight genera in the steppic flora. The 
largest genus, Allium, is represented by ten species occurring in the 
steppic region under a wide variety of conditions, from moist places along 
streams to extreme xeric habitats of saline depressions and mountain 
steppe. Two species, which seem to endure drought longer than other 
species, were studied. 

ALLIUM ROTUNDUM L. (fig. 4) is a tall slender plant reaching a height 
of 40-50 cm. The leaves are filiform, tapering gradually to the tips. 
Although the leaves are typically unifacial in their ontogeny, the adaxial 
surface is flat, the abaxial one ridged (cf. Esau 1953 and Hayward 1938 
for the ontogeny of the leaf of Allium cepa). There is no central cavity 
in the blade. Upon wilting, involution takes place through a folding action 
along the median bundle of the lamina halves. The leaves are 2—3 mm. 


50 MADRONO [Vol. 16 


wide and 280-300 microns thick. The thickness decreases very slightly 
toward the margin of the blade. The small bulbs are oval in outline. 

The epidermis, as seen from the surface, consists of long, tapering 
cells with straight lateral walls. The lower epidermis bears a much heavier 
cuticle than the upper and its outer walls are characterized by lamella- 
tion. The lumina of the lower epidermal cells are extremely small. Bulli- 
form cells of the upper epidermis extend along the groove above the 
median bundle and form a row of 5—7 cells. They are not accompanied by 
any specialized hinge cells in the underlying tissue. Stomata occur on 
both upper and lower surfaces and are sunken below the epidermal level. 
Subsidiary cells are not present, and trichomes in the epidermis are 
missing. 

Mesophyll includes a uniseriate palisade on both sides. The abaxial 
palisade is strongly differentiated, while the adaxial one is weakly devel- 
oped. The spongy mesophyll is composed of small isodiametric paren- 
chyma cells which form 10-15 layers with conspicuous intercellular 
spaces. A well differentiated water-storing parenchyma in the central 
portion of the mesophyll consists of enlarged cells lacking chloroplasts. 
Articulated laticifers and patches of tracheoid idioblasts are abundant 
in the mesophyll and are distributed at random. Well developed vascular 
bundles are densely arranged in longitudinal directions. They are trans- 
connected by means of small bundles. Reversal of the bundles in regard 
to the position of xylem and phloem is obvious. Apparently this situation 
is caused by the peculiarity of the leaf apex (cf. Esau 1953, fig. 16.19). 
Clearly differentiated bundle sheaths have cells characterized by promi- 
nent central vacuoles and peripherally arranged chloroplasts. 

Fibers associated with the vascular bundles are strongly developed, 
the xylem fibers, especially, forming very thick and compact strands along 
major veins. 


ALLIUM STAMINEUM Boiss. (fig. 5) is a small plant about 20-25 cm. 
high with tunicated oval bulbs. The leaves are smaller and more tender 
than those of the previous species. They are 1.5—2 mm. wide and appear 
reniform in transverse section. Involution upon wilting is not pronounced. 
The thickness of the leaves ranges from 450 to 550 microns. Asin Allium 
rotundum, the leaves are unifacial, but both the upper and lower surfaces 
are ridged. The epidermal cells are tabular in form, and their outer walls 
are extremely thick and heavily cutinized throughout both surfaces of the 
leaves. Consequently the lumina of the epidermal cells are very small. 
Stomata occur in deep depressions of abaxial and adaxial surfaces. Uni- 
cellular, short, conical trichomes with very thick cell walls are arranged 
in regular rows along the ridges. Trichomes are living cells, and their 
nuclei are always situated at the bottom of the enlarged basal portions 
of the cells. 

The mesophyll is quite different from that of A. rotundum. First of 
all it has a well developed biseriate palisade on both upper and lower sides, 


_— Trichome 
Lumen 
Nucleus 


4 ays. oe 
TE PNG 
f : bee os 


Palisade parenchyma 


Spongy parenchyma 
Laticiferous tubes —— 


a( Rass 
On: ) 
emesis 


Sx e5¢5 ~~ Bundle sheath 
Eveuna - Laticiferous tubes 


Cuticle 
Adax. epid. 


AN > Se : eee. AC f. 
Palisade par. APY. ve” aie ites oe ‘i : \ aN ses 2S aes 


Ky 


HOY 
eens 


A AS 
oN fa) ° 
yaaa ; VON 


Phloem A O") ARE 
Bundle sheath —#./ 


Palisade par. LOR Nd & ‘ ») J ADEN A | 
A ALE IAQ Ke nad ae 

gs ZTE 

ZSZ= CO ZS 


Adaxial epidermis 

Intercellular space \ 

Spongy parenchyma ~~: 
Outer bundle sheath 

inner bundle sheath 


Ve 


aes 


i ea! =< stomata 


cae Ce Abaxial epidermis 
ie Ps ee Abs 
Ce cae] Palisade parenchyma 


ran 
GES 


Fics. 5-7. Transverse sections of leaves: 5, Allium stamineum ; 6, Colchicum 
biebersteinii; 7, Gagea arvensis var. semiglabra. 


52 MADRONO [Vol. 16 


followed by a layer of articulated laticiferous tubes. These laticifers pos- 
sess perforation plates similar to sieve plates. The spongy parenchyma 
is very weakly developed, forming a layer of 2—3 cells inside the “ring” 
of laticifers. As the leaves reach maturity, remarkably large water-storing 
parenchyma cells in the center of the leaf collapse and form lacunae. Col- 
lateral bundles are surrounded by bundle sheaths. “Parallel” running 
major bundles are more or less spaced from each other, but they are 
transconnected by minor bundles. There are a few xylem fibers associated 
with the bundles, but phloem fibers are very rare. Sclerenchymatous 
idioblasts are abundant and seem always to be associated with inter- 
connecting transverse bundles or to occur terminal to freely ending 
veinlets. 


COLCHICUM BIEBERSTEINII Rouy (fig. 6) is a stout plant which grows 
in foothills and on mountain slopes protected from the wind. It blooms 
as early as in February, right after a few sunny winter days. The basal 
leaves arising from tunicated bulbs lie on the ground. They are flat, 
narrow, dark green and coriaceous in texture, 8-10 mm. wide and 
380-450 microns thick. Involution does occur, although the leaves are 
not provided with any specialized motor cells. A similar situation was 
observed in Oryzopsis hymenoides (Roem. et Schult.) Ricker by Shields 
(1951b). The epidermal cells are long tabular cells with straight anticlinal 
walls. Abaxial and adaxial sides of the epidermis are coated with a thick 
cuticle layer which exhibits very fine dentation as seen in transectional 
view. Outer tangential walls of the upper and lower epidermis are exceed- 
ingly thick and are marked with distinct lamellation (25 microns or even 
thicker) while the inner tangential walls are moderately thickened. Radial 
walls of the epidermal cells however do not show a noticeable thickening. 
The stomata occur on both surfaces, but are more frequent on the abaxial 
surface. They are not sunken, but lie at the same level with cuticle and 
thick outer walls of the epidermal cells. The guard cells also possess very 
thick cell walls and are provided with well developed inner and outer 
ledges. Subsidiary cells are not present. Unicellular, short, conical tri- 
chomes with blunt tips occur along leaf margins only. These trichomes 
are dead cells with extremely thick walls and practically no lumens. 

The leaves of Colchicum biebersteinii present an isolateral organization 
with a biseriate palisade on both upper and lower sides of the mesophyll. 
Palisade parenchyma is strongly developed at the expense of the spongy 
parenchyma. It occupies two thirds of the mesophyll. Palisade paren- 
chyma has thin walls in general, but the outer cell walls of the outer 
parenchyma layer are remarkably thickened toward the leaf margins. 
The spongy parenchyma is scanty and represented by 2—5 layers of iso- 
diametric cells in the central portion of the mesophyll. Prismatic crystals 
occur in both palisade and spongy parenchyma cells. Large and small 
vascular bundles imbedded within the spongy mesophyll are sheathed 
by border parenchyma. The median bundle is not different from other 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 33 


bundles. The general pattern of venation is striate, but the veins run- 
ing along the longitudinal axis of the leaf are interconnected by small 
bundles at frequent intervals so that a strongly developed and compact 
network of veins with many vein islets of varying sizes results. Isolated 
tracheids with spiral thickenings are found occasionally outside the 
bundles. Vascular bundles are not accompanied by fibers. Strands of 7-8 
collenchyma cells with highly reduced lumens occur along leaf margins 
under the epidermis. 


GAGEA ARVENSIS Dum. var. SEMIGLABRA Beck (fig. 7) is a bulbous 
plant occurring commonly in fields, fallow lands and in thickets under 
the protection of small steppic shrubs. The smooth, flat, lanceolate, light 
green, tender leaves are 7—8 cm. long, 5-6 mm. wide and 350-380 microns 
thick. Involution occurs upon wilting, although there are no specialized 
motor cells. 

The epidermis consists of long tabular cells with straight radial walls. 
The outer tangential walls of epidermal cells are 12—14 microns thick 
on both surfaces. However, the outer tangential walls of the upper epi- 
dermis project and become almost papillate especially in the cells adjacent 
to the stomata. Apparently the adaxial epidermis is involved in water 
storage. A finely striate cuticle covers the entire leaf. The stomata are 
not sunken, and they occur on both the upper and lower epidermis at 
nearly equal frequencies. The guard cells are 20 microns wide, 45 microns 
long and 20 microns thick. They are equipped with well developed inner 
and outer ledges. Subsidiary cells are not present. The long unicellular 
trichomes occurring along leaf margins are dead, thick walled cells 
with papillate projections all around them. The leaf structure is dorsi- 
ventral, and the mesophyll is provided with a single layer of palisade 
on the abaxial side. The palisade parenchyma is poorly differentiated, 
although sometimes it appears to be biseriate. The greater part of the 
mesophyll consists of large spongy parenchyma cells, 4—5 cell layers 
thick. These cells are more or less isodiametric and variable in size, but 
cells with a diameter of 100-120 microns are not uncommon. The spongy 
parenchyma cells, with large central vacuoles filled with sap, are mainly 
responsible for the succulent texture of the leaves. Crystal sand is found 
in the spongy parenchyma cells, but no specialized idioblasts occur. 

Vascular bundles are not accompanied by fibers, but interestingly 
enough are sheathed by two layers of border parenchyma, thin walled 
cells which appear round in transectional view. The inner bundle sheath 
consists of smaller cells than those of the outer bundle sheath. There are 
no sheath extensions on either side of the bundles. Vascular bundles 
present a typical striate pattern. They converge near the leaf apex, but 
interconnecting veinlets have not been observed. They are more wide- 
ly spaced than in any other species studied. Isolated vascular strands lie 
parallel to adjacent vascular bundles. These strands consist of a few 
helical tracheids and terminate in undifferentiated procambial cells at 


54 MADRONO LVol. 16 


both ends. There is no evidence of bundle sheaths around the isolated 
vascular strands. 

Leaves of Gagea do not present striking xeromorphic features. They 
represent a leaf type at the border line between mesomorphic and xero- 
morphic structures. 


MERENDERA TRIGYNA (Adam) Woron. (fig. 8) is a bulbous plant with 
strap shaped, flat leaves which are 3-4 mm. wide and 300-350 microns 
thick. It occurs in the mountain steppe and blooms during February and 
March. The leaves are isolateral and involution occurs upon wilting. 

The epidermis consists of rectangular cells, 14 microns wide and 70 
microns long. Upper and lower epidermis are coated by a striate cuticle 
2-3 microns thick. The outer tangential walls of the epidermal cells are 
remarkably thick, and cell lumina are extremely reduced. Both upper 
and lower epidermal cells contain chloroplasts. The stomata are sunken 
and distributed on both adaxial and abaxial surfaces. Guard cells are 12 
microns wide and 32 microns long as seen in face view. Inner and outer 
ledges are well developed, the outer ones being cutinized. Subsidiary cells 
are not present. Short unicellular trichomes occur along leaf margins. 
The cell walls of the trichomes are very thick and appear refringent 
under polarized light. 

The mesophyll has a biseriate palisade on both adaxial and abaxial 
sides. Palisade parenchyma cells are small and densely arranged. They 
occupy two thirds of the mesophyll as seen in transverse section. Spongy 
parenchyma consists of 5—6 cell layers between upper and lower palisade. 
Crystal sand occurs in spongy parenchyma cells. There are no special- 
ized idioblasts in the mesophyll. 

Vascular bundles are closely spaced and transconnections occur very 
frequently. Border parenchyma cells containing very few chloroplasts 
are restricted to outer edges of xylem and phloem. Consequently they do 
not form a continuous sheath around bundles. There are no fibers 
associated with the vascular bundles, but strands of partially lignified, 
hypodermal fibers extend along leaf margins. 


Muscari comosuM Mill. (fig. 9) is a bulbous plant which grows in 
mountain steppe as well as in cultivated lands. Strap-shaped leaves, 
slightly deflexed at their tips, are 10-15 cm. long, 1—1.5 cm. broad and 
250-350 microns thick. Incomplete involution takes place upon wilting. 

The upper epidermis is smooth, while the lower epidermis exhibits a 
wavy surface although there are no ridges on the abaxial side. Epidermal 
cells are almost prosenchymatous, 10-15 microns wide and 150-200 
microns long. The cuticle is thin and the outer tangential walls of the 
epidermal cells are moderately thick. Stomata occur on both surfaces of 
the leaves. They are nearly at the same level with the rest of the epi- 
dermal cells of the adaxial surface, but the stomata of the abaxial surface 
are hidden within the furrows caused by the plication of the epidermal 


Adaxial epidermis 

Chloroplasts 
8 Palisade parenchyma 
Spongy parenchyma 


Pa 


Abaxial 
epidermis 


so: 23 NS eeachioroplests 


Palisade parenchyma 


; a ara, 
Palisade parenchyma 1O0Ou 
St 


Lacuna ; 
Idioblast with Raphides 
—— Water storing par. 


\ 
Palisade parenc 


Abaxial ridges a _ Adaxial epidermis 
 \wte— Spongy parenchyma 
eo ® Hy Raphides 
By Lacuna 
Bundle sheath Ce ‘ 
Xylem 


Phloem 


Xe — Ny wy 
Bundle sheath 
> Palisade par. 


Abaxial epid. 


Fics. 8-10. Transverse sections of leaves: 8, Merendera trigyna; 9, Muscari 
comosum; 10, Muscari racemosum. 


56 MADRONO [Vol. 16 


tissue on this side. Guard cells are not accompanied by subsidiary cells 
and there are no trichomes. 

Leaves are typically isolateral; mesophyll is provided with a uniseriate 
palisade on adaxial and abaxial sides. Palisade parenchyma cells are 
nearly cylindrical in shape and are compactly crowded. Cells of the 
abaxial palisade are almost twice as long as the cells of the adaxial pali- 
sade. Together the two palisade layers occupy about one third of the 
thickness of mesophyll. Spongy mesophyll consists of 7—8 layers of 
cells which are loosely arranged, leaving conspicuous intercellular spaces 
between them. Chlorenchymatous cells are located around vascular 
bundles and adjacent to the inner edges of the palisade layers. Spongy 
parenchyma cells, located centrally in the mesophyll, are larger than the 
cells of the peripheral spongy parenchyma and contain few chloroplasts 
and large vacuoles. This water storing parenchyma gives a fleshy texture 
to the leaves. It is interesting to note that this tissue becomes lacunate 
as the leaves reach maturity. Cylindrical idioblastic cells containing 
raphides are abundant in the spongy mesophyll. 

Vascular bundles are surrounded by a single layer of bundle sheath. 
Large and small bundles which are distantly spaced and which alternate 
with each other run along the longitudinal axis of the leaves. Small veins 
interconnecting the ‘‘parallel’’ veins are common. Sclerenchyma cells 
are not present. 

With the exception of isolateral structure of leaves and presence of 
water storing tissue, Muscari comosum does not present any appreciable 
xeromorphic feature and lies rather on the mesomorphic side, especially 
in respect to the presence of lacunae and prominent intercellular spaces 
and high proportion of spongy parenchyma as compared to the propor- 
tion of palisade parenchyma. It can well be regarded as a mesoxero- 
morphic leaf. 


MuscarI RACEMOSUM Mill. (fig. 10) is a larger bulbous plant than 
Muscari comosum Mill. It grows in the mountain steppe. The leaves are 
lanceolate to almost ovate, tapering gradually toward apex. They are 
coated with a waxy substance which gives a glaucous appearance to the 
leaves. The upper surfaces of the leaves are flat and the lower surfaces 
are ridged. Involution takes place upon wilting although there are no 
bulliform cells in the epidermis. 

Epidermal cells are prosenchymatous, 18—20 microns wide and 220-280 
microns long. The cuticle is thicker than the cuticle of the preceding 
species. Outer walls of the epidermal cells are very thick and are char- 
acterized by lamellate cutinization. The wall thickness is more prominent 
in the epidermal cells on the abaxial surface than in those on the adaxial 
surface. Inner tangential walls of the abaxial epidermal cells are also 
thickened considerably. Portions of the abaxial epidermis beneath vas- 
cular bundles have exceedingly thick outer cell walls reaching 25 microns 
in thickness. These thick-walled epidermal cells contribute to the for- 


1961] KASAPLIGIL: FOLIAR XEROMORPHY a7 


mation of abaxial ridges on the leaves since there are no hypodermal 
fibers along ridges. Outer cell walls of the abaxial epidermis adjacent to 
the mesophyll between vascular bundles are relatively thin walled. Con- 
sequently thin and thick walled portions of the lower epidermis alternate 
with each other following the spacing of vascular bundles and dorsal 
ridges as seen in transverse section. Stomata are sunken and occur on 
both surfaces. There are no subsidiary cells or trichomes. 

Unlike the previous species, the leaves of Muscart racemosum are 
dorsiventral with a biseriate palisade on the lower side. The central por- 
tion of mesophyll is occupied by an extensive water storing tissue which 
takes up two thirds of the thickness of the mesophyll between vascular 
bundles. Water storing parenchyma cells contain very few chloroplasts. 
Assimilatory cells of the spongy parenchyma are situated at the periph- 
eries of the water storing tissue and are very loosely arranged, leaving 
conspicuous air spaces between them. Sizable lacunae are present in the 
centers of water storing tissues. Although these lacunae seem to be schizo- 
genous cavities, there is ample evidence for collapsed cell walls, suggest- 
ing that they are partly lysigenous in origin (cf. Newcombe 1894). 
Bizarre idioblasts containing raphide bundles are abundant in the meso- 
phyll. 

Vascular bundles are surrounded by bundle sheaths and are devoid 
of fibers. However, converging bundles of the leaf apex are associated 
with many tracheoid idioblasts. Vascular bundles are widely spaced, as 
in the preceding species. Transverse veinlets occur frequently, either 
connecting the longitudinal bundles or terminating blindly in mesophyll. 

The leaves of Muscari racemosum also can be considered as meso- 
xeromorphic, especially considering the presence of large lacunae. This 
lacunate condition, however, may also be considered as an ancestral 
feature, if we follow the classical belief that the xeromorphic leaves are 
derived from mesomorphic leaves during evolution. 

Ornithogalum is represented by ten species in the steppic flora. Only 
two species are considered in this paper. 


ORNITHOGALUM ARMENIACUM Bak. (fig. 11) is a bulbous plant which 
grows in the valleys of the mountain steppe. Linear leaves are 2—3 mm. 
wide and 250-400 microns thick. The upper leaf surface is flat, the 
lower one ridged by prominent thickenings of the outer walls of epidermal 
cells. Involution occurs upon wilting (fig. 11B). 

Epidermis is covered by a heavy cuticle. Epidermal cells of both 
surfaces are more or less tabular in shape, 20-22 microns wide and 247— 
330 microns long. Stomata are not sunken and occur on both surfaces. 
Guard cells are thick-walled and equipped with cutinized external ledges 
only. Guard cells are 13 microns wide and 39 microns long, and are not 
accompanied by subsidiary cells. Trichomes are not present. 

The leaf is isolateral and mesophyll is provided by a uniseriate palisade 
on adaxial and abaxial sides. It is interesting to note that the adaxial 


58 MADRONO [Vol. 16 


palisade is interrupted by water storing parenchyma cells above the 
median vascular bundle (fig. lla and b). Assimilatory parenchyma 
cells of the spongy mesophyll form one or two layers bordering the pali- 
sade layers. The central portion of mesophyll is occupied by water 
storing parenchyma and schizogenous lacunae. Raphide containing excre- 
tory idioblasts are abundant particularly in the spongy mesophyll and 
along the leaf margins. 

Vascular bundles lack sclerenchyma and are sheathed by bundle 
sheaths. Large bundles are closer to the adaxial leaf surface than are the 
aiternating small bundles. Transverse veinlets interconnect large ‘‘paral- 
lel” veins obliquely or perpendicularly, very often without making any 
contact with the alternating smaller longitudinal bundles. 


ORNITHOGALUM NARBONENSE L. var. PYRAMIDALE Boiss. (fig. 12) is 
a bulbous plant which grows in fields and fallow lands as well as in the 
mountain steppe. Strap-shaped, fleshy leaves are 20-30 cm. long, 1—1.5 
cm. wide and 500-600 microns thick. Upper surfaces of the leaves are 
flat; lower surfaces are more prominently ridged than those of O. armeni- 
acum. Slight involution occurs upon wilting. 

The epidermis consists of very narrow cells, tapering toward the ends. 
Epidermal cells are 618-825 microns long, 25-30 microns wide, and 
35-38 microns deep. Superficial walls of the epidermal cells project on 
both adaxial and abaxial sides. Outer walls of the epidermal cells are 
thick and heavily cutinized. Stomata are distributed throughout both 
surfaces of leaves and are sunken distinctly on the lower epidermis. 
Guard cells are 12 microns wide and 32 microns long; they are not ac- 
companied by subsidiary cells. Two kinds of unicellular trichomes have 
been observed: very thick walled conical trichomes 100-110 microns 
long which occur along the abaxial ridges; and cylindrical trichomes 
345—450 microns long which are confined to the leaf margins. 

Leaves are isolateral and the mesophyll has a uniseriate, well-developed 
palisade on abaxial and adaxial sides. Again the adaxial palisade is inter- 
rupted by a few large cells without chloroplasts along the median groove. 
These cells may function as motor cells together with other cells between 
median bundle and epidermal cells lining the groove. Spongy parenchyma 
is more prominent than it is in the leaves of the preceding species. The 
central portion of mesophyll is occupied by a water storing tissue which 
consists of very large irregular parenchyma cells of 4-5 layers. Schizo- 
genous lacunae develop within this water storing tissue in mature leaves. 
Excretory idioblastic cells containing druses or raphides of calcium oxa- 
late are abundant in spongy mesophyll. Individual raphide crystals are 
4—5 microns thick and 206-230 microns long. Very long isolated individ- 
ual fibers occur in the mesophyll. These fibers are about 20 microns thick 
and 8-14 millimeters long, extending along the longitudinal axis of the 
leaf, not far from large vascular bundles. 

The vascular bundles are more closely spaced than those of the pre- 


Adaxial Bere 
epidermis 


Xylem 
Phloem 


\— Water storing par. 
Raphides 
Idioblast 


lOOu 


Stoma 
Adaxial epidermis 


Palisade par. 
Bundle sheath 


Lacuna 


5 
AST 


Raphides 


Idioblast 


I2 


“0 Co, mil 
eee an 
Abaxial epidermis ———————— we 
Palisade parenchyma 
Spongy parenchyma 
Water storing parenchyma 


Water storing parenchyma 
Palisade par. 


Spongy par. 


mg: 


ia RS ye Re ER | 

ie ; a | 

(ashy ia ee 

Baten Sanne ee 
29) 


SS 
Sete: 
WA : 
C4 
ae 
5. 
oer 
Noi 
Sas 


by ary 
oe 
BO 
oe 


Pee SN Stomata 
BG J 


Trichome 


Abax. epid. Phloem fibers 


Palisade par. 


Fics. 11-13. Transverse sections of leaves: 11, Ornithogalum armeniacum ; 
12, Ornithogalum narbonense var. pyramidale ; 13, Crocus ancyrensis. 


60 MADRONO LVol. 16 


ceding species. Large and small veins are interconnected by small veins 
which always run into major veins obliquely and never perpendicularly. 
Bundle sheaths are well developed around large and small bundles. 


IRIDACEAE 


This family is represented by Crocus, Gladiolus and Iris in the steppic 
flora. The genus Crocus, which has tunicated, fleshy, underground corms, 
is represented by six species, two of which will be considered in this paper. 
Gladiolus possesses a tunicated corm which is not so deeply buried as 
the corms of crocuses. Gladiolus is represented by a single species. The 
genus /ris is represented by four species in the steppic flora, all of them 
having fleshy rhizomes which become slightly woody in age. 


CROCUS ANCYRENSIS Maw (fig. 13) is an endemic species abounding 
in the mountain steppe and blooming from February to April. It has 
very narrow, stiff, dark green and shiny leaves which are not differenti- 
ated into petiole and lamina. These leaves, which remind one of pine 
needles, represent an extremely reduced leaf type among geophytic mono- 
cotyledons of the steppic flora. Their length ranges from 5-8 cm., their 
width is 800-900 microns and their thickness in the middle portion of the 
leaf, across the dorsal ridge, is 400-450 microns. The transectional out- 
line of the leaf is almost “‘T’’-shaped. Lateral flaps of the leaf are revo- 
lute, 120-162 microns thick. Possibly, change of turgor pressure in the 
abaxial epidermal cells of the lateral flaps is responsible for the revolu- 
tion mechanism. 

The epidermis consists of very narrow and elongated cells which appear 
almost fusiform in surface view. Epidermal cells are 290-300 microns 
long, 14-16 microns wide and 18—24 microns deep. The cuticle is 1-2 
microns thick and covers the entire leaf surface. Both outer and inner 
tangential walls of the epidermal cells are strongly thickened, the outer 
walls being much thicker than the inner ones. The lumina of the epidermal 
cells are highly reduced, particularly along the margins of the lateral 
flaps of the leaf. The stomata are sunken and appear on the abaxial sur- 
face of the foliar flaps and along the upper portions of the thickened 
abaxial ridge of the leaf. The adaxial surface of the leaf and the abaxial 
surface of the dorsal ridge are devoid of stomata. Guard cells are very 
small, 7-8 micrcns wide and 18-20 microns long. Very few unicellular 
trichomes occur along the margins of foliar flaps and along lateral cor- 
ners of the dorsal ridge. 

Leaves are isolateral in a peculiar way. A strongly developed, biseriate 
palisade consists of small and tightly arranged cells. Palisade is inter- 
rupted by water storing parenchyma cells on the adaxial side. Spongy 
mesophyll is differentiated into assimilatory and water storing tissues. 
Small chlorenchymatous cells of the spongy mesophyll form 2—5 rows 
in foliar flaps as well as in dorsal ridge, beneath palisade parenchyma. 
The central portion of the mesophyll is occupied by water storing tissue 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 61 


which extends from the dorsal ridge to the adaxial epidermis in a 
“V-shape as seen in transectional view. Water storing parenchyma cells 
lack chloroplasts. They appear as a white band on the adaxial sides of 
the leaves. Crystals and idioblasts are not present. 

Vascular bundles are strongly developed and run very close to each 
other and to the abaxial epidermis. A median bundle is not present. Each 
foliar flap and each “corner” of dorsal ridge is provided with a major 
bundle which is characterized by the presence of a well-developed bundle 
cap. Minor bundles have small amounts of phloem fibers or none. The 
minor bundles anastomose highly and interconnect the major bundles. 
Xylem fibers are not present and bundle sheaths are not clearly differen- 
tiated. 


CROCUS SUTERIANUS Herb. (fig. 14) is another endemic species which 
occurs in the mountain steppe of Asia Minor. It has a spherical corm 
protected by a fimbriate tunic. The leaves are linear, 10-15 cm. long, 
2-3 mm. wide and 200-300 microns thick in foliar flaps. With the excep- 
tion of a white adaxial stripe, the leaves are dark green and stiff. The 
dorsal ridge is more pronounced than that of the preceding species and 
the transectional outline of the leaf is T-shaped (fig. 14b). The foliar 
flaps are revolute and roll backwardly upon wilting. Ridged middle 
portion of leaves is 1-1.5 mm. thick, which is twice or three times as 
thick as that of Crocus ancyrensis. 

Epidermal cells, which are very slender and fusiform as seen in face 
view, are 130-243 microns long, 20-26 microns wide and 25—28 microns 
deep. The entire leaf surface is covered by a thick cuticle which reaches 
a thickness of 4 microns on the adaxial epidermis and on the abaxial 
epidermis of the dorsal ridge. Inner and outer tangential walls of the 
epidermal cells are very thick. Abaxial epidermal cells of foliar flaps 
are characterized by dome-shaped outer walls. These highly vacuolated 
cells are relatively thin-walled and may function as motor cells. Stomata 
are sunken and restricted to the abaxial epidermis of foliar flaps and 
to the lateral sides of the dorsal ridge. Guard cells are very small, 14 
microns long and 7 microns wide. There are no subsidiary cells. Uni- 
cellular, dead trichomes occur only along the margins of the foliar flaps. 
Trichomes are 120 microns long and 15 microns thick. 

The leaves are peculiarly isolateral as described in the preceding 
species, The adaxial palisade consists of narrow cylindrical cells which are 
tightly arranged and biseriate, but the cells tend to be in three layers 
near water storing tissue. The palisade tissue is strongly developed at 
the expense of spongy parenchyma in the foliar flaps and occupies more 
than half of the thickness of the mesophyll. Again the spongy mesophyll 
is differentiated into assimilatory and water storing tissues (fig. 14a 
and b) as described for Crocus ancyrensis. Assimilatory spongy paren- 
chyma cells are small, tightly arranged and form four layers on the 
abaxial side of the foliar flaps and in dorsal ridge. Water storing paren- 


62 MADRONO — [Vol. 16 


chyma cells are large and have prominent air spaces between them. Idio- 
blasts containing large prismatic crystals are abundant in the assimila- 
tory spongy tissue. These crystals are 15-16 microns thick and 97-146 
microns long. The idioblasts are situated around the bundle caps and 
occasionally occur also ‘“‘scattered” within the assimilatory spongy meso- 
phyll. 

Vasculation of the leaves is similar to that of Crocus ancyrensis with 
minor differences. The major bundles along the margins of foliar flaps 
and those at the “corners” of the dorsal ridge are accompanied by xylem 
and phloem fibers. Furthermore, the bundle sheaths are more distinctly 
differentiated and the vascular bundles more compactly arranged than 
those of C. ancyrensis. 


GLADIOLUS ATROVIOLACEUS Boiss. (fig. 15) is a cormous plant 30—55 
cm. high. It has a wide distribution in the Middle East. It grows in fields, 
fallow lands as well as in mountain steppe. The leaves are strap-shaped, 
firm, strongly ribbed, 15-20 cm. long, 4-5 mm. wide and about 0.5 mm. 
thick. From the morphological point of view, these leaves present a very 
peculiar structure. Typically, the leaves are unifacial, with inverted 
bundles, but the presence of two marginal bundles and a prominent 
marginal groove (fig. 15a and b) along one edge of the blade and the 
presence of a single bundle with a well developed ridge along the other 
edge of the blade suggests the folding and fusion of two halves of the 
blade during ontogeny. However, there is no evident line of fusion in the 
mesophyll. The problem may be solved by an ontogenetic study. Corms 
are ovoid, tunicated and buried superficially in ground. 

If the interpretation of the foliar structure given above be true, the 
epidermis of upper and lower leaf surfaces may represent the abaxial 
epidermis only. The epidermal cells are tabular in form, 48 microns long, 
30 microns wide and 20 microns deep. The cuticle is 2 microns thick on 
the blade surface and 4-5 microns thick along leaf margins. Epidermal 
cells form 2—5 papillose projections. Stomata are sunken and occur on 
both leaf surfaces. Epidermis of the projecting ribs has no stomata or 
trichomes. 

The leaves are isolateral in the sense that abaxial and adaxial sides 
of the leaves present identical structure, but the mesophyll is not differ- 
entiated into palisade and spongy parenchyma. The mesophyll consists 
of very small, tightly arranged, elliptical cells which resemble palisade, 
but they lie parallel to the leaf surface. Water storing parenchyma cells 
are situated between opposite major ribs of the blade (fig. 15b). Idio- 
blasts containing prismatic crystals are distributed at random within 
mesophyll. Crystals are 16 microns thick and 65-160 microns long. 
Tracheoid idioblasts occur commonly in mesophyll. 

Major vascular bundles are accompanied by bundle caps and project 
on leaf surfaces as well as along leaf margins. Major bundles extend 
“parallel” but the minor bundles which are imbedded in mesophyll anas- 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 63 


Adaxial epidermis 


Water storing parenchyma 
Palisade parenchyma 


Xylem Spongy parenchyma 


Phloem 


Phloem fibers 


Crystal 


4 


Stomata 
Abaxial epidermis —— 


Trichomes PEERS SADE, 

Ae ee yA FEES SNe 

el ——\ wh Ana anpnasndel stem, 
Spongy par. BABI re PaaS ase Src SSO ae 


Phloem eas on 


We Pease > 


senate) e 
ap ae oseeeee at 


oo oe : ee pecan es DOR ee tN 
Xylem SDE S =» en ai eee eee eee emer bundles 
SBS aS REO et eee SOE en 
tes uk Y SC 
Crystal Stine pat ae SS Sa SOTO 
[oan Eee gaa eee eee 
Water pomtel ee ES Se) 
y Es: Sg aac ach uD) as 
Phloem BP EA Chey Corin MEN aeRO RR DV 
Bey A ONE OERO a oe MENT Sibeee nok PNY 
Mec ZR Fy aera A Bundle sheath 
¥ wire SOPs Rey eseceu a 
yes chess Xylem 
gE eG Phloem 
\OOu 


Fics. 14-15. Transverse sections of leaves: 14, Crocus suterianus ; 
15, Gladiolus atroviolaceus. 


tomose frequently and interconnect the major bundles. Minor bundles 
lack fibers and are surrounded by bundle sheaths. As a whole, the leaves 
present a compact vascular system. 


Irts APHYLLUS L. (fig. 16) is a rhizomatous plant occurring in the 
mountain steppe and in the openings of forest remnants of the steppic 
region. Bluish-green, sword-shaped, erect leaves are 10-15 cm. long, 1—1.5 
cm. broad and 800 microns thick in the lower sheath portion, 400 microns 
thick in the upper blade portion. Unifacial structure of leaves with in- 
verted vascular bundles is seen clearly (fig. 16a, b, and c). 

Epidermal cells are 227 microns long, 48 microns wide and 39 microns 


64 MADRONO a8 [Vol. 16 


deep and rectangular in form. Stomata are sunken and occur on both sur- 
faces of the blade. However, there are no stomata on the adaxial epidermis 
of the flaps in sheathing lower portions of leaves. Considering the devel- 
opment of these unifacial leaves, it is understandable that a similar epi- 
dermis occurs all around the upper blade portions of leaves; therefore the 
actual distribution of stomata would be confined to abaxial epidermis 
only. The guard cells are typically reniform, 50 microns long and 25 mi- 
crons wide. There are no subsidiary cells. Unicellular, thick walled tri- 
chomes occur on the abaxial epidermis only and there are no trichomes 
on the adaxial epidermis of sheathing flaps. The trichomes are 40 microns 
long and conical in shape. 

The blades are isolateral in the sense that both sides of the blades are 
similar in regard to uniform mesophyll and inverted bundles, although 
there is no palisade tissue. On the other hand sheathing flaps present a 
dorsiventral structure since mesophyll is differentiated into water storing 
parenchyma on the adaxial side and assimilatory parenchyma on the aba- 
xlal side. Furthermore xylem tissues of vascular bundles face the adaxial 
epidermis in sheathing flaps (fig. 16b). The mesophyll of the blade con- 
sists of elliptical spongy parenchyma cells lying with their longitudinal 
axes parallel to leaf surfaces. This is quite similar to the situation found 
in Gladiolus leaves except that the parenchyma cells in /ris leaves are at 
least twice as big as the assimilatory parenchyma cells of Gladiolus leaves. 
Extensive water storing tissue occupies the central part of the mesophyll 
in lower sheathing portions of leaves. Idioblasts containing prismatic 
crystals are abundant in mesophyll. These crystals are very similar to 
those found in the leaves of Crocus suterianus and Gladiolus atrovio- 
laceus. 

Vascular bundles of the leaves are strongly developed, but are not as 
compact as the bundles in Crocus and Gladiolus leaves. The bundle caps 
also are strongly developed along major veins, but do not project as ribs 
on the leaf surface. Xylem tissues of the major veins possess a few fibers. 
Phloem fibers are lignified while thick walls of xylem fibers do not show 
any sign of lignification. Veinlets interconnecting the major bundles occur 
frequently. Some of the veinlets terminate blindly in mesophyll and very 
often are associated with tracheoid idioblasts at their tips. Strands of 
fibers occur along the margins of sheathing blades as well as along both 
edges of blades (fig. 16b and c). 


ORCHIDACEAE 

This family is represented by a number of terrestrial orchids which 
grow in moist places of the steppic region. The specimens of Limodorum 
abortivum and Orchis mascula subsp. pinetorum were collected from an 
open pine forest in Beynam at an altitude of 1450 meters. 


LIMODORUM ABORTIVUM (L.) Sw. (fig. 17) is a saprophyte with well 
developed fleshy rhizomes. Scale-like leaves are small, very tender and 
nearly surround the scape. Apical portions of these leaves slightly diverge 


t——-—H 
|OOu 
Trichomes “eee 5 ©. a) C) 

Stoma mois tie ae AN OA IKEA 

Epidermal A a RSE ita Phloem fibers 
Bey ererstins : BCP phloem 
Bundle sheath aes 2 : 


meee Aylem-: 25. 


€ 
Cd. 
ie | 
<\ ie, oe : 
Bg oe 
SOO) eee eee Xylem fibers 
Onerme 


eee Xylem 
J S\ Bundle sheath 
ae ey, 2 Ca Pe NEE me 
Ge. Cae: Be ASN? ce) 
ZS XY Page oe Bee Phloem fibers 
ay Cee pe (2 ORE We 
Se ce 


q CL EC 


pS Papillose epidermal 
Ts ER on 6°23 )s cell 
ER: er ETO) £3 S2 Cy 


2 YoRo XD 

Te S— Spongy parenchyma 
Idioblast with raphides 
= “= — Bundle sheath 

Se ae 


<a Tie rs) 
BO Abax. epidermis 


Adaxial epidermis —~>—~____ 


Ks) 


Vacuole 


Choroplast 
Nucleus 


Bundle 


Cx Adee: sheath 


Idioblast 48 PASSA: 
wih ee 
raphides CICS 
£\ 


os Guard cell 
Ledge 


Abax. epid. 


Fics. 16-18. Transverse sections of leaves: 16, Iris aphyllus; 17, Limodorum 
abortivum; 18, Orchis mascula subsp. pinetorum. 


66 MADRONO LVol. 16 


from the scape and form a short blade about 2-3 cm. long, 1—1.5 cm. broad 
and 250-300 microns thick. Involution occurs slightly upon wilting. 

The adaxial epidermis consists of large tabular cells with papillose outer 
walls. Water storage takes place in upper epidermis. The abaxial epidermis 
consists of much smaller cells than those of the adaxial epidermis. Stomata 
occur on the abaxial epidermis only and are not sunken. Guard cells are 
65 microns long and 23 microns wide. Each stoma is surrounded by six 
subsidiary cells, each guard cell being subtended by three of them. The 
presence of four Tvadescantia-type subsidiary cells was reported by Sole- 
reder and Meyer (1930) in some orchidaceous genera. The radial walls 
of subsidiary cells extend toward the stomatal opening, the stomatal ap- 
paratus as a whole resembling a rosette in face view. 

Mesophyll is not differentiated into palisade and spongy parenchyma 
and consists of more or less uniform isodiametric parenchyma cells ar- 
ranged tightly. Patches of tracheoid idioblasts, fibers and mucilage- or 
raphide-containing excretory idioblasts are present in mesophyll. The oc- 
currence of such a great variety of idioblasts together in one species was 
not observed in other species of the present study. 

Major vascular bundles present a striate pattern, but minor bundles 
anastomose frequently and form also peculiar zigzags extending back and 
forth and interconnecting major bundless eventually. 


ORCHIS MASCULA L. subsp. PINETORUM Boiss. (fig. 18) has tuberous 
fleshy roots protected by a dark brown cork tissue. The leaves are basal, 
flat and very tender, 10-15 cm. long, 1.5—2 cm. wide, 350-450 microns 
thick. Upper leaf surface is grooved along the median bundle and the 
lamina halves fold over along the median bundle when a water deficit 
develops. 

The adaxial epidermis consists of remarkably large tabular cells 340— 
550 microns long, 100—150 microns wide and 120-200 microns deep. These 
cells are characterized by thick outer walls, peripheral cytoplasm and 
large vacuoles. Their nuclei are pushed against inner tangential walls. 
They contain chloroplasts and concentric starch grains. Epidermal cells 
along the adaxial groove are specialized as motor cells and are much 
smaller than adjacent water storing epidermal cells. Adaxial epidermis is 
mainly responsible for the fleshy nature of the leaves. The occurrence of 
water storing epidermis was reported by Metzler (1924) for other orchi- 
daceous genera such as Dendrobium, Otochilus, Pholidota and Pleione. 
The abaxial epidermis consists of much smaller cells. Stomata occur on 
the abaxial epidermis only and are not sunken. Guard cells are 70 microns 
long, 35 microns wide and are equipped with strongly developed inner and 
outer ledges. Subsidiary cells and trichomes are not present. 

Mesophyll consists of small isodiametric parenchyma cells which are 
arranged rather tightly. Idioblasts containing protein crystalloids (Kues- 
ter 1935) and raphide bundles associated with mucilage occur at random 
in the mesophyll. There is no sclerenchyma, but a strand of a few hypoder- 


1961] KASAPLIGIL: FOLIAR XEROMORPHY 67 


mal collenchyma cells occurs occasionally on the dorsal side of the ridge. 

Vascular bundles are widely spaced. Major and minor veins anastomose 
frequently. Bundle sheaths are differentiated distinctly around major 
bundles. 

The leaves of Limodorum abortivum and Orchis mascula subsp. pine- 
torum do not show striking xeromorphic features with the exception of 
folding of blades, epidermal water storage, compactness and decreased cell 
size in the mesophyll, and abundance of excretory idioblasts. On the other 
hand they exhibit a prevailingly mesomorphic structure, and may be con- 
sidered as mesoxeromorphic. 

The present study shows that not all xeromorphic characters occur uni- 
versally in the leaves of the geophytic monocotyledons investigated. Each 
plant exhibits a different combination of xeromorphic features. Further- 
more, there are quantitative differences in the degree of development of 
each particular xeromorphic feature. I believe many of these xeromorphic 
characters are genetically fixed as the result of a natural selection in this 
semi-arid environment. Consequently these plants are morphologically 
and physiologically adapted to tolerate the factors of the environment. 
Drought is one of the most severe factors and the plants in question are 
well adapted to survive drought. On the other hand some of the features 
described may represent xeroplastic characters which may be subject to 
qualitative and quantitative changes under varied conditions of the envi- 
ronment. It would be very desirable to conduct an experimental study 
using controlled conditions to determine the extent of plasticity of the 
foliar xeromorphic features. 


SUMMARY 


The foliar histology of eighteen species of geophytic monocotyledons 
representing Araceae, Liliaceae, Gramineae, Iridaceae and Orchidaceae 
has been described. The research materials were collected wholly from the 
central Anatolian steppic region. They exhibit certain uniformities such 
as geophytism, shallow root distribution, leaf shape, osmotic concentra- 
tions, and phenological periodicity. 

Xeromorphic characters observed in gross morphology and in anatomi- 
cal structure do not occur universally in the leaves of all plant species 
studied, but each species exhibits combinations of certain foliar xeromor- 
phic features as follows. Reduction of the leaf surface: Allium stamineum, 
Crocus ancyrensis, C. suterianus; revolute margins: Crocus ancyrensis, 
C. suterianus; unifacial leaves: Allium rotundum, A. stamineum, Gladio- 
lus atroviolaceus, Iris aphyllus; involution or folding of blade upon wilt- 
ing: Hordeum bulbosum, Poa bulbosa, Allium rotundum, Colchicum bie- 
bersteinii, Gagea arvensis, Merendera trigyna, Muscari comosum, M. 
racemosum, Orchis mascula; increased thickness of the blade: Allium 
stamineum, Crocus ancyrensis, C. suterianus; strong cutinization of epi- 
dermis: Allium rotundum, A. stamineum, Colchicum biebersteinii, Gagea 
arvensis, Merendera trigyna, Muscari racemosum, Ornithogalum arment- 


68 MADRONO LVol. 16 


acum, O.narbonense, Crocus ancyrensis, C. suterianus, Gladiolus atrovio- 
laceus; increased thickness of the epidermal cell walls: Allium rotundum, 
A. stamineum, Colchicum biebersteinu, Merendera trigyna, Ornithogalum 
armeniacum, O. narbonense, Crocus ancyrensis, C. suterianus, Orchis mas- 
cula; depression of stomata: Poa bulbosa, Allium rotundum, A. stamine- 
um, Merendera trigyna, Muscarit racemosum, Ornithogalum narbonense, 
Crocus ancyrensis, C. suterianus, Gladiolus atroviolaceus, Iris aphyllus; 
the occurrence of trichomes: Allium stamineum, Gagea arvensis, Meren- 
dera trigyna, Ornithogalum narbonense, Crocus suterianus, Gladiolus 
atroviolaceus, Iris aphyllus; epidermal water storage: Gagea arvensis, 
Limodorum abortivum, Orchis mascula; isolateral leaf: Allium rotundum, 
A. stamineum, Colchicum biebersteinu, Merendera trigyna, Muscari co- 
mosum, Ornithogalum armeniacum, Crocus ancyrensis, C. suterianus, 
Gladiolus atroviolaceus, Iris aphyllus; strongly developed palisade paren- 
chyma at the expense of spongy parenchyma: Allium stamineum, Meren- 
dera trigyna, Ornithogalum armeniacum, Crocus ancyrensis, C. suteri- 
anus; compactness of the tissues: Poa bulbosa, Allium rotundum, Meren- 
dera trigyna, Crocus ancyrensis, C. suterianus; decreased size of cells: 
Poa bulbosa, Allium rotundum, Merendera trigyna, Crocus ancyrensis, 
C. suterianus, Gladiolus atroviolaceus, Orchis mascula; water storing tis- 
sue in mesophyll: Allium rotundum, A. stamineum, Gagea arvensis, Mus- 
cart comosum, M.racemosum, Ornithogalum armeniacum, O.narbonense, 
Crocus ancyrensis, C. suterianus, Gladiolus atroviolaceus, Iris aphyllus, 
Limodorum abortivum; relative compactness of vascular bundles: Hor- 
deum bulbosum, Poa bulbosa, Allium rotundum, Colchicum biebersteinit, 
Merendera trigyna, Ornithogalum narbonense, Crocus ancyrensis, C. su- 
terianus, Gladiolus atroviolaceus; strongly developed fibers: Hordeum 
bulbosum, Poa bulbosa, Allium rotundum, Crocus ancyrensis, C. suteri- 
anus, Gladiolus atroviolaceus, Iris aphyllus. 

The leaves of Arum orientale, Gagea arvensis, Muscari comosum, Limo- 
dorum abortivum and Orchis mascula have been designated as mesoxero- 
morphic, since they exhibit structural features intermediate between meso- 
morphic and xeromorphic leaves. 

In spite of the adaptive responses of any particular tissue to the xeric 
environment, the anatomical structures of the monocotyledonous leaves 
studied are highly specialized and the pattern of tissue organization is 


quite distinct even between species of the same genus. 
Department of Biology, Mills College, California 


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REVIEWS 


Die Evolution der Angiospermen. Von PROFESSOR DR. ARMEN TAKHTAJAN, 
Leningrad. Aus dem Russischen iibersetzt von W. Hoppner, Berlin. viii -- 344 pages, 
43 figures. VEB Gustav Fischer Verlag, Jena. 1959. 

The review of this book by our fellow-member of the California Botanical Society, 
Dr. Lincoln Constance (Science 132:801. 1960), led me to order it from Messrs. 
Stechert-Hafner. It is not available in this country. Sent from East Germany, it 
arrived in about three months, with a bill for $11.65, including postage. 

About two thirds of the work may be described as prolegomena. The workings of 
evolution in general are described; as details, the frequency of neoteny (this term is 
distinguished from paedomorphosis, and preferred to it), and of parallel development, 
are noted; the hypothesis of a profound difference in mechanism between macro- 
evolution and microevolution is denied. The generally accepted hypotheses as to the 
usual directions of evolution in the plant body, the xylem, phloem, vascular anatomy 
of the node, the leaf, the flower and its parts, the fruit and seed, are duly set forth. 
The telome theory, which denies the foliar nature of stamens and carpels, is rejected. 

The flowering plants are held to be genuinely a natural group, being of a single 
evolutionary origin. The original flowering plants are believed to have been large- 
leaved trees of tropical mountains. 

Surely, the original flowering plants were homoxylous, i.e., without vessels in 
the wood. The plants now living which are homoxylous as a primitive character 
(Lemnaceae, at least, are homoxylous by reduction) are primitive in all of their 
characters. Nevertheless, these few appear to be related to widely divergent hetero- 
xylous groups. Takhtajan suggests the independent origin of vessels in at least five 
distinct lines of descent. Winteraceae, allied to Magnoliaceae, appear to represent 
the origin of the bulk of the dicots. Trochodendraceae and Tetracentraceae appear 
to lead into Amentiferae, and Amborella and Sarcandra into minor lines. The 
Nymphaeaceae appear to represent the ancestry of the monocots. Thus certain 
hypotheses maintained by Bailey and Cheadle are at the same time applied and 
convincingly supported: the presentation is elegant in the sense in which our math- 
ematical and physical colleagues use the term. 

Referring primarily to the body of opinion which has just been sketched, Con- 
stance remarks, “If in all this there is very little that is startlingly new to Western 
students of plant evolution, it is interesting to discover that the climate of opinion 
is not radically different between East and West.” I think that there is something 
more to say. The Russian botanists know everything that we know; Takhtajan cites 
Bailey and Cheadle, Gundersen and Cronquist. We, on the other hand, can not be 
certain that political and linguistic barriers have not withheld from us much pertinent 
information. How many American botanists have cited Koso-Poljansky ? 

These same opinions of Takhtajan are available also in two essays which have 
been translated into English by Mrs. Olga Hess Gankin, edited by G. Ledyard Steb- 
bins, published by the American Institute of Biological Sciences, and reviewed by 
Constance, along with the book, as cited above. In one of these translations, the 
mere skeleton of Takhtajan’s system of the flowering plants is included as an 
appendix. 

In the book, Takhtajan’s conception of the phylogeny and the system of the 
flowering plants constitutes the remaining one-third of the text. The extent of this 


1961] REVIEWS 71 


presentation makes possible a statement of the reasons which have guided him in 
the placement of each family. I did not note among the prolegomena a statement 
that no division of the flowering plants into major natural groups, beyond the 
distinction of dicots and monocots, can be made by any small number of definite 
characters; that among apetalae, choripetalae and sympetalae, between woody plants 
and herbaceous, or in any similar system, at least some of the groups are artificial. 
Neither did I find the correlative statement, that the true system is perceived by 
recognizing the connections between groups, no matter what may be the characters 
which indicate these connections. Without stating these principles, Takhtajan has 
evidently been guided by them. Some years ago, I dared predict that the main 
outlines of the natural system of the flowering plants would become essentially 
completely known within our lifetime. I consider that Takhtajan has essentially 
fulfilled this prediction. Delaying discussion of the taxonomic pattern which he 
has produced, I shall sketch the phylogeny as Takhtajan sees it, interpolating remarks, 
among which only the few references to plants which I have personally studied have 
any claim to authority. 


As noted, Takhtajan derives the Amentiferae from something of which the Tro- 
chodendraceae and Tetracentraceae are the surviving representatives; these being 
immediate derivatives of the unknown original flowering plants. The Hamameli- 
daceae-Platanaceae group is an important secondary center among the Amentiferae, 
which include also the Juglandaceae and associated families, and the odd monotypic 
American family Leitneriaceae. 

Lauraceae and their allies, including Amborella, and the Englerian Piperales, 
including Sarcandra, are treated as minor lines of descent as ancient as any among 
flowering plants; and Aristolochiaceae (among which the most ancient have apo- 
carpous flowers), with associated families, are treated as another such line. 


The remainder of the Englerian Ranales (Winteraceae, Magnoliaceae, Ranuncu- 
laceae, Berberidaceae, etc.) lead to the remainder of the dicots. A first offshoot 
includes the Centrospermae (including Cactaceae) and Papaveraceae. 


Starting again from Ranales, the Dilleniaceae-Theaceae group is recognized as 
a second node in the main line of descent of the dicots. Offshoots at this node include 
Ericales, Ebenales, a wide range of Parietales (including Salicaceae), and Malvales, 
with Euphorbiaceae in the same neighborhood. 


I was surprised to find Cyrillaceae in Ericales. Cyrillaceae have many of the 
characters of Bicornes, but seemed excluded by formal characters, notably by the 
presence of a typical endothecium in the anthers. Takhtajan’s Ericales is an extended 
group, including, with the authority of Schnarf, Saurauiaceae and Actinidiaceae. 
It now seems to me that if he had placed Cyrillaceae near these families, rather than 
near Empetraceae, he would have given satisfactory expression to the apparent 
relationships. 

Capparidaceae and Cruciferae are placed near the Parietales. I would have thought 
that Eames’ study of carpels had fixed the position of these families near Papa- 
veraceae. 


From the Dilleniaceae-Theaceae group, the main line of descent of the dicots 
goes on to a third note in Rosales sensu lato. From this group spring (1) Myrtales; 
(2) Rutales-Sapindales-Geraniales-Polygonales (I question this last) ; (3) Umbelli- 
florae; and (4) Celastrales-Rhamnales-Santalales. The great family Proteaceae, 
rather strange to us, is placed near Santalaceae. Since the Proteaceae have simple 
pistils, this association appears mistaken. 


It is generally agreed that the bulk of the Sympetalae—the old orders Contortae, 
Tubiflorae, Rubiales, and Campanulatae—belong together. Cronquist treated them 
so. Without strongly committing himself, he appeared disposed to derive them from 
the Dilleniaceae-Theaceae group. Takhtajan derives them from the neighborhood 
of Celastraceae. To us, to whom Celastraceae are known merely by cultivated plants 
of Euonymus japonica, the connection is not evident. 


MADRONO [Vol. 16 


~“TI 
bo 


A system of the monocots—derived from Nymphaeaceae, these in turn having 
as old an origin as Amentiferae, Laurales, Piperales, and the main line of the dicots— 
is duly set forth. 

So much for Takhtajan’s conception of the phylogeny of the flowering plants, 
in which only some three or four points have appeared to call for serious question. 
We shall not abandon detailed studies of species and groups; I expect, however, that 
the results of these studies will tend to strengthen the hypotheses of relationship which 
Takhtajan has set forth very much more frequently than they will tend to weaken 
them. 

In the taxonomic expression of his phylogenetic hypotheses, Takhtajan has divided 
several families; recognized numerous orders of few families (of dicots and monocots 
together, he has provided eighty-two orders, grouped in eighteen superorders) ; and 
designated every order by the stem of a generic name with the termination -ales. These 
practices are the current style or mode-trend. I deprecate them. Is a well-founded 
opinion, that some two or three families belong together, of sufficient significance to 
require expression in the category of orders? On most occasions, I think not; what 
we wish to know of each family is to what group of a dozen or more families it 
belongs. And as to names, to write Theales (or Guttiferales), Ericales, Malvales, or 
Gentianales, is to contribute to erasing from memory some of the facts of history, 
namely that these groups were known, during a period of a century or more, as the 
orders Guttiferae, Bicornes, Columniferae, and Contortae——HERBERT F. COPELAND, 
Sacramento City College, Sacramento 22, California. 


Blakeslee: The Genus Datura. By AMos G. AVERY, SOPHIE SATINA, and JACOB 
RIETSEMA. xli+289 pp., frontispiece, 67 figs, 34 tables. The Ronald Press Co., New 
York, 1959. $8.75. 

This volume is a monumental review of investigations on the genus Datura. The 
authors, all collaborators of Dr. Alfred Francis Blakeslee, internationally recognized 
expert in genetics and leader of a devoted group of associates, have given a complete 
account of more than 40 years of scientific research on this justly famous genus. 
Focused on genetical studies, the research program was one of the broadest and most 
complete ever made of plants comprising a single genus. Included is a complete Datura 
bibliography of the 228 published papers of Dr. Blakeslee and associates. An inter- 
esting feature is a historical review of the taxonomy of the genus including descrip- 
tions, figures, and notes on the ten known species included in the sections Stramonium, 
Dutra, and Ceratocaulis. Section Brugmansia, regarded by many taxonomists as a 
a separate genus, is not considered. The sole published summary of the Datura nomen- 
clature from 1753 is also included. Nearly 1,400 collections of D. stramonium, the 
most widely investigated species, from many localities scattered all over the temperate 
zones were grown and studied. 

The advanced student of genetics and the tyro alike should find much of interest 
in the preface, the foreword, and the historical review. The former will be reminded 
of the tremendous contributions to our understanding of fundamental genetical phe- 
nomena made through the investigation of one genus, of the ramifications which 
result from following up promising leads, and of the influence these researches have 
had on subsequent biological thinking and experimentation. If the tyro reads no more 
than the historical review by A. G. Avery, with its account of the medical, cere- 
monial, and cultural influences of this widely known solanaceous genus, his imagina- 
tion should be stimulated. He should come away with an appreciation of the depth 
of genetical roots as he visualizes Kolreuter, Gaetner, Naudin, Godron, de Vries, and 
Bateson at work on these puzzling plants. In this account, as well as in Dean Sinnott’s 
biographical sketch of Dr. Blakeslee, the career of a distinguished teacher, organizer, 
and investigator stands revealed in a stimulating manner, while attention is drawn to 
a magnificent cooperative venture. 

A short chapter on the alkaloids of Datura by Edward Leete refers to the chem- 
istry, distribution, pharmacology, and biogenesis of these useful substances and clearly 
indicates the need for further information at both practical and theoretical levels. 


5 


1961] REVIEWS 73 


For many readers the chapters on “Chromosome Number and Morphology,” 
“Polyploidy,” and “Extra-Chromosomal Types” will strike responsive chords. They 
will be reminded of the use of Belling’s iron-acetocarmine technique, of plant breed- 
ing methods, of the production of induced polyploids, and of the use of cold, injury, 
radiation, growth substances, and various other chemicals, including the pioneering 
work on the use of colchicine, as aids in experimentation. Interesting and valuable 
ideas were developed through combined cytological and genetical examination of the 
polyploids and extra-chromosomal types. Segmental interchange and ring formation, 
comparisons of the effects of single additional chromosomes with the duplication of 
complete sets of chromosomes, and the evolution of genetic systems come readily to 
mind as one flips the pages concerned with the details of these and other important 
phenomena. 

Only two allelomorphic pairs of characters had been described prior to 1919 when 
Blakeslee and Avery reported the first of numerous mutations observed by the Cold 
Spring Harbor group. The total number of known gene pairs approaches 550 and, 
among plants, is probably exceeded only by those in Zea. These mutations affect all 
parts of the plant, have appeared either spontaneously or, more often, as a result of 
a variety of treatments, and are discussed at some length in the chapter on “Gene 
Mutations.” Because of the enormous amount of work required, relatively little at- 
tempt was made to locate most of the genes. Nevertheless, genes responsible for 81 
distinct mutations representing twelve linkage groups have been located. Methods for 
constructing maps of the several chromosomes are described. Certain markers have 
been very useful in various investigations of particular interest, including those inves- 
tigations affecting male and female sterility. 

Miss Satina’s short, compact, and beautifully illustrated and organized chapter 
on “Chimeras’”’ deserves careful reading. Here emphasis on the advantage of utilizing 
chimeras for obtaining information on the histogenesis of plant structures merits 
reiteration. The controversy on the origin, nature, and organization of apical meri- 
stems and other initials has engaged the attention of many botanists over a long 
period of years. Perhaps further study of chimeras along the lines suggested by the 
Datura work will help elucidate this perennial problem and will also offer evidence 
for the diverse functions of the epidermis which have been suggested. The use of 
periclinal chimeras as a tool in investigating a variety of morphogenetic problems 
should receive wider recognition. 

A chapter on “Radiation Experiments” by A. G. Avery and Jean Cummings gives 
considerable detail on both methods and results. In this review, already becoming too 
lengthy, the chapter summary seems to be an efficient device for giving its essence: 

In Datura, as in so many other organisms, radiations have been very use- 
ful as inducers of genic and chromosomal mutants, and these in turn have 
been helpful in clarifying the answers to many questions of morphogenesis 
and physiology. On the other hand, it has been possible to use the knowledge 
gained from irradiated plants in the comparison of the effectiveness of dif- 
ferent types of radiations--thermal and fast neutrons from various sources. 
To a great extent the conclusions seem clear-cut. All the radiations so far 
tested seem to cause the same types of both genic and chromosomal aberra- 
tions, but the effectiveness of the different radiations is quite different. Neu- 
trons cause much greater effects, both genic and chromosomal, than do either 
X-rays or gamma rays for equivalent energies. 


Geneticists have long since learned the need for a thorough knowledge of the life 
history of any experimental organism and the Datura team is no exception. Clear 
descriptions of the growth and activity of the tissues associated with reproduction, 
fertilization, and of the development of the embryo, endosperm, and seed coats are 
given by Satina and Rietesema in chapters 10 and 11. 

“Barriers to Crossability: Prefertilization” and “Barriers to Crossability: Post- 
fertilization” are two important chapters full of suggestions for future research. An 
inventive and imaginative approach to many experimental problems is presented. 


74 MADRONO [Vol. 16 


Extensive studies of pollen viability, germination, and pollen-tube growth are sum- 
marized. The fate of embryos and endosperm in incompatible crosses, the growth of 
ovular tumors, and the physiological aspects of seed abortion are among the topics 
discussed. 

In later years special attention has been paid to the vital link between genera- 
tions, the seed. Problems of incompatability, sterility, and abortion required attention 
and led to the development of a method for the culture of embryos. Many observa- 
tions have led to a partial understanding of the numerous complex processes going on 
simultaneously in the growing seed. Here are dozens of unsolved problems for which 
the Datura investigators suggest promising lines of attack. 

In the chapter on “Segmental Interchanges and the Species Problem,” Miss Satina 
assembles considerable material on prime types, racial differences, interspecific hy- 
brids, and the characteristics of hybrids from incompatible crosses. The problems of 
chromosome analysis, ring formation, and of chromosome-end arrangements are 
examined. The abundant occurrence of segmental interchange present in the various 
races of Datura is unusual, but the condition is known in some other plants. In spite 
of intensive study, the exact relationship of the phenomenon to speciation remains 
obscure. We may agree with Blakeslee when he says that, “The frequency of inter- 
change of chromosomal fragments in D. stramonium and the relation of this phenom- 
enon to the formation of new pure-breeding types has led to the hypothesis that 
segmental interchange has accompanied the changes responsible for the formation of 
species in the genus Datura. Nevertheless, in spite of very intensive study the rela- 
tionship of the phenomenon to speciation remains obscure.’”’ We may hope that some 
day Datura will be a valuable instrument in helping us work out the relationship 
between genes and chromosomes which will further our understanding of the evolu- 
tionary picture. 

The scientific world owes its gratitude to Smith College and the National Science 
Foundation for contributing assistance, facilities, and finances towards the completion 
of this work. Congratulations are due the Ronald Press for its part in this fine enter- 
prise, for the volume is pleasing in all aspects. High praise is due the committee of 
the Genetics Society of America which catalyzed the reaction which resulted in the 
publication of this labor of love. 

Attention should be called to page three of the volume, which carries an invitation 
to investigators interested in securing material of the Jimson weed in order to add 
further chapters to our knowledge of the members of this fascinating genus ——ALTON 
H. Gustarson, Department of Biology, Bowdoin College, Brunswick, Maine. 


Vascular Plants of the Pacific Northwest. By C. Leo HitcHcock, ARTHUR CRON- 
QUIST, MARION OwnBeEy, and J.W. THompson. Illustrated. University of Washing- 
ton Press. Part 5, pp. 1-343. 1955. $7.50. Part 4, pp. 1-510. 1959. $12.00. 


Present or future students of the Pacific Northwest flora will find their time well 
spent in carefully looking through the two volumes now available of the projected 
five-volume “Vascular Plants of the Pacific Northwest.” It is a credit to the authors 
that they have drawn on their wide experience in the western flora to point out and 
discuss specific problems such as unusual variation patterns, possible hybridization, 
disjunct or vicarious distributions, and a host of other phenomena which suggest a 
number of areas requiring the attention of biosytematists, plant geographers, gene- 
cologists, and cytologists. This is merely a bonus added to a sound taxonomic treat- 
ment of the 4000 vascular plants (upon completion) either native or introduced in 
“all of Washington, the northern half of Oregon, Idaho north of the Snake River 
Plains, the mountainous portion of Montana, and an indefinite southern fringe of 
British Columbia.” The area circumscribed is a natural floristic unit and excludes 
most of the interesting but large Great Basin flora occurring in the southern portions 
of Idaho and Oregon as well as the sizable cluster of endemic or California-centered 
species characteristic of southwestern Oregon. 

The two volumes published to date are Parts 4 and 5 of the series: Part 5 is a 


1961] REVIEWS eS 


monograph of the Compositae by Arthur Cronquist and Part 4 includes treatments 
by Cronquist (Polemoniaceae through Campanulaceae, except Castilleja), C. Leo 
Hitchcock (Ericaceae through Cuscutaceae), and Marion Ownbey (Castilleja). Al- 
though J. W. Thompson has not yet contributed texts, his valuable and extensive 
collections in the Northwest serve as a substantial basis for the present knowledge of 
the northwestern flora, hence his inclusion as a co-author is fully justified even in 
the absence of such textual material. The families are arranged in the Englerian 
sequence; the genera and species within a genus are arranged alphabetically. This 
practice eliminates the need for an index, although one to the synonyms of the 
larger genera and to common names is included. The chief difficulty with this alpha- 
betical arrangement is that closely related species and genera are placed together 
only rarely, thus rendering a character-by-character comparison of two taxa (as is 
often done during the course of identification) rather more difficult. Taxonomic con- 
cepts are admittedly conservative, which results in the volumes being attractive for 
purposes of identifying species of such name-ridden genera as Aster, Castilleja, 
Erigeron, and Senecio. 

In Part 5, Cronquist outlines his philosophy of taxonomy, which we assume to 
be the philosophy of the other authors as well in the absence of any statements to 
the contrary. His discussion is worth reading since he presents particularly well his 
views as an “orthodox” taxonomist on some of the attitudes held in the past by 
experimentally oriented workers. The term variety is preferred to subspecies when 
only one infraspecific level is recognized. In a few instances where it has been necessary 
to utilize two infraspecific levels, variety is subordinate to subspecies; forma is not 
used. It is a reflection of the authors’ conservatism that infraspecific categories are 
widely used for entities which other workers prefer to recognize as species. In attempt- 
ing to conform to their system, numerous changes in rank of many taxa have been 
made, involving transfers from subspecies to variety. Fortunately, these changes have 
been rendered somewhat inconspicuous by their placement among the synonvmy of 
the species. The synonymy of each species is laudably complete and gives the full 
bibliographical citation, the collector, locality, and date of collection for the type of 
each svnonym. 

The volumes issued to date have served as vehicles for the publication of 
three highly localized species: Chaenactis thompsonii, Hackelia davisti, and Luina 
serpentina—apovropriately endemic to Washington, Idaho, and Oregon respectively. 
In addition to these new svecies, numerous varieties are described for species of manv 
genera. This emphasizes the fact that the Pacific Northwest is far from completely 
known botanically. In addition to these novelties, numerous changes of rank and 
transfers from one taxon to another are made. Hence this work can hardly be called 
an automatic compilation; rather it is an original, critical treatment of all the 
species it covers. The geographical ranges of the taxa are specifically outlined and, 
where known, ecological data regarding the habitat are included as well. 

Although the authors have made extensive use of existing taxonomic monographs 
(manv of which are cited in the text), they have obviously felt no obligation to 
follow these works in their own treatments. Undoubtedly thev will receive criticism 
for their handling of many groups, but the citation of monographs and the lists of 
synonvms will make it easy for those who do disagree on some count to find other 
interpretations of the group in question. The Menyanthaceae and Gentianaceae are 
considered separate families; on the other hand, the Lobeliaceae are included in the 
Camnanulaceae. Genera of the Vacciniaceae, Pvrolaceae, and Monotropacere are 
found within the Ericaceae. lpomopsis is relegated to Gilia and Cacaliopsis nardosmia 
will be found as a Luina. 

The work serves as an interesting chronicle of the fate of many weedy introduc- 
tions into the area. For instance, we Jearn that Anthemis mixta L. and A. altissima L. 
have both been collected only on the famous ballast heaps at Portland and pre- 
sumably have failed to become members of the naturalized flora of the Pacific 
Northwest. This is quite different behavior from that of their cogeners Anthemis 


76 MADRONO [Vol. 16 


cotula L. and A. arvensis L., which have been extremely successful in becoming estab- 
lished in the region as weeds. Conyza floribunda H.B.K. and Tournefortia sibirica L. 
have apparently only persisted at their point of introduction and are not expanding 
in range. But by far the largest number of weeds have become widely distributed in 
the region and most, if not all of these are discussed in the text. These different 
patterns of behavior subsequent to introduction suggest a fertile field for gene- 
cological studies. 

Notes on the cultivation and ornamental value of many of the native species 
included in Part 4 have been supplied by two accomplished Seattle horticulturalists: 
Carl English and Brian O. Mulligan. These men and a number of other northwestern 
gardeners have’ demonstrated the high desirability of native plants in the garden 
when they are grown properly. The ornamental value of the northwestern flora is 
not as widely appreciated in this country as it deserves to be; it is probably easier 
to purchase seeds of Northwest Pacific Coast indigens from British and European 
nurserymen than from American ones. It is true that attractive species such as 
Gaultheria shallon, Arctostaphylos uva-ursi, and Arbutus menziesii are rather widely 
grown, yet too few gardeners are aware of the potentialities of equally attractive 
species such as Menziesia ferruginea, Vaccinium ovatum and V. parvifolium, and 
Polemonium carneum. The majority of these species can be propagated or obtained 
in a manner which does not involve denudation of the countryside. Understandably, 
horticultural notes have not been included for the Compositae, whose contribution 
trom the temperate zone of a large and diverse assemblage of weeds largely outweighs 
its contribution of a depauperate ornamental flora. 

Chromosome numbers are included for most of the taxa in which they are known, 
although the literature sources for these numbers are not given. In the instances of 
circumboreal or polytypic species, or of polyploid complexes, it would be valuable to 
know the geographical source of the plants on which the counts are based. 

Despite the necessity of using the same type throughout the entire work, the 
format of the book is enlivened by consistent use of indentations, capital letters, and 
underlinings for various kinds of information repeatedly appearing in each species 
description. Part 5 appears to be free of typographical errors; there are, however, 
several errors in Part 4 but none has been found which seriously impairs the mean- 
ing or usefulness of any section of the book. Each species discussed is illustrated ; 
the Compositae were drawn by Dr. John Rumely and the taxa in Part 4 were done 
by Jeanne R. Janish, who is well known in the west for her work on Abrams’ 
“Tllustrated Flora of the Pacific States.” Both artists have provided a felicitous com- 
bination of accurate scientific illustration with esthetically pleasing artistry. Dr. 
Rumely’s drawings lack some of the three-dimensional qualities of Mrs. Janish’s and 
are more obviously based on herbarium specimens; this may, however, prove to be 
something of a virtue in view of the high likelihood that most botanists would prefer 
to bring their specimens back from the field in press, rather than carry these bulky 
volumes with them in the field. 

The keys are quite usable and generally include a number of characters which 
can be utilized in identifying an unknown species. Since the series is not yet complete, 
no glossary of terms or key to the families is provided. However, the publishers 
promise a family key in Part 1, and the authors promise that “a more nearly natural 
arrangement of the families and orders of dicotyledons will be presented at the begin- 
ning of Part 2.” The high caliber of the work that has so far appeared makes us 
eagerly await the completion of this valuable contribution to the knowledge of the 
flora of the Pacific Northwest.—RosBERT ORNDUFF, Department of Botany, University 
of California, Berkeley. 


INFORMATION FOR CONTRIBUTORS 


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the author assume the complete cost of publication. 

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

Institutional abbreviations in specimen citations should follow Lanjouw 
and Stafleu’s list (Index Herbariorum. Part 1. The Herbaria of the World. 
Utrecht. Second Edition, 1954). 

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

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ADRONO 


VOLUME 16, NUMBER 3 JULY, 1961 


Contents 


PAGE 
THE GENUS LEPIDIUM IN CANADA, 
Gerald A. Mulligan 77 


ESCHSCHOLZIA COVILLEI GREENE, A TETRAPLOID 
SPECIES FROM THE MOJAVE DESERT, 


Theodore Mosquin 91 
ABNORMAL FRUITS AND SEEDS IN ARCEUTHOBIUM, 

Frank G. Hawksworth 06 
To ALBERT W. T. C. HErRRE, /ra L. Wiggins 102 


CHROMOSOME COUNTS IN THE GENUS MIMULUS 
(SCROPHULARIACEAE), Barid B. Mukherjee and 
Robert K. Vickery, Jr. 104 


SPHENOPHYLLUM NYMANENSIS SP. NOV. FROM THE 
Upper PENNSYLVANIAN, J. F. Davidson 106 


A NEw NAME IN THE ALGAL GENUS PHORMIDIUM, 
Francis Drouet 108 


NoTES AND NEwS: PLAGIOBOTHRYS AUSTINAE (GREENE) 
JounstTon: A NEw ADDITION TO THE OREGON FLORA, 
Francia Chisaki and Robert Ornduf; 

STEGNOSPERMA CUBENSE AND GOSSYPIUM KLOTZSCHIA- 
NUM DAVIDSONII NOT KNOWN IN THE REVILLAGIGEDOS, 
Reid Moran 108 


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 
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 
MitpreD E. Martuias, University of California, Los Angeles 24 
Marion OwNnBEY, State College of Washington, Pullman 
REED C. ROLLINS, 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—JoHN H. THoMAS 
Dudley Herbarium, Stanford University, Stanford, California 


1961] MULLIGAN: LEPIDIUM 77 


THE GENUS LEPIDIUM IN CANADA'™ 


GERALD A. MULLIGAN 


A world monograph of the genus Lepidium was published by Thellung 
(1906) and a further study of the genus by Hitchcock (1936). Accord- 
ing to Hitchcock, Thellung did not have enough North American material 
at his disposal for an accurate interpretation of our plants. This prompted 
Hitchcock’s comprehensive treatment of the genus Lepizdium in the 
United States. However, Hitchcock did not see any material from 
Canadian herbaria and relatively few Canadian specimens were repre- 
sented in his material from United States herbaria. Consequently it was 
not surprising to find on studying Canadian specimens that some of the 
taxa present in Canada are not included in even the most recent floras or 
lists. Apparently as the result of not having many Canadian specimens 
for study, Hitchcock included L. bourgeauanum, a common plant in the 
Canadian prairies, under L. ramosisstmum. He erroneously applied Thel- 
lung’s name, L. bourgeauanum, to another plant, L. densiflorum var. 
bourgeauanum. Specimens of L. heterophyllum, a species introduced 
from Europe, and previously unreported for North America, were found 
in the material studied. 

This paper includes keys to all the Lepidium present in Canada and a 
description and discussion of each taxon. The life durations given are 
mostly based on information obtained by growing plants in nursery plots 
at Ottawa. The chromosome numbers given for Canadian material were 
determined from somatic root tip cells. The root tips studied were pro- 
cessed as in Mulligan (1959). Distribution maps (figs. 12 and 13) were 
prepared by mapping all the herbarium specimens seen, except where 
localities were closely duplicated. 

A total of 935 herbarium specimens, exclusive of duplicates, was ex- 
amined from the following Canadian herbaria: Department of Agricul- 
ture, Ottawa (DAO); National Museum of Canada, Ottawa (CAN); 
British Columbia Provincial Museum, Victoria (V); University of 
British Columbia, Vancouver (UBC). Type specimens were obtained 
from the Gray Herbarium, Harvard University, Cambridge (GH), the 
New York Botanic Garden, New York (NY); also seen were McCabe’s 
British Columbia collections from the University of California, Berkeley 
(UC). I wish to express my appreciation to the curators of these herbaria 
for the loan of material. I am also indebted to workers at the Plant Re- 
search Institute, Canada Department of Agriculture, Ottawa, for their 
encouragement and assistance in this study. 


LEPIpIuM L., Sp. Pl., 643. 1753; Gen. Pl. 291. 1754. 
Annual to perennial herbs, glabrous to hirsute with simple hairs. 


1 Contribution 79 from the Plant Research Institute, Research Branch, Canada 
Department of Agriculture, Ottawa, Ontario. 


Maprono, Vol. 16, No. 3, pp. 77-108. July 12, 1961. 


78 MADRONO [Vol. 16 


Flowers small, white to sulfur yellow in dense terminal racemes. Sepals 
usually somewhat pubescent along back. Petals lacking, or to twice length 
of sepals. Stamens 2, 4 or 6. Ovary with 2 ovules, style short, stigma 
capitate, sometimes 2 lobed. Fruit a dehiscent silicle, strongly keeled or 
winged (silicle indehiscent, not keeled or winged in closely related Car- 
daria). Usually one seed attached to the apex of each cell. 


Key TO THE SPECIES OF LEPIDIUM IN CANADA 


a. Middle and upper leaves suborbicular, deeply cordate clasping with a closed sinus 
and slightly overlapping lobes, thus appearing as if perfoliate . 1. L. perfoliatum 
aa. Middle and upper leaves narrower, linear to broadly lanceolate, if clasping, not 
appearing as if perfoliate. 
b. Silicles 5 to 6 mm. long. 
c. Middle and upper leaves clasping the stem, silicles on spreading pedicels. 

d. Annual or biennial with usually a single erect stem; anthers yellow; silicles 
covered with small white vesicles, style included to slightly exserted from 
shallow apical notch... . , 3 6 2a aleseanipesine 

dd. Perennial with numerous ascending ian mth violet; silicles with few 
or no vesicles, style mostly exserted from shallow apical notch. 

3. L. heterophyllum 
cc. Middle and upper leaves not clasping, silicles on strongly ascending to ap- 
pressed pedicels .-“..8 @ 68) ee ee ete earziniy 

bb. Silicles 2 to 3.5 mm. long. 
e. Glaucous perennial 50 to 130 cm. high, with rhizomes; leaves thickish and 


rugose, lanceolate to broadly lanceolate . . . .  . 5.2L. latifolium 
ee. Annual or biennial, 5 to 40 cm. high, leaves not thickish and rugose, linear to 
lanceolate. 


ft. Silicle bidentate at apex, the sinus well developed and broad, its projecting 
shoulders abruptly contracted into Messe divergent teeth, pedicels sig- 
moid. . . . Ss gus “PUD Ds oxycarpum 
ff. Silicles merely vanes: or ean A apex ani a shallow sinus, narrowed to 
abruptly curved into apical teeth; pedicels straight to arching. 
g. Silicles puberulent, at least on margin. 

h. Silicles 2.5 to 3 by 1.5 to 2 mm., nearly elliptic, narrowed into acute 
apical teeth; inflorescence congested into numerous axillary racemes 
as well as terminal ones. .. .  .  . 11. L. ramosissimum 

hh. Silicles 3 to 3.5 by 2.5 to 3 mm., oes obcordate to short oblong- 
obovate, rounded to abruptly Aaeeed into obtuse apical teeth; in- 
florescence a single raceme or of sparsely branched racemes 

9. L. densiflorum 
gg. Silicles glabrous. 

i. Silicles oval, orbicular to rotund; petals conspicuous, as long or slightly 


longer than sepals . .. .  . 8, L. virginicum 
. Silicles ovate, obovate to round seondate: petals shorter than sepals 
or lacking. 


j. Silicles ovate to obovate, narrowed into acutish apical teeth. 
k. Middle and upper cauline leaves blunt tipped, lower cauline and 
rosette leaves bipinnatifid, petals absent . . 6. L. ruderale 
kk. Middle and upper cauline leaves acute tipped, lower cauline and 
rosette leaves incised, oe present, usually about half length of 
sepals .  .. .  . 10. L. bourgeauanum 
jj. Silicles round apeord ate to ee obovate, rounded to abruptly 
curved into obtuse apical teeth . . . . 9.2L. densiflorum 


1961] MULLIGAN: LEPIDIUM 79 


1. LEPIDIUM PERFOLIATUM L., Sp. Pl., 643. 1753. 

Annual or winter annual with single erect stem 1—5 dm. high, sparsely 
hairy, usually branched above; lower leaves bipinnate, the middle and 
upper leaves suborbicular, deeply cordate clasping; petals pale yellow, a 
little longer than the sepals; stamens usually 6; silicles usually glabrous, 
rhombic-ovate, on spreading-ascending pedicels, nearly as broad as long, 
3-4 mm. long and 3—4 mm. broad; pedicels terete; style usually project- 
ing beyond the shallow apical notch. 2n = 16 (voucher: grown at Ottawa 
from seed collected at Lethbridge, Alberta, Mulligan 1527, DAO, fig. 1). 

Rare along roadsides and in waste places in Ontario, Saskatchewan 
and Alberta. Occasional along roadsides in the Okanagan Valley of British 
Columbia and rare elsewhere in the Province (fig. 12). This plant, intro- 
duced from Eurasia, was first collected in Canada at Cranbrook, British 
Columbia, in 1931. 


Representative material seen. ONTARIO: York County, at county line of Ontario 
County, Shumovich 976 (DAO). SASKATCHEWAN: Swift Current, Budd in 1937 
(DAO). ALBERTA: Lethbridge, Bzbbey 12 (DAO). BRITISH COLUMBIA: Cran- 
brook, Groh in 1931 (CAN) ; Kelowna, McCalla 11598 (UBC, V) ; Osoyoos, Lindsay 
é& Woodbury 1128 (DAO). 


2. LEPIDIUM CAMPESTRE (L.) R. Br., Ait. Hort. Kew, ed. 2,4:88. 1812. 

Annual to biennial with dense short spreading hairs throughout, stem 
usually solitary, erect, 2-6 dm. high, branched above the middle, the 
branches ascending; lower leaves entire or lyrate, narrowed into a short 
petiole, the middle and upper leaves narrowly triangular, sessile, clasping 
the stem with long narrow pointed basal lobes; petals white, a little longer 
than sepals; stamens 6 with yellow anthers; pedicels spreading, slightly 
flattened; silicles densely covered with small white vesicles that become 
scale-like when dry, silicles oblong-ovate, 5-6 mm. long and 4 mm. broad: 
style included to slightly exserted from the shallow apical notch. 2n = 16 
(voucher: grown at Ottawa from seed collected in southwestern Ontario, 
Mulligan 1499, DAO, fig. 2). 

Common in fields, roadsides and waste places in southern Ontario, 
Quebec and British Columbia. Sporadic along roadsides and in waste 
places in Newfoundland, Prince Edward Island, Nova Scotia, New Bruns- 
wick and Alberta (fig. 12). Introduced from Eurasia. 


Representative material sen. NEWFOUNDLAND: Gander, Bassett 383 (DAO). 
PRINCE EDWARD ISLAND: Souris, Kings County, Erskine and Smith 2046 
(DAO); Charlottetown, Dore & Gorham 45.314 (DAO). NOVA SCOTIA: South 
Sydney, Cape Breton, Macoun in 1886 (CAN); Kentville, Lewis in 1944 (DAO) ; 
Mabou, Smith et al 8669 (DAO). QUEBEC: Grosse-Ile, Comté de Montmagny, 
Marie-Victorin et al 40129 (CAN); Mont-Rolland, Marie-Anselm 14 (DAO); 
Bristol, Bassett and Mulligan 1140 (DAO) ; Montreal, Bernard in 1952 (CAN, UBC). 
ONTARIO: Milton West, Mulligan and Lindsay 818 (DAO); Snelgrove, White in 
1897 (CAN); St. Thomas, James 2478 (DAO); Kemptville, Lindsay and Bassett 
213 (DAO); Port Arthur, Garton 2339 (DAO). ALBERTA: between Macleod and 
Pincher, McCalla 11070 (DAO). BRITISH COLUMBIA: Chilliwack, Faris 32 
(DAO); Koksilah, V.I., Tice in 1937 (UBC, V); Sandspit, Moresby Island, Queen 
Charlotte Islands, Calder 21111 (DAO). 


80 MADRONO [Vol. 16 


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Fics. 1-11. Somatic chromosomes of Lepidium, camera lucida drawings, * 2150. 
1, L. perfoliatum, 2n=16; 2, L. campestre, 2n—=16; 3, L. latifolium, 2n—=24; 4, L. 
virginicum (eastern material), 2n=32; 5, L. virginicum (western material), 2n=32; 
6, L. densiflorum var. densiflorum, 2n = 32; 7, L. densiflorum var. macrocarpum, 
2n=32; 8, L. densiflorum var. elongatum, 2n=32; 9, L. densiflorum var. pubicarpum, 
2n=32; 10, L. bourgeauanum, 2n=32; 11, L.ramosissimum, 2n=64. 


3. LEPIDIUM HETEROPHYLLUM (DC.) Benth., Cat. Pl. Pyr. 95. 1826. 
L. smithu Hook., Brit. Fl., ed. 3, 300. 1835. 

Perennial herb with short spreading hairs on leaves and stem; stems 
many, ascending, 1.5—4.5 dm. high, often branched below as well as above 
the middle, the branches ascending; lower leaves oblanceolate or elliptical, 
narrowed into a short petiole, the middle and upper leaves narrowly 
triangular, sessile, clasping the stem with long narrow basal lobes; petals 
white, a little longer than sepals; stamens 6, anthers violet; pedicels 
spreading, slightly flattened; silicles with vesicles lacking or few, oblong- 
ovate, 5-6 mm. long and 4 mm. broad; style mostly exserted from the 
shallow apical notch. 2n—16, European material (FI. Brit. Isles, 175. 
1952). 

Occasional along roadsides, in fields and waste places on Vancouver 
Island, British Columbia (fig. 12). This plant was first collected near 
Victoria in 1908. Lepidium heterophyllum, introduced from Europe, was 
first recognized as occurring in North America by Dr. C. Frankton in 


1961] MULLIGAN: LEPIDIUM 81 


1956 when he identified a specimen, collected near Courtenay, British 
Columbia, as L. smithiz. 


Material seen. BRITISH COLUMBIA. VANcouvER ISLAND: vicinity of Victoria, 
Macoun, May 20, 1908 (CAN), June 19, 1908 (CAN); Telegraph Bay, Copley 6657 
(V); Mt. Finlayson, Copley 6658 (V); Alberni, Carter 2196 (V); Millstream Road, 
Hardy 7558 (V); S. Saanich, Newcombe 8917 (V); Sooke, Hardy 22768 (V); 
Courtenay, Molyneux 73 (DAO, UBC, V); 2 miles east southeast Langford, Calder 
et al 20795 (DAO). 


4, LEPIDIUM SATIVUM L., Sp. Pl., 644. 1753. 

Annual with a solitary erect stem 2—8 dm. high, glabrous; lower leaves 
long-stalked, lyrate with toothed obovate lobes, the middle and upper 
leaves pinnatipartite or bipinnatipartite, occasionally entire and linear; 
petals white or reddish, up to twice as long as the sepals; stamens 6; 
silicles glabrous, broadly elliptical or nearly orbicular, 5—6 mm. long and 
3—4 mm. broad; pedicels appressed, flattened; style not projecting beyond 
the deep apical notch. 2n—16, European material (Jaretzky 1932). 

Rare along roadsides and in waste places in Prince Edward Island, 
Nova Scotia, New Brunswick, Quebec, Ontario, Saskatchewan, Alberta, 
British Columbia and Yukon Territory (fig. 12). Introduced from Eur- 
asia as early as 1882 but still only a casual escape from cultivation. 


Representative material seen. PRINCE EDWARD ISLAND: 4 miles northwest 
Charlottetown, Campbell 150 (DAO). NOVA SCOTIA: Harbourville, Kings County, 
Lewis in 1944 (DAO). NEW BRUNSWICK: St. Quentin, Groh in 1937 (DAO). 
QUEBEC: Ste. Annes des Monts, Gaspé, Macoun in 1882 (CAN). ONTARIO: 
Ottawa, Scott in 1890 (DAO, CAN). SASKATCHEWAN: Yorktown, Macoun and 
Herriot in 1906 (CAN). ALBERTA: Beaverlodge, Brooks in 1930 (DAO). BRITISH 
COLUMBIA: Victoria, Newcombe 9259 (V); Nelson, Eastham 3065 (UBC). 
YUKON TERRITORY: Dawson, Macoun in 1902 (CAN). 


5. LEPIDIUM LATIFOLIUM L. Sp. Pl., 644. 1753. 

Perennial herb with subterranean rhizomes, each branch of the rhizome 
giving rise to a single erect stem 5—13 dm. high, glabrous, much branched 
above; lower leaves long-petioled, simple and ovate with a toothed margin 
or pinnately lobed with a large terminal and 2 or more smaller lateral 
lobes, the lobes all rounded, the middle and upper leaves sessile, ovate 
or ovate-lanceolate, acute, entire or with distant teeth, the uppermost 
leaves often bract like and white margined near the apex; petals white, 
up to twice as long as sepals; stamens 6; silicles glabrous to pubescent, 
elliptical to orbicular, 2 mm. long and 2 mm. broad; pedicels ascending, 
terete; style very short with large rounded stigma, apical notch very 
slight or lacking. 2n = 24 (voucher: grown at Ottawa from seed collected 
at Lethbridge, Alberta, Mulligan 2147, DAO, fig. 3). 

This plant, introduced from Eurasia, was first collected in 1934 but 
has remained localized around Quebec City and Lethbridge, Alberta 
(igs 12). 


Representative material. QUEBEC: Quebec, Marie-Anselm in 1934 (DAO). AL- 
BERTA: Lethbridge, Moss in 1940 (CAN). 


82 MADRONO [Vol. 16 


L. campestre | 


SRG ye ia 3 
Woy , 
- 7 
. / 
, 
b A 


ae ee 


Sao) 
ny 


' 
aca Ses 


Fic. 12. Distribution maps of Lepidium. 


1961] MULLIGAN: LEPIDIUM 83 


6. LEPIDIUM RUDERALE L., Sp. Pl., 643. 1753. 

Annual to biennial with single erect or ascending stem 1—3 dm. high, 
plant almost glabrous, with occasionally a few short spreading hairs; 
stem branched above, the branches ascending; lower leaves long-petioled, 
deeply bipinnately divided into narrow entire segments, the middle and 
upper leaves sessile, narrowly oblong, entire, rounded at apex; petals 
usually absent; stamens usually 2; silicles glabrous, ovate or broadly 
elliptical, 2-2.5 mm. long and 1.5—2 mm. broad; pedicels spreading to 
ascending, slightly flattened; style at base of the shallow apical notch. 
2n = 32, European material (Jaretzky 1932). 

Rare along roadsides and in waste places in Nova Scotia, New Bruns- 
wick, Quebec, Ontario and Manitoba (fig. 12). Introduced from Eurasia 
as early as 1868. 

Representative material seen. NOVA SCOTIA: North Sydney, Macoun in 1883 
(CAN). NEW BRUNSWICK: Bass River, Kent County, Fowler in 1868 (CAN). 
QUEBEC: Montreal, Rolland-Germain 46008 (DAO, CAN). ONTARIO: Welling- 
ton, Montgomery and Shumovich 997 (DAO). MANITOBA: Winnipeg, Frankton 
and Bibbey 60 (DAO). 

7. LEPIDIUM OXYCARPUM T. & G., Fl. N. Am. 1:116, 688. 1838. 

Slender, nearly glabrous annual, 0.5-2 dm. high, with many semi- 
erect stems branched above the middle; lower leaves linear, often with 
2—4 pairs of linear lobes, middle and upper leaves usually linear and 
entire; petals white, rudimentary; stamens 4; silicles on slender some- 
what sigmoid and flattened pedicels; silicles ovate, glabrous, and finely 
reticulate, 2.5-3.5 mm. long and 2—2.5 mm. broad, abruptly contracted 
at apex into a pair of widely divergent teeth; style at base of large apical 
notch. 

The only Canadian specimen of L. oxycarpum seen (fig. 12) was col- 
lected at Cadboro on Vancouver Island (Macoun in 1893, CAN). 


8. LEPIDIUM VIRGINICUM L. sensu lat., Sp. Pl., 645. 1753. 

Annual, freely branched, erect to spreading, 1.5—-6 dm. high, glabrous 
to strongly pubescent; lower and middle leaves irregularly toothed or 
incised to pinnatifid, the divisions often again dissected, the upper leaves 
much reduced, usually entire or remotely toothed; petals white, equalling 
to much longer than the sepals; stamens usually 2; silicles glabrous, 
oval, orbicular to rotund, 2.5-4 mm. long and 2.5—4 mm. broad; pedicels 
spreading to ascending, slightly flattened to terete; stigma included in 
the shallow apical notch. 2n = 32 (vouchers: grown at Ottawa from seed 
collected at St. Thomas, Ontario and Saanichton Spit, British Columbia, 
Mulligan 2420 and 2421, DAO, figs. 4 and 5). 

In Canada, L. virginicum sensu lat. is represented by eastern and 
western elements (fig. 12). The positions of the cotyledons in the seeds 
of Canadian material, as in the United States material (Hitchcock 1936), 
are accumbent in eastern plants and oblique to incumbent in western 
plants. Eastern plants occur sporadically in Newfoundland, Prince Ed- 
ward Island, Nova Scotia, New Brunswick, Quebec and Ontario. These 


84 MADRONO [Vol. 16 


plants are L. virginicum var. virginicum and are introduced from further 
south in the eastern United States. Western plants of L. virginicum 
sensu lat. are found only on Vancouver Island, and the adjacent islands 
and mainland. They are undoubtedly native to this area. Most of these 
western plants have morphological characters tending towards the varie- 
ties pubescens and medium as treated by Hitchcock (1936). However, 
it appears that two and possibly three varieties of L. virginicum sensu lat. 
come together in the southwestern corner of British Columbia and at this 
northern limit of their range, there is extreme morphological variability 
in the population. An understanding of the British Columbia plants would 
require an extensive study of all the western North American material of 


L. virginicum sensu lat. Such a study is outside the limits of this treat- 
ment. 


Representative material seen. NEWFOUNDLAND: St. John’s, Green 1517 (DAO). 
PRINCE EDWARD ISLAND: Charlottetown, Erskine 2332 (DAO). NOVA 
SCOTIA: Wolfville, Groh in 1932 (DAO). NEW BRUNSWICK: Fredericton, Dore 
and Gorham 45165 (DAO). QUEBEC: Shawinigan Falls, Groh in 1927 (DAO, 
CAN). ONTARIO: near St. Thomas, Macoun in 1907 (CAN). BRITISH CO- 
LUMBIA: Saanich Spit, Eastham in 1939 (DAO, UBC); Parksville, Vancouver 
Island, Carter 2195 (V); Jessie Island, Departure Bay, Macoun in 1908 (CAN) ; 
Mitlenatch Island, Sweeney 15567 (V). 


9. LEPIDIUM DENSIFLORUM Schrad. sensu lat., Ind. Sem. h. Gotting. 4. 
1832. 

Annual to biennial, puberulent to pubescent; stem erect, 1—5 dm. 
high, usually branched above the middle, sometimes simple; lower leaves 
mostly oblanceolate, coarsely toothed to pinnatipartite, the divisions also 
toothed, the middle and upper cauline leaves reduced, slightly toothed 
or entire; petals white, rudimentary to sometimes equalling the sepals 
in western varieties; stamens 2; silicles glabrous to puberulent in some 
of western varieties, round-obcordate to short oblong-obovate, rounded 
to abruptly curved into obtuse apical teeth, 2-3.5 mm. long and 1.5 to 
3 mm. broad; pedicels slightly ascending to nearly appressed, slightly 
to conspicuously flattened; stigma included in the narrow apical notch. 


KEY TO VARIETIES OF L. DENSIFLORUM 


a. Silicles averaging 2.5 mm. long, glabrous; pedicels slightly flattened, crowded, 
more than 9 pedicels per cm. Bra Fe ee os Qa. var. densiflorum 
aa. Silicles averaging 3-3.5 mm. long, puberulent except in var. macrocarpum,; 
pedicels conspicuously flattened, less crowded, usually less than 9 pedicels per cm. 


b.Silicles glabrous . . . . . . . .  . «Ob. var. macrocarpum 
bb. Silicles puberulent. 
c. Silicles puberulent only on margins . . . .  . 9c. var. elongatum 
cc. Silicles uniformly puberulent . . . .  .  . = 9d. var. pubicarpum 


9a. L. DENSIFLORUM Schrad. var. DENSIFLORUM. L. densiflorum var. 
typicum Thell., Bull Herb. Boiss., ser. 2, 4:706. 1904. 

Plant erect, 1-5 dm. high, annual or winter annual with glabrous 
silicles, averaging 2.5 mm. long and 2 mm. broad, smaller than all western 


1961] MULLIGAN: LEPIDIUM 85 


ie shine var. densiflorum 


L. _densiflorum vor. pee L. densiflorum var. 


’- macrocarpum J 
‘ NA ea | 


o 


; “a 
L. bourgeauanum 
bS ° x 
XN 


Fic. 13. Distribution maps of Lepidium. 


86 MADRONO [Vol. 16 


varieties. 2n = 32 (voucher: grown at Ottawa from seed collected at 
Ottawa, Mulligan 1528, DAO, fig. 6). 

Widely distributed in all types of disturbed habitats: Newfoundland, 
Prince Edward Island, Nova Scotia, New Brunswick, Labrador, Quebec, 
Ontario, Manitoba, Saskatchewan, Alberta, British Columbia, Yukon 
Territory and Mackenzie District, Northwest Territories (fig. 13). Na- 
tive to the Prairie Provinces, interior of British Columbia and probably 
some localities in eastern Canada. Weedy throughout its range. 


Representative material sen. NEWFOUNDLAND: Gander, Bassett 462 (DAO). 
PRINCE EDWARD ISLAND: Bideford, Prince County, Smith 319 (DAO) ; French 
River, Fernald et al 7508 (CAN). NOVA SCOTIA: Boylston, Hamilton in 1890 
(CAN); Wolfville, Groh in 1928 (DAO). NEW BRUNSWICK: Point du Chene, 
Bassett and Mulligan 2964 (DAO); Woodstock, Macoun in 1899 (CAN) ; Edmuns- 
ton, Malte 332 (CAN). LABRADOR: Goose Bay, Gillett and Findley 5883 (DAO, 
UBC). QUEBEC: Nominique, Lucien in 1924 (CAN); Magog, Bassett and Hamel 
2322 (DAO); Shawville, Mulligan and Lindsay 382 (DAO); Ville Marie, Baldwin 
5940 (CAN). ONTARIO: Leamington, Macoun in 1901 (CAN) ; Moosonee, Baldwin 
1453 (CAN); Goderich, Senn et al 4759 (DAO); Point Pelee, Bassett 1112 (DAO). 
MANITOBA: Douglas, Lindsay 490 (DAO); Duck Mountain, Scoggan and Baldwin 
7793 (CAN); Fort Ellice, Macoun in 1879 (CAN); The Pas, Krivda 1223 (DAO). 
SASKATCHEWAN: Dundurn, Campbell 54 (DAO); Prince Albert, Macoun in 
1876 (CAN); Cypress Hills, Breitung 5001 (DAO); Bjorkdale, Van Blaricom in 
1941 (DAO). ALBERTA, 7 miles north Fort Fitzgerald, Cody and Loan 3863 
(DAO); 20 miles west Selba, McCalla 12313 (UBC); Fort Saskatchewan, Turner 
41873 (CAN). BRITISH COLUMBIA: 141 Mile House, Cottle in 1949 (UBC); 
Grand Forks, Tice in 1933 (V); Yahk, Bassett and Cumming 3970 (DAO). NORTH- 
WEST TERRITORIES: Fort Simpson, Cody and Matte 8109 (DAO); Alexander 
Falls, Hay River, Lewis 558 (DAO). YUKON TERRITORY: Watson Lake, 
Gillett 2585 (DAO). 


9b. L. DENSIFLORUM Schrad. var. macrocarpum var. nov. L. densiflor- 
um var. bourgeauanum sensu Hitchcock, Madrono, 3:279, 1936, nec L. 
bourgeauanum Thellung. 

Herba biennis erecta, saepius 1-3 dm., siliculis glabris, 3.0-3.5 mm. 
long., 2.5-3.0 mm. lat., 2n = 32 ex canadensibus. 

Plant erect, 1-3 dm. high, biennial with glabrous silicles 3.0-3.5 mm. 
long and 2.5-3.0 mm. broad. 2n = 32 (voucher: grown from seed col- 
lected at Cache Creek, Mulligan 2416, DAO, fig. 7). 

Native on dry open soil in western Saskatchewan, Alberta and British 
Columbia, as far north as Prince George, British Columbia (fig. 13). 

Type. Lethbridge, Alberta, Platieres de la riviére Sainte-Marie pres 
de son embouchure, 23 juin 1958, Boivin, Perron and Harper 12197 
(DAO), fig. 14. 


Material seen. SASKATCHEWAN: Webb, 7 miles au nord, Boivin et al 12005 
(DAO); Saskatchewan Landing, Russell S58099 (DAO); 7 miles au sud de la 
Station Expérimentale de Manyberries, Boivin and Alex 9651 (DAO). ALBERTA: 
1 mile east of Canmore, south of Peace River, Macoun in 1903 (CAN); Canyon 
Creek, Boivin and Perron 12744 (DAO); Lethbridge, Bozvin and Perron 12166 
(DAO). BRITISH COLUMBIA: Tranquille, Groh 246 (DAO); Lillooet, Luyat in 
1928 (V), Anderson 2197 (V), Macoun in 1916 (CAN); Kamioops, Davidson in 
1912 (UBC), Tisdale 40-410 (DAO), Wattie in 1915 (UBC) ; Kamloops, Thompson 


1961] MULLIGAN: LEPIDIUM 87 


River Flats, Brink in 1935 (UBC), a mixture of 1 plant var. macrocarpum and 2 
plants var. elongatum,; Spences Bridge, AZacoun in 1899 (CAN); Hamilton Com- 
monage, Nicola Valley, Zisdale in 1935 (DAO), 40-409 (DAO); Cache Creek, 
Mulligan and Woodbury 1617 (DAO); Hat Creek Valley, Thompson and Thompson 
221 (DAO); Yahk, Bassett and Cumming 3988 (DAO); Cecil Lake, Merten in 
1958 (DAO); Riley’s Ranch, Big Bear Creek, Copley 6430 (V) ; Fairmont, Anderson 
225 (V); Merritt, Copley 7312 (V); Nelson, Eastham 3057 (UBC); Macalister, 
Taylor and Lewis 286 (UBC); Prince George, Eastham 14735 (UBC); Nanaimo, 
Eastham 3058 (UBC); Lytton, Dawson in 1876 (CAN); Crow Nest Pass, Macoun 
in 1897 (CAN); 2% miles south Merritt, McCabe 4523 (UC); 21% miles south 
Williams Lake, McCabe 1312 (UC). 


9c. L. DENSIFLORUM Schrad. var. ELONGATUM (Rybd.) Thell., Bull. 
Herb. Boiss., Ser. 2, 4:706. 1904; Monog. Lepid. 235. 1906. L. elongatum 
Rydb., Bull. Torr. Bot. Club, 29:234. 1902. L. simile Heller, Bull. Torr. 
Bot. Club, 26:312. 1899. 

Plant erect, 1-3 dm. high (rarely taller), biennial with silicles puberu- 
lent only on margins, 3—3.5 mm. long and 2.5—3 mm. broad. 2n = 32 
(voucher: grown at Ottawa from seed collected at Ashnola River, Flat- 
iron Mountain, British Columbia, Mulligan 2422, DAO, fig. 8). 

Native on dry open soil in interior of British Columbia and as far 
north as Kamloops. Apparently also native along the Mackenzie River 
in Yukon Territory and in the northwestern corner of British Columbia 
(fig. 13). 

Representative material seen. BRITISH COLUMBIA: 1 mile east Fort Steele, 
Calder and Savile 9149A (DAO); Fernie, Bassett and Cumming 3986 (DAO); 
Goodfellow Creek, Hardy 18.875 (V); Revelstoke, Macoun in 1890 (CAN) ; Windy- 
Arm, Yukon Boundary, Gervaise in 1914 (UBC); 2 miles north Skookumchuck, 


McCabe 5031 (UC). YUKON TERRITORY: Carcross, Gillett 3384 (DAO) ; White- 
horse, Gillett 3508 (DAO); island in Klondike River, Macoun in 1902 (CAN). 


Od. L. DENSIFLORUM Schrad. var. PUBICARPUM (Nelson) Thell., Bull. 
Herb., Boiss., Ser. 2, 4:706. 1904; Monog. Lepid., 235. 1906. L. pubi- 
carpum Nelson, Bot. Gaz. 30:189. 1900. 

Plant erect, 1-3 dm. high (rarely taller), annual or winter annual 
with puberulence scattered over all of silicle, 3-3.5 mm. long and 2.5-3 
mm. broad. 2n = 32 (voucher: grown at Ottawa from seed collected at 
Osoyoos, British Columbia, Mulligan 2412, DAO, fig. 9). 

Known to occur in Canada only around Osoyoos and Penticton, 
British Columbia (fig. 13). 


Material seen. BRITISH COLUMBIA: 19 miles east Osoyoos, Mulligan and 
Woodbury 2010 (DAO); Osoyoos, Lindsay and Woodbury 630 (DAO); Penticton, 
Eastham 3056 (UBC), 7067 (UBC); Okanagan Valley at U.S. Boundary, McCabe 
5848 (UC). 


10. LEPIDIUM BOURGEAUANUM Thell., Monog. Lepid., 237, 1906. L. 
fletchert Rydb., Bull. Torr. Bot. Club, 34:428. 1907. 

Biennial, 1.5—6 dm. high, sparsely to densely puberulent throughout; 
stem erect, with many ascending to nearly appressed branches bearing 
usually less than 5, rarely up to 10 racemes; lower leaves incised, middle 


[Vol. 16 


— 


MADRONO 


88 


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1961] MULLIGAN: LEPIDIUM 89 


leaves incised or sometimes slightly toothed; upper leaves linear, entire, 
rarely slightly toothed; petals white, up to 34 length of the sepals; 
stamens 2; silicles glabrous, ovate to obovate, 2.5—-3 mm. long, and 1.5—2 
mm. broad; pedicels spreading to ascending, slightly flattened; style 
included in the apical notch. 2n = 32 (vouchers: grown at Ottawa from 
seed collected at St. Simeon, Province Quebec, Alexander Falls and 
Norman Wells, Mackenzie District, Mulligan 2423, 2418 and 2419, DAO, 
fig. 10). 

Fairly common on open soil in Manitoba, Saskatchewan, Alberta, 
British Columbia, Yukon Territory and Northwest Territories, and 
probably native in all these areas. It also occurs at a few locations in 
Newfoundland, New Brunswick, Ontario and Quebec, where it has 
probably been introduced (fig. 13). 

Lepidium bourgeauanum, described by Thellung (1906), was based 
on a collection of Bourgeau [Saskatchewan, 1857-8, Bourgeau (Pal- 
lisers Brit. N. Am. Exped.)—Herb. Petersburg|. Hitchcock (1936) 
applied this name to his L. densiflorum var. bourgeauanum, a plant that 
is relatively rare on the Canadian prairie. However, Thellung’s descrip- 
tion obviously refers to the plant here being treated (see fig. 14), not 
Hitchcock’s L. densiflorum var. bourgeauanum. A Bourgeau specimen 
[labelled Lake Winnipeg Valley, 1857 (Pallisers Brit. N. Am. Exped.) | 
in the Gray Herbarium, Harvard University, is L. bourgeauanum. This 
specimen is possibly an isotype of L. bourgeauanum with more complete 
label data than the holotype in the Petersburg Herbarium. L. bourgeau- 
anum has been included under L. ramosissimum by most botanists, but 
in addition to the differences in morphology and geographic distribution, 
the former plant has 32 somatic chromosomes while the latter plant 
has 64. 

Representative material seen. NEWFOUNDLAND: Deer Lake, Rouleau 1160 
(DAO). NEW BRUNSWICK: 2 miles north northeast Edmunston, Madawaska 
County, along railroad tracks, Mulligan and Spicer 2538 (DAO), not mapped on 
fig. 13. ONTARIO: Prescott, Grenville County, single plant near grain elevator, 
Dore 18299 (DAO), not mapped on fig. 13. QUEBEC: 2 miles west St. Simeon, 
Bassett and Hamel 2190 (DAO); Ellis Bay, Anticosti Island, Johansen in 1923 
(CAN). MANITOBA: Lake Winnipeg Valley, Bourgeau in 1857 (GH, possibly 
isotype of L. bourgeauanum) ; Brandon, Macoun in 1896 (CAN) ; Churchill, Beckett 
3852 (DAO); Winnipeg, Fletcher in 1905 (DAO, isotype of L. fletcheri). SAS- 
KATCHEWAN: Cherryfield, Macoun and Herriot 69881 (CAN, paratype of L. 
fletcheri) ; Dana, Senn et al 2745 (DAO); 16 miles west Saskatoon, Macoun and 
Herriot in 1906 (CAN) ; Lee’s Lake Reservoir, Bird 1560 (DAO). ALBERTA: Fort 
McMurray, Cody and Gutteridge in 1953 (DAO) ; Beaverlodge, Jenkins 123 (DAO) ; 
Calgary, Macoun in 1897 (CAN). BRITISH COLUMBIA: Sinkut Lake, Eastham 
16959 (UBC, V); 54 miles south Williams Lake, Mulligan and Woodbury 1776 
(DAO). NORTHWEST TERRITORIES. Mackenzie District: Wrigley Harbour, 
Brabant Island, Lewis 998 (DAO) ; 2 miles east Trout River, Cody and Matte 8637 
(DAO) ; Indian Village on north shore of Mackenzie River, Cody and Matte 8622 
(DAO). YUKON TERRITORY: West Dawson, Calder and Billard 4627 (DAO). 


90 MADRONO [Vol. 16 


11. LEPIDIUM RAMOSISSIMUM Nelson, Bull. Torr. Bot. Club, 26:124, 
1899. L. ramosissimum var. robustum Thell., Monog. Lepid., 236. 1906. 

Biennial, 1-4 dm. high, sparsely to densely puberulent; stem erect, 
usually profusely branched throughout, with many spreading to ascend- 
ing branches bearing usually more than 10, occasionally as few as 5 
racemes; lower and middle leaves sessile, pinnately or bipinnately parted; 
upper cauline leaves usually with at least one pair of linear lobes towards 
the apex, rarely entire; petals white, up to 34 length of the sepals; 
stamens 2; silicles puberulent, at least along margins, ovate to obovate, 
2.5—3 mm. long and 1.5—2 mm. broad; pedicels spreading to ascending, 
slightly flattened; style included in the apical notch. 2n = 64 (vouchers: 
grown at Ottawa from seed collected at Stirling and Edmonton, Alberta 
and Yellowknife, Mackenzie District, Mulligan 2129, 2424 and 2417, 
DAO, fig. 11). 

Fairly common on open soil in Manitoba, Saskatchewan and Alberta; 
rare in western Ontario, British Columbia and Mackenzie District, North- 
west Territory. Native in the Prairie Provinces, but possibly introduced 
elsewhere (fig. 13). The first Canadian collection seen was collected by 
Bourgeau, at Fort Ellice, Manitoba, in 1857. 


Representative material seen. ONTARIO: Schreiber, Hosie et al 689 (CAN). 
MANITOBA: Snowflake, Bassett and Kemp 3504 (DAO) ; Norway House, off north 
end of Lake Winnipeg, Scoggan 4233 (CAN); Churchill, Beckett 3 (DAO); Buttes 
de Sables au Fort Ellice, Bourgeau in 1857 (GH). SASKATCHEWAN: Scott, Groh 
in 1933 (DAO); Twelve-Mile Lake, Wood Mountain, Macoun in 1895 (CAN); 
Saskatchewan, Bourgeau in 1858 (GH, isotype of L. ramosissimum var. robustum) ; 
Scott, Groh in 1933 (DAO). ALBERTA: Edmonton, Frankton 895 (DAO); Fort 
Saskatchewan, Turner 4948 (DAO, UBC); Craigmyle District, Brinkman in 1921 
(CAN); Frank, Bassett and Cumming 3975 (DAO). BRITISH COLUMBIA: 
Windermere Slough, Columbia Valley, Eastham 16288 (V, UBC); Windermere, 
McCabe 6365 (UC); Fernie, Bassett and Cumming 3971 (DAO). NORTHWEST 
TERRITORIES. Mackenzie District: Yellowknife, Cody and McCanse 3045 
(DAO). 

Plant Research Institute, Research Branch, 
Canada Department of Agriculture, Ottawa, Canada 


LITERATURE CITED 


Hitcucock, C. L. 1936. The genus Lepidium in the United States. Madrono 3:265- 
820; 

JarETzKy, R. 1932. Beiziehungen zwischen Chromosomenzahl und Systematic bei 
den Cruciferen. Jahrb. wiss. Botan. 76:485-527. 

Mutuican, G. A. 1959. Chromosome numbers of Canadian weeds. II. Can. Jour. Bot. 
37:81-92. 

THELLUNG, A. 1906. Die Gattung Lepidium (L.) R. Br. Eine monographische Studie. 
Mitteilungen bot. Mus. Univ. Zurich. 28:1-340. 


1961) MOSQUIN: ESCHSCHOLZIA 91 


ESCHSCHOLZIA COVILLEI GREENE, A TETRAPLOID SPECIES 
FROM THE MOJAVE DESERT' 


THEODORE MOSQUIN 


The purpose of this paper is to establish the validity of Eschscholzia 
coville: Greene (Papaveraceae) as a taxon of specific rank on the basis 
of a comparative study of morphological variation in relation to chromo- 
some number and geographical distribution. Eschscholzia covillei is one 
of a group of closely related taxa in the deserts of southwestern United 
States and adjacent Mexico that has frequently been treated as con- 
specific with E. minutiflora Watson (e.g., Jepson, 1922, 1925; Munz, 
1935; Abrams, 1944; Munz, 1959). 

Lewis and Snow (1951) pointed out that E. minutiflora is hexaploid 
(n=18) and that a diploid species, F. parishii Greene (n=—6), for- 
merly considered a variety of E. minutiflora, is readily distinguishable 
from the latter on morphological grounds. They also pointed out that 
plants intermediate between these two taxa in Inyo County, California, 
might be tetraploid and genetically distinct from both E. minutiflora and 
E. parishii. This suggestion was confirmed in 1957 when a collection of 
the intermediate material from the White Mountains (Lew7s 1084) was 
determined to be tetraploid (n=12). More recently Ernst (1959) has 
reported the tetraploid number of chromosomes for two collections from 
the same area (Ernst 561, 564). From study of my own collections, I 
have found that these intermediate specimens are consistently tetraploid 
and morphologically distinguishable from both the diploid, E. parishiz, 
and the hexaploid, FE. minutiflora. Consequently the tetraploid should 
be recognized as a distinct species. An examination of the literature and 
of the type specimens concerned indicates that the earliest specific name 
for the tetraploid is /. coviller Greene. This was clearly designated on 
the United States National Herbarium sheet (number 3340) by Greene. 


ESCHSCHOLZIA COVILLEI Greene, Pittonia 5:275. 1905. Type: from 
Pete’s Garden to 1000 feet below, Johnson Canon, Panamint Mountains, 
Inyo County, California, elevation 1700 meters, Coville & Funston 519 
(US). E. minutiflora var. darwinensis M. E. Jones, Contr. West. Bot. 
8:2-3, 1898. Type: on mesas, Darwin, Inyo County, California, Jones 
in 1897 (POM). 

Glabrous annual herb, to 40 cm. tall, freely branched throughout; 
basal rosettes well-developed with leaves coarsely divided, numerous, 


1T am grateful to Dr. Harlan Lewis for suggesting this problem to me and for 
critical review of the manuscript. Special thanks are due to Dr. Peter H. Raven 
for his assistance in checking types and for other helpful suggestions. I also wish to 
thank Dr. Richard Snow for permission to publish his previously unreported chro- 
mosome number determinations, and for permission to examine the specimens in their 
care the curators of the following herbaria: the University of California, Berkeley ; 
Pomona College; Rancho Santa Ana Botanic Garden; and the San Diego Museum 
of Natural History. 


G2 MADRONO LVol. 16 


glaucous, 6-13 cm. long, the blade 0.5—4.5 cm. long, 0.5—-4 cm. wide; 
upper leaves strongly reduced; mature buds elliptical, 6-9 mm. long, acu- 
minate; pedicels 1-8 cm. long; torus turbinate; petals obovoid-cuneate, 
golden-yellow, 7-17 mm. long; stamens 8-15 per flower, 3.5—5 mm. long; 
pollen with 7-10 grooves (usually 8 or 9), 24-37 microns in diameter; 
seeds with finely reticulate grey-brown coat; chromosome number, 
Tice 7 

Distribution. Slopes and washes of desert mountains, Inyo and San 
Bernardino counties, California (fig. 1). 

Representative specimens. CALIFoRNIA. Inyo County: Panamint Valley, 11 miles 
southwest of Ballarat on road to Ridgecrest, Mosquin & Lewis 3241 (LA, UC); 
0.7 mile from junction to Darwin on road to Darwin Falls, Mosquin & Lewis 3251 
(LA, UC); Panamint Valley, 7.2 miles east of junction to Trona on road to Stove- 
pipe Wells, Mosquin & Lewis 3255 (LA, UC); Emigrant Canyon, Mosquin & Lewis 
3256, 3257 (LA, UC); 0.6 mile west of Bradbury Well entrance to Death Valley 
National Monument, Mosquin & Lewis 3258-1 (UC); Westgard Pass road, Lewis 
1084 (LA) ; Nelson Range, Austin in 1906 (UC) ; Pleasant Canyon, Panamint Moun- 
tains, Hall & Chandler 6965 (UC); Hole-in-the-Rock Spring, Epling et al. in 1930 
(LA, UC); Hanaupah Canyon, Panamint Mountains, in 1922 (collector unknown, 
SD); Shepherds Canyon, Argus Mountains, Keller 126 (SD) ; Black Canyon, White 
Mountains, Duran 2668 (LA, UC); Bishop Creek, 5,200 feet, Hall & Chandler 7249 
(UC); Darwin, 4,600 feet, Jones, April 28, 1897 (POM); from Pete’s Garden to 
1060 feet below, 1,700 meters, Coville & Funston 519 (US). San Bernardino County: 
7 miles east of Daggett, Munz & Keck 7843 (POM); 10 miles southwest of Garlic 
Springs, Munz & Keck 7878 (POM). 

Eschscholzia covillei is usually readily distinguishable from EF. minuti- 
flora (table 1), especially when the two are found in adjacent or mixed 
colonies. Where they occur in mixed colonies the two are distinguished 
by flower size and habit. It is perhaps more difficult to distinguish 
E. covillei from E. parish, but the two are not known to grow together 
(fig. 1). In general, the latter two differ consistently in stamen number 
and in the number of grooves on the pollen. The specimens from San 
Bernardino County that are identified as /. covillei are geographically 
closest to E. parishii and it would be desirable to have additional chromo- 
some number determinations from this area in order to confirm their 
identification. The hexaploid species, E. minutiflora, also grows sym- 
patrically with FE. parishii, and in such localities plants of the two species 
are readily distinguished, as is also true of most herbarium specimens, 
by the larger flowers and greater stamen number of E. parishi. All three 
species are found on comparatively moist alluvial slopes and fans, but 
unlike the other two species, the hexaploid E. minutiflora extends onto 
the desert floor. 

Plants of Eschscholzia parishi from near Randsburg, Kern County 
(Lewis & Mosquin 1117; Heller 7683), the only locality for this species 
on the Mojave Desert, are intermediate in several morphological traits 
between FE. parishii from the Colorado Desert and E. covillei. They may 
have as few as 14 stamens per flower, and they have an intermediate 
pollen morphology and stamen number. In the Heller collection, pollen 


1961] MOSQUIN: ESCHSCHOLZIA 


93 
\y i 
gh ae aon eas NEVADA | / | Me 
e q al {3 ’ 
Pe eee aie. a 
Ye ee \ ie 
wkd ih jr 1 4 ; i He eer 
San Francisco-—~— : : a raceme | - er 
St Vite Heo at a a eae 
me 
\ 
} | 
| % 
' 
' 
= = 
— | : es 
—— 


i i ae 


—_—_—_—_—_—_——+4 


ARIZONA 


Eschschoizia 


BAJA CALIFORNIA 
minutiflora 


covillet 
parishit 


Fic. 1. General distribution 
E. covillei and E. parishii. 


~ 
oes 
aw 
<a 
2 
: 
| 


\ 
} 
| 


of Eschscholzia minutiflora and selected localities of 


94 MADRONO [Vol. 16 


size varied from 17—21 microns, and the two buds examined had, respec- 
tively, 14 and 15 stamens. The pollen had 7 and 8 grooves in approxi- 
mately equal frequencies. In the Lewis and Mosquin collection, for 
which the chromosome number has been determined, pollen diameter 
varied from 20-32 microns, the pollen had 7, rarely 6, grooves per grain, 
and the stamen number varied from 16-18 per flower. In view of these 
morphological traits this might be considered a local subspecies of E. 
parishit. 

The chromosomes of all three species are similar in size and morphol- 
ogy, with ring bivalents being more frequent than rods. The following 
list includes all known chromosome counts? made in this group: 


ESCHSCHOLZIA PARISHII (n = 6). CALIFORNIA. Kern County: about 2 miles south- 
east of Searles Station, Lew7zs & Mosquin 1117 (3 plants, 1 count from somatic cells). 
Riverside County: Morongo Wash, Snow 11 (5 plants); about 5 miles west of road 
to Cottonwood Springs on United States Highway 60, Lewis & Ernst in 1949 
(3 plants)»; Joshua Tree National Monument, Lewis in 1949»; Box Canyon, 
Snow 7%; Raven 11478 (somatic count). 


ESCHSCHOLZIA COVILLEI (n= 12). CaLtForNIA. Inyo County: Mosquin & Lewis 
3241 (2 plants), 3251 (3 plants), 3255-3, 3255-4 (2 plants), 3256 (2 plants), 3258-1; 
Lewis 1084; Westgard Pass road, 2.8 miles west of Zurich, Ernst 561°, 1.6 miles west 
of Zurich, Ernst 5644, 


EsCHSCHOLZIA MINUTIFLORA (n= 18). CaLiForRNIA. Imperial County: road to 
17 Palms, 0.3 miles east of United States Highway 99, Lewis in 1952. Inyo County: 
Panamint Valley, 15 miles north of road to Darwin on road to Stovepipe Wells, 
Mosquin & Lewis 3240; 4.6 miles from junction of State Highway 190 with road to 
Darwin Falls, Mosquin & Lewis 3249, 3250 (total of 3 plants); 4.2 miles north of 
road to Darwin Falls on road to Darwin Springs, Mosquin & Lewis 3254; Panamint 
Valley, 7.2 miles east of junction to Trona on road to Stovepipe Wells, Mosquin & 
Lewis 3255-1, 3255-2 (2 plants) ; 0.6 mile west of Bradbury Well entrance of Death 
Valley Natl. Mon., Mosquin & Lewis 3258-2; east of Darwin, Snow 26 (approxi- 
mate count)®; just below Darwin Falls, Raven 12114. Kern County: 2 miles south- 
east of Searles Station, Lewzs & Mosquin 1117-4. Los Angeles County: 0.5 mile 
north of Pearblossom, Mosquin 3265. Riverside County: about 5 miles west of road 
to Cottonwood Springs on United States Highway 60, Lewis & Ernst in 1949 
(3 plants); road to Cottonwood Springs, 7.2 miles north of United States Highway 
60, Snow 102; Box Canyon, Snow 51*. San Bernardino County: 0.8 mile north of 
Atolia, Lewis in 1950; about 2 miles west of Lucerne Valley, Snow 12%; 10.2 miles 
east of Barstow, Snow 232%; United States Highway 395, 22 miles south of Inyo 
County line, Snow 25-12; 15.9 miles south of Kramer Junction, Lewis & Mosquin 
1114; 2 miles north of Needles, Raven 13891 (approximate count). San Diego County: 
Mason Valley, near Vallecito Station, Ernst 258°. Baya CALIForNIA, Mexico. 14.8 
miles south of Mexican Highway 2 on road to San Felipe, Raven 11630. 


Eschscholzia minutiflora also occurs in the South Coast Ranges of 
California (Axelrod 260, UC; Axelrod 9170, POM; Schreiber 1045, UC). 
The identification of this species is based on morphological considera- 


2 Counts by Snow (unpublished) indicated by *, those of Lewis & Snow, by ?, 
those of Ernst, 1958, by ©, and those of Ernst, 1959, by *. Vouchers for chromosome 
number determinations not previously reported are on file in the herbarium, Univer- 
sity of California, Berkeley, or in the herbarium, University of California, Los 
Angeles. The first set of my own collections are deposited at the herbarium of the 
University of California at Berkeley. 


1961] MOSQUIN: ESCHSCHOLZIA 95 


TABLE 1. MORPHOLOGICAL COMPARISON OF THREE SPECIES* OF ESCHSCHOLZIA 


E. parishi E. covillei E. minutiflora 
(i==6) (n==12)) (n=18) 
Habit (rosette) Poorly Well de- Well developed 
developed veloped (Colorado Desert) 
or lacking 


(Mohave Desert) 


Habit (branching) Slender, Much- Much-branched 
delicate branched 
Length of longest petals 8-22 7-18 4-10 


(range in mm.) 


Length of mature buds 7-16 6-9 2-7 
(range in mm.) 


Bud apex Acuminate Acuminate Blunt (Colorado Des- 
ert or acuminate 
(Mohave Desert) 


Number of stamens 16-37 8-15 4—15 
(range) 

Length of longest stamens 4-7.5 3.5-5 2-4 
(range in mm.) 

Number of pollen 5.5-7 7.5-9.1 8.2-10.4 
grooves** 


(range of means) 


Diameter of pollen 20-32 24-37 25-44 
(range in microns) 


* Only considering plants from which the chromosome number has been deter- 
mined, 9 of EF. parishii, 12 of E. covillei, and 27 of E. minutiflora. 
** Mean of each plant based on 10 grains. 


tions. One plant (Axelrod 9170) which was examined in detail had only 
10 stamens per bud, a pollen diameter of 40 to 44 microns and usually 
11 rarely 10 grooves per pollen grain. I have examined the pollen of the 
diploids E. californica Cham., E. caespitosa Benth., E. glvptosperma 
Greene, and E. californica var. peninsularis (Greene) Munz, and have 
found these plants to have a pollen variation comparable to E. parishii 
and out of the range of the pollen of E. minutiflora. A comparison of the 
pollen traits of E. minutiflora in the South Coast Range to those of 
E. parishii as shown in table 1 can leave little doubt that these Coast 
Range plants are hexaploid. The presence of this desert hexaploid in 
dry areas of the South Coast Ranges is not too surprising for a similar 
pattern of distribution is known for other desert annuals, e.g. Linanthus 
parryae (Gray) Greene, Streptanthella longirostris (Wats.) Rydb., Erio- 


96 MADRONO LVol. 16 


gonum trichopes Torr., Chaenactis xantiana Gray and Salvia colum- 


bariae Benth. 
Department of Botany, 
University of California, Los Angeles 


LITERATURE CITED 
ABRAMS, L. R. 1944. Illustrated flora of the Pacific States. Vol II. Stanford. 
Ernst, W. R. 1958. Chromosome numbers of some western Papaveraceae. Contr. 
Dudley Herb. 5:109-115. 
. 1959. Chromosome numbers of some Papaveraceae. Contr. Dudley Herb. 
5:137-139. 
Jepson, W. L. 1922. A flora of California, Vol. I. Berkeley. 
. 1925. Manual of the flowering plants of California. Berkeley. 
Lewis, H., and R. Snow. 1951. A cytotaxonomic approach to Eschscheltzia. 
Madrono 9:141-143. 
Muwnz, P. A. 1935. Manual of southern California botany. Claremont, Calif. 
. 1959. A California flora. Berkeley. 


ABNORMAL FRUITS AND SEEDS IN ARCEUTHOBIUM! 


FRANK G. HAWKSWORTH 


The normal Arceuthobium fruit, as described in the literature (Thoday 
and Johnson 1930, Dowding 1931, Gill 1935, Kuijt 1955, 1960), consists 
of a single seed containing one embryo. This paper describes abnormal 
fruits with two seeds and seeds with two embryos and endosperms as 
found in some specimens of A. americanum Nutt. ex Engelm. and 4. 
vaginatum {. cryptopodum (Engelm.) Gill. 

The fruit of Arceuthobium and other members of the Loranthaceae 
differs from other angiosperms in that there are no true ovules. The 
ovarian cavity becomes nearly filled by an undifferentiated mound of 
tissue termed the mamelon, nipple, or ovarian papilla. Two embryo sacs 
are borne within the ovarian papilla. Usually only one embryo sac de- 
velops, but occasional diembryonic seeds have been reported in a number 
of species (Peirce 1905, Weir 1914, and Heinricher 1915). The process 
of fertilization in Arceuthobium has not been precisely described. How- 
ever, the development of the embryo sac after fertilization is apparently 
similar to that in most dicotyledonous plants. As the fruit matures, the 
dominant embryo sac develops into a copious endosperm with a small 
embryo. The remnants of the ovarian papilla become crushed, and in 4. 
pusillum they form a distinct ‘“‘crest”’ at the base of the seed (Thoday and 
Johnson 1930). The crest was not well defined in the mature, normal 4. 
americanum (fig. 1A) and A. vaginatum f. crvptopodum fruits examined. 
However, a small mass of tissue which is presumed to be analogous to the 


1 Acknowledgment is expressed to Job Kuijt, Department of Biology and Botany, 
University of British Columbia, for reviewing the manuscript and to William Schacht, 
School of Forestry, Duke University, for providing some of the abnormal fruits 
described. 


1961] HAWKSWORTH: ARCEUTHOBIUM 97 


A. AMERICANUM A. vaGINATUM 


MILLIMETERS 


Fic. 1. Semi-diagrammatic drawings of longitudinal sections through Arceuthobium 
americanum fruits. A. Normal fruit with a single seed containing one embryo. The 
tissues labeled are: v.c., viscin cells; per., pericarp; e., endocarp of the seed; emb., 
embryo; end., endosperm; a.J., abscission layer; and ped., pedicel. B-F. Abnormal 
fruits; these are described in the text. 


crest in A. pusillum was observed in most fruits. At maturity the Arceu- 
thobium fruit is severed from its pedicel, and the seed is forcibly ejected. 
ABNORMAL FRUITS 


Fruits of A. americanum with more than one stigma (figs. 1, 2) were 
noticed from plants in several areas of the Medicine Bow National Forest 


98 MADRONO [Vol. 16 


Fic. 2. Fruits of Arceuthobium americanum: normal, D and H; abnormal, A-C, 
E-G. Scale below is millimeter rule. 


in southern Wyoming and the Roosevelt National Forest in northern 
Colorado. In a sample of 803 fruits from one locality in the latter forest, 
seven, or 0.9 per cent, had two stigmas. Dissection of 16 abnormal fruits 
collected in August revealed four general types. 

Type 1. Fruits with two stigmas and two normal seeds (fig. 1B; fig. 
2F and 2G). A wall of tissue separating the two seeds was sometimes 
present (fig. 1F) and sometimes not (fig. 1B). Nine of the sixteen fruits 
dissected were of this type. 

Type 2. Fruits with two stigmas, one normal seed and one aborted seed 
(fig. 1C and 1D; fig. 2A and 2C). Four specimens had a small aborted 
seed (fig. 1C), but only one was found with two full-sized chambers 
(fig. 1D). 

Type 3. Fruit with one stigma but two normal seeds (fig. 1E; fig 2B). 
Only one such fruit was found. 

Type 4. Fruit with three stigmas and two normal seeds (fig. 1F; fig. 
2E). One of the seeds had two embryos. Only one fruit of this type was 
found. 

The abnormal fruits averaged about the same length as normal ones 
(3.6 mm.) but were about 50 per cent wider (2.7 compared with 1.9 mm.). 
The seeds from the multiple fruits measured 0.9 x 2.1 mm. compared 
with 1.0 x 2.2 mm. for seeds from normal fruits on the same plants. 


1961] HAWKSWORTH: ARCEUTHOBIUM 99 


MILLIMETERS 


Fic. 3. Seeds of Arceuthobium americanum and A. vaginatum ft. cryptopodum. 
The upper seed in each column is normal and of average size for the species. The 
lower three seeds in each column represent abnormal forms with two elements, each 
with an endosperm (light stippling) and embryo (dark stippling) within a common 
endocarp. 


Intensive search yielded only one fruit of A. vaginatum f{.cryptopodum 
with more than one stigma. This fruit, from near Estes Park, Colorado, 
had three stigmas and three distinct chambers. Two of these, the outer 
ones, contained normal seeds, but the central chamber had an apparently 
aborted seed (similar to that in the right chamber in fig. 1D). 

Fruits with multiple stigmas have not been reported previously in 
Arceuthobium. Unfortunately, there has been no opportunity to observe 


100 MADRONO [ Vol. 16 


the development of these abnormal fruits. Possibly the double fruits 
arise as fasciations and each stigma is pollinated separately. The resulting 
seeds seem to develop more or less independently of each other. The two 
seeds are enclosed in separate endocarps. 

Usually both seeds develop at about the same rate (type 1, fig. 1B), 
but sometimes one is suppressed (type 2, figs. 1C and 1D). I am unable 
to explain satisfactorily the development of the fruit bearing a single 
stigma but containing two normal seeds (type 3, fig. 1E) ; however, both 
embryo sacs may have developed as each seed became enclosed in a 
separate endocarp. The most unusual fruit was that containing three 
stigmas and two seeds, one with two embryos (type 4, fig. 1F). The 
diembryonic seed had embryos at opposite ends. Both embryos were about 
normal size, but the accessory one was somewhat irregular in shape. 


ABNORMAL SEEDS 


Peirce (1905) described a seed of A. campyvlopodum f. campylopodum 
(A. occidentale) that had two embryos (one about normal size and the 
other one third normal size) but within a single endosperm. Weir (1914) 
reported diembryonic seeds in A. vaginatum {. cryptopodum (3 of 20 
seeds), A. douglasi (4 of 30 seeds), and A. americanum, and although he 
did not describe them in detail, he stated that they were morphologically 
similar to normal seeds but “occasionally below average size.’’ Heinricher 
(1915, Plate 1, fig. 6) illustrated an unusual diembryonic seed in the 
European A. oxycedri; the seed itself appears to be similar to normal 
seeds, but it has two hypocotyls. 

Diembryonic seeds have been found by the writer in both A. america- 
num and A. vaginatum {. cryptopodum. They differ from the diembryonic 
seeds previously described in the literature (see above) in that they also 
contain two endosperms (fig. 3). Apparently both embryo sacs develop 
so that there are two units each of embryo and endosperm, both enclosed 
within a common endocarp. (These differ from the seeds shown in fig. 1E 
which are enclosed in separate endocarps.) The two units differ in size, 
the embryo in the larger unit being about normal size. The embryo in the 
second unit is smaller, the reduction being approximately proportional to 
that of the endosperm. No seeds of this type were found in the abnormal 
fruits dissected, therefore it is assumed that they are formed in normal 
appearing fruits. 

Counts of Arceuthobium vaginatum f. cryptopodum seeds in various 
localities showed that 1.0 per cent were of this abnormal type (Table 1). 

No counts have been made on the frequency of abnormal seeds in A. 
americanum, but they appear to be about as rare as in A. vaginatum f. 
cryptopodum. It has not been determined whether or not these abnormal 
seeds will produce two hypocotyls. However, Heinricher (1915) and Weir 
(1914) observed formations of double hypocotyls in the species of 
Arceuthobium which they studied. 


1961] HAWKSWORTH: ARCEUTHOBIUM 101 


TABLE 1. ABNORMAL SEEDS OF ARCEUTHOBIUM VAGINATUM 
F. CRYPTOPODUM FROM VARIOUS LOCALITIES. 


SEEDS EXAMINED ABNORMAL 
LOCALITY NUMBER PERCENT 
Sandia Mountains, New Mexico 500 2.4 
Manzano Mountains, New Mexico 925 ye 
Flagstaff, Arizona 3,950 0.8 
Roosevelt National Forest, Colorado fiz 0.7 
TOTALS 6,147 1.0 


DISCUSSION 


The formation of two seeds in the fruit of Avceuthobium has the ad- 
vantage of increased reproductive capacity. However, this is presumably 
accompanied by decreased efficiency of the seed dispersal mechanism. 

Polyembryony is common in the Loranthaceae. Its possible significance 
in the dioecious mistletoes is discussed by Allard (1943). He suggests 
that male and female plants may arise from different embryos within a 
seed. If this is true, it is possible that a mistletoe population could de- 
velop in a new area from a single seed. 


Rocky Mountain Forest and Range Experiment Station, 
Forest Service, U.S. Department of Agriculture, 
with headquarters at Colorado State University, 

Fort Collins, Colorado 


LITERATURE CITED 


ALLARD, H. A. 1943. The eastern false mistletoe (Phoradendron flavescens) ; when 
does it flower? Castanea 8:72-78. 

Dowp1nc, E. SILvER. 1931. Floral morphology of Arceuthobium americanum. Bot. 
Gaz. 91:42-54. 

Griz, L. S. 1935. Arceuthobium in the United States. Trans. Conn. Acad. Arts and 
Sci. 32:111-245. 

HEINRICHER, E. 1915. Die Keimung und Entwicklungsgeschichte der Wacholdermistel, 
Arceuthobium oxycedri, auf Grund durchgeftihrter Kulturen geschildert. Akad. 
Wien. Sitz-ber. 124:319-352. 

Kurt, Jos. 1955. Dwarf mistletoes. Bot. Rev. 21:569-626. 

. 1960. Morphological aspects of parasitism in the dwarf mistletoes (Arceu- 
thobium). Univ. Calif. Publ. Bot. 30(5) :337-436. 

PEIRCE, GEORGE J. 1905. The dissemination and germination of Arceuthobium occi- 
dentale, Eng. Ann. Bot. 19:99-113. 

THopay, D. and JoHNson, EMMA Trevor. 1930. On Arceuthobium pusillum, Peck. 
II. Flowers and Fruit. Ann. Bot. 44:813-824. 

WEIR, J. R. 1914. The polyembryony of Razoumofskya species. Phytopathology 
4:385-386. 


102 MADRONO [ Vol. 16 


TO ALBERT W. €. T. HERRE 


Dr. Herre, the California Botanical Society congratulates you on your 
forthcoming ninety-third birthday, September 16, 1961. This close ap- 
proach to the century mark in itself excites admiration among us, for few 
indeed possess the heritage that makes such an accomplishment possible. 
But attainment of this outstanding age is but one of the attributes that 
places you high in our esteem, for your accomplishments as a naturalist 
in the fields of ichthyology, lichenology, and ecology through three quar- 
ters of a century set you apart as a scientist extraordinary. 

As we realize that lichenology has claimed only a segment of your 
thought, study, and publishing activities, and that your list of papers 
dealing with taxonomic ichthyology, ecology, and geographical distribu- 
tion of fishes includes several hundred titles, we marvel that one man has 
been able to accomplish so much in his life-time, long and busy though 
it has been. Nor, we realize, has writing scientific papers, monographs, 
and textbooks claimed all of your time and energy. Individuals well be- 
yond middle age can recall that their early grade and high school training 
occurred under your supervision while you served as teacher, principal, 
and superintendent of schools. Others are equally aware of your admin- 
istrative and research activities in the Philippines and your promotion of 
careful work on fishes and other natural resources in that part of the world 
before you returned to the United States in 1928. 

We remember, also, that you accepted a challenging appointment with 
the Fish and Wildlife Service after you became Curator Emeritus of the 
Ichthyological Collections at Stanford University in 1947, and spent a 
strenuous year in your old area of field operations, the Philippine Islands, 
carrying on extensive collecting activities. 

Adding further to your laurels, you followed the Philippine work with 
a dozen years as Ichthyologist and Curator of Tropical Fishes at the Uni- 
versity of Washington, carrying forward work on large collections of 
lichens during your ‘‘spare” time. Then, when Mrs. Herre’s health was 
jeopardized by the cool, damp climate along the Washington coast, and 
you moved to Santa Cruz, some thought you would be content to reduce 
your work load and withdraw from active participation in biological re- 
search. Others, who knew better your penchant for continuous work, were 
not greatly surprised that instead, you launched into the final stages of 
preparing a monograph on the genus Usnea as represented in North Amer- 
ica. Few, indeed, have the strength, the desire, and the will to take up 
such an arduous task when nearly ninety years of age. Still fewer success- 
fully apply to the National Science Foundation for a grant to enable 
them to visit over a score of herbaria and private collections to carry the 
undertaking to completion! 

In celebration of your ninety-third birthday, we congratulate you on 
your scientific accomplishments, admire your physical stamina and 
mental alertness, wish we could consistently display your cheerfulness, 


1961] WIGGINS: HERRE 103 


and hope that one of us may have the privilege of preparing a congratu- 
latory message to you seven years hence. We sincerely hope that you will 
enjoy yet more years of extraordinary good health and continue your 
interest in the various phases of natural history that have fascinated you, 
and to which you have contributed so much during a large portion of a 
century.—Ira L. Wiccins, Stanford University. 


104 MADRONO [Vol. 16 


CHROMOSOME COUNTS IN THE GENUS MIMULUS 
(SCROPHULARIACAE) 


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


Although our long range investigation concerns the evolution of species 
in sections Simiolus and Erythranthe of the genus Mimulus (Vickery, 
1951), we have recently made genetical and cytological studies of several 
species belonging to other sections of the genus. The crossing results 
have already been given (Vickery, 1956), and this paper presents the 
cytological findings. 


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Fic. 1. Meiotic chromosomes of Mimulus ringens (5074), M. aurantiacus (6085), 
M. moschatus (6086) and M. floribundus (6110). All configurations are in or near 
second metaphase. Camera lucida drawings were made at 2,520 and reduced to 
1,260 in reproduction. 


The same method of bud fixation was employed as in previous investi- 
gations (Mukherjee and Vickery, 1959, 1960). Each chromosome num- 
ber determination was based on counts from an average of nine pollen 
mother cells. Herbarium specimens of each culture will be deposited in 
the Garret Herbarium of the University of Utah (UT). 

Four species, representing three sections of the genus, were studied: 
Mimulus ringens L. of section Eumimulus; M. aurantiacus Curt. of 
section Diplacus; and M. moschatus Dougl. and M. floribundus Dougl., 
both of section Paradanthus (see table 1 and figure 1). 

The count of n= 12 for blue flowered M. ringens of eastern North 
America, type species of the genus, differs from any previously reported 
for the genus (Campbell, 1950; Carlquist, 1953; Darlington and Wylie, 
1955; McMinn, 1951; Mukherjee and Vickery, 1959, 1960; Mukherjee, 
Wiens, and Vickery, 1957a, 1957b; and Vickery, 1955). However, 
M. ringens is the only species so far counted in section Eumimulus, and 
additional counts of n = 12 are possible in this section. 

Shrubby WM. aurantiacus of the chaparral areas of central and north- 
ern California was found to have n= 10 chromosomes, as do other 


1961] MUKHERJEE & VICKERY: MIMULUS 105 
TABLE 1. CHROMOSOME COUNTS IN THE GENUS MIMULUS. 


n—12 M.ringens L., section Eumimulus. 
St.-Jean, St.-Jean County, Province of Quebec, altitude ca. 200 feet, 
M. Raymond and J. Kucyniak, summer 1951 (5074). 

n=10 M. aurantiacus Curt., section Diplacus. 
Near Round Top, Alameda County, California, altitude 1,200 feet, 
Vickery 990 (6085). 

n=16 WM. moschatus Dougl., section Paradanthus. 
Mill Creek Canyon, Salt Lake County, Utah, altitude 7,400 feet, 
Vickery 1036 (6086). 

n=—16 MM. floribundus Dougl., section Paradanthus. 
Hog Ranch, Mather, Tuolumne County, California, altitude 4,600 feet, 
Vickery 1372 (6110). 


species of section Diplacus previously reported by McMinn (1951). 
McMinn’s extensive crossing studies suggest that this number, not found 
in other sections of the genus, is characteristic of section Diplacus. 

Mimulus moschatus and M. floribundus, both widespread in western 
North America, were found to have n = 16 chromosomes. Both species 
have small yellow flowers and are low-growing with viscid-pubescent 
leaves and stems, but they differ markedly in leaf size and shape and 
in duration (M. floribundus is annual; M. moschatus is perennial). The 
two species hybridized readily in the garden. The Fi hybrids were vig- 
orous, but completely sterile (Vickery, 1956). Despite the cytological 
and some morphological similarities, these two entities would appear 
to be genetically and taxonomically distinct. 

Department of Genetics and Cytology 


University of Utah 
Salt Lake City 12, Utah 


LITERATURE CITED 


CAMPBELL, G. R. 1950. Mimulus guttatus and related species. El Aliso 2:319-335. 

CarLQuIsT, S. 1953. in Documented chromosome numbers of plants. Madrono 12:31. 

Dariincton, C. D. anp A. P. Wyite. 1955. Chromosome atlas of flowering plants. 
Allen and Unwin Ltd. London. 519 pp. 

McMinn, H. E. 1951. Studies in the genus Diplacus (Scrophulariaceae). Madrono 
(335123. 

MUKHERJEE, B. B. anv 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. Madronfto 15:239-245. 

MuxKHeERjJEE, B. B., D. Wiens, AND R. K. Vickery, Jr. 1957a. Chromosome counts 
in the section Simiolus of the genus Mimulus (Scrophulariaceae). II. Madrono 
14:128-131. 

. 1957b. Chromosome counts in the section Erythranthe of the genus Mimu- 
lus (Scrophulariaceae). Madrofio 14:150-153. 

VickeErY, R. K., Jr. 1951. Genetic differences between races and species of Mimulus. 
Carn. Inst. Wash. Year Book 50:118-119. 

. 1955. Chromosome counts in section Simiolus of the genus Mimulus (Scro- 
phulariaceae). Madrofio 13:107—110. 

. 1956. Data on intersectional hybridizations in the genus Mimulus (Scro- 
phulariaceae). Proc. Utah Acad. 32:65—71. 


106 MADRONO [Vol. 16 


SPHENOPHYLLUM NYMANENSIS SP. NOV. FROM THE 
UPPER PENNSYLVANIAN? 


J. F. DAviIpsoN 


While coal is not found in commercial quantities in Nebraska, there 
are a number of thin seams exposed in the southeastern counties of the 
state, and investigation of these has proved to be quite profitable from a 
paleo-botanical viewpoint. In the majority of sites investigated, a layer 
of limestone immediately above the coal has precluded the finding of 
anything but highly coalified fossils. However, in the clay pit of the 
Western Brick Company at Nebraska City in Otoe County, the exposed 
Nyman coal shales out, and plant fossils are abundant. Plant material is 
so abundant in fact that the specimens, frequently with cuticle intact, are 
almost impossible to separate. It was from this site that representatives 
of the Pennsylvanian Sphenopsida of the order Sphenophyllales, includ- 
ing the present Sphenophyilum, were collected. 

According to Condra and Reed (1943), the Nyman coal is found to- 
ward the top of the Langdon shale formation of the Richardson Sub- 
group, Wabaunsee Group, Virgil Series of the Pennsylvanian Sub-system. 
In part of the clay pit, as in the other sites mentioned above, the Nyman 
coal lies immediately below the Dover limestone. As the coal begins to 
shale out, the plant remains are separated by such minute quantities of 
shale as to be almost impossible to recover. However, as shaling continues, 
and the amount of shale increases, the plant remains are more readily 
defined. Preservation is very good in the fine sediment, and, as reported 
by Barbour (1914), many compressions retain their cuticular coverings 
which may be floated free, cleared, and mounted for study. 

In the deposit, specimens of Sphenophyllum are fairly common, al- 
though few show more than two or three nodes. Sphenophvllum cunei- 
folium Sternb., S. emarginatum Brong. and S. majus Brong. are repre- 
sented, as well as another large-leaved taxon which was at first consid- 
ered to be a variant of S. majus. Closer examination, and a comparison 
of a number of these large-leaved specimens with specimens of S. majus 
indicate that two taxa are involved, and the name Sphenophyllum nyma- 
nensis is hereby proposed for the novelty. 

Sphenophyllum nymanensis sp. nov. (Fig. 1) Leaves in whorls of 6 
per node, 12-17 mm. long, 5-10 mm. wide; veins branching 3—5 times 
from the base, terminating at the rounded to somewhat truncate apex; 
stems fairly robust for a Sphenophyllum, about 2 mm. in diameter, with 
nodes swollen to 3 mm.; internodes subequalling the leaf length, 12—17 
mm. long. 

Locatity. Clay pit, Western Brick Company, Nebraska City, Otoe 
County, Nebraska. 


1 This work was supported by a grant from the University of Nebraska Research 
Council. 


1961] DAVIDSON: SPHENOPHYLLUM 107 


Horizon. Nyman coal, and Langdon shale above, Virgil Series, Upper 


Pennsylvanian. 
Type. Paleobotanical collection, University of Nebraska State Museum. 


METRIC ] 


MO CT 


Fic. 1. Sphenophyllum nymanensis J. F. Davidson. 


Sphenopyllum nymanensis, in terms of size most closely resembles 
S. majus from which it is readily distinguished by the following char- 
acters: 


S.nymanensis S. majus 
6 leaves per node 8—10 leaves per node 
leaf apex rounded to slightly leaf apex truncate 
truncate 
leaf margin entire leaf margin with each vein term- 


inating in a small deltoid tooth. 


Department of Botany, 
University of Nebraska, 
Lincoln, Nebraska. 


LITERATURE CITED 


Barsour, E. H. 1914. Plant tissue in the Carboniferous shales of Nebraska. Nebr. 
Geol. Surv. 4, part 16, pp. 231-232. 

Conpra, G. E. and E. C. REEp. 1943. The geological section of Nebraska. Nebr. Geol. 
Surv. Bull. 14. 


108 MADRONO [Vol. 16 
A NEW NAME IN THE ALGAL GENUS PHORMIDIUM 
FRANCIS DROUET 


Phormidium anabaenoides, nom. nov. P. thermale Drouet. Publ. 
Field Mus. Bot. 20(6):138. 1942. A new name is necessary for this alga 
of hot springs of Lake and Sonoma counties, California, because of the 
discovery in the literature of another P. thermale described by Professor 
V. Vouk (Prirod. Istr. Hrvatske i Slavon., Jugosl. Akad., Mat.-Prirod. 
Razr. 8:9. 1916). The research involved here was supported by the Na- 
tional Science Foundation. 


Department of Botany, College of Agriculture 
University of Arizona, Tucson 


NOTES AND NEWS 


PLAGIOBOTHRYS AUSTINAE (GREENE) JOHNSTON: A NEW ADDITION TO THE OREGON 
Friora.—In April, 1959, the distinctive Plagiobothrys austinae (Greene) Johnston, 
formerly believed endemic to the Great Valley of California with a range of distribu- 
tion from Stanislaus to Shasta counties, was collected in the botanically interesting 
Agate Desert west of Camp White, Jackson County, Oregon (Ornduff 5043A, UC, 
OSC, WTU). The locality in Oregon where this species occurs is separated from its 
nearest station in California near Redding, Shasta County, by about 140 miles of 
the Klamath-Cascade mountain complex. In many aspects of vegetation and topog- 
raphy, the Agate Desert is strongly reminiscent of parts of the northern Sacramento 
Valley in California; consequently, intensive collecting in the future may be expected 
to reveal additional Californian floral elements in the Agate Desert —FRANCIA CHISAKI 
and ROBERT ORNDUFF, Department of Botany, University of California, Berkeley. 


STEGNOSPERMA CUBENSE AND GOSSYPIUM KLOTZSCHIANUM DAVIDSONII NOT KNOWN 
IN THE REVILLAGIGEDOS.—On the expedition of the California Academy of Sciences 
to the Revillagigedo Islands in 1925, plant collections were made not only there but 
also en route (Proc. Calif. Acad. ser. 4, 18:393-484, 1929). Labels of way specimens, 
headed “Expedition to the Revillagigedo Islands,” have led evidently to one and ap- 
parently to two erroneous reports. 

Rogers (Ann. Missouri Bot. Gard. 36:476, 1949) reported Stegnosperma cubense 
A. Richard from the Revillagigedos on the basis of Mason 1846; but Mason’s field- 
book shows that this collection is from Isabel Island, just off the Mexican mainland. 

Hutchinson (in Hutchinson, Silow, and Stephens, The evolution of Gossypium 
and the differentiation of the cultivated cottons, 1947, p.23) reported Gossypium 
klotzschianum var. davidsonii (Kellogg) Hutchinson from the Revillagigedos, though 
without citing a specimen. This report has been repeated elsewhere. Upon inquiry, 
Dr. Hutchinson wrote that the report appeared to be erroneous, based on a specimen 
from an expedition to the Revillagigedos but collected in Baja California. Very likely 
he was misled by the same label heading (Mason 1936, 1937 from Magdalena Bay). 

Since it does not seem feasible at present to square the facts with the reports by 
introducing these two plants into the Revillagigedos, perhaps the best expedient is 
this note——RrE1p Moran, Natural History Museum, San Diego, California. 


INFORMATION FOR CONTRIBUTORS 


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Shorter items, such as range extensions and other biological notes, 
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Institutional abbreviations in specimen citations should follow Lanjouw 
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Utrecht. Second Edition, 1954). 

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

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MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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i 


MADRONO 


VOLUME 16, NUMBER 4 OCTOBER, 1961 


Contents 


PAGE 


EVOLUTION OF THE GALIUM MULTIFLORUM COMPLEX 
IN WESTERN NorTH AMERICA. I. DIPLOIDS AND 
POLYPLOIDS IN THIS DIoEcIoUS GROUP, 
Friedrich Ehrendorfer 109 


A NEw SPECIES oF LyctuM IN NEVADA, 
Cornelius H. Muller 122 


SOME RECENT OBSERVATIONS ON PONDEROSA, JEFFREY 
AND WASHOE PINES IN NORTHEASTERN CALIFORNIA, 
John R. Haller 126 


INFLUENCE OF TEMPERATURE AND OTHER FACTORS ON 
CEANOTHUS MEGACARPUS SEED GERMINATION, 
Elmer Burton Hadley 7 132 


REviEws: John Hunter Thomas, Flora of the Santa 
Cruz Mountains of California. A Manual of the Vas- 
cular Plants (Mary L. Bowerman); R. W. Allard, 
Principles of Plant Breeding (Thomas W. Whitaker) 138 


Notes AND NEws 140 


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 
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 
Mitprep E. Matutas, 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—JoHN H. THOMAS 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Baki Kasapligil, Deparment of Biology, Mills College, California. 
First Vice-president: Lawrence R. Heckard, Department of Botany, University of 
California, Berkeley, California. Second Vice-president: Kenton L. Chambers, De- 
partment of Botany, Oregon State College, Corvallis, Oregon. Recording Secretary, 
Mary L. Bowerman, Department of Botany, University of California, Berkeley, Cali- 
fornia. Corresponding Secretary: Wallace R. Ernst (January-June), Lauramay 
Dempster (July-December), Department of Botany, University of California, Berke- 
ley, California. Treasurer: John H. Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1961) EHRENDORFER: GALIUM 109 


EVOLUTION OF THE GALIUM MULTIFLORUM COMPLEX 
IN WESTERN NORTH AMERICA 


I. DIPLOIDS AND POLYPLOIDS IN THIS DIOECIOUS GROUP 


FRIEDRICH EHRENDORFER 


The Galium multiflorum complex comprises a group of races which are 
spottily distributed through the interior of western North America. They 
occupy, for the most part, dry and rocky talus slopes and cliffs, and they 
range from the Larrea desert zone to alpine timberline. Life forms change 
correspondingly from low xerophytic desert shrubs to reduced alpine 
hemicryptophytes. All members of this racial complex are dioecious and 
sexually reproducing. Their mostly rotate, yellowish or greenish flowers 
are insect pollinated. Long silky hairs on the fruits facilitate wind dis- 
persal. 

The Galium multiflorum complex belongs to the section Lopho-Galium 
K. Schum. of this rubiaceous genus. Circumscription, differentiation, and 
first detailed taxonomic treatment along classical lines have been worked 
out by Ehrendorfer (1956). Further contributions, including a somewhat 
changed taxonomical evaluation, have been published by Dempster 
(1959). 

Within the frame of a broad scientific project aiming at an understand- 
ing of the main evolutionary features of this world-wide genus along com- 
parative lines (for publications available so far, see Ehrendorfer 1958), 
work on the Galium multiflorum complex has been continued and ex- 
panded since 1959. This has been made possible by financial support from 
a Rockefeller grant and especially from National Science Foundation 
grant number 9825. Under the guidance and extremely stimulating co- 
operation of Professor G. L. Stebbins, the author carried out intensive 
research work in 1959; this included study of habitats and accompanying 
vegetation for nearly all of the chief divisions of the G. multiflorum com- 
plex, collection of transplant material and extensive population samples 
for statistical analyses, cytological research, and comparison of practi- 
cally all herbarium material available. Later this project was joined and 
continued by Lauramay T. Dempster, whose help in various respects is 
gratefully acknowledged. 

The present research on the Galium multiflorum complex aims at a 
reconstruction of its evolution in time and space as part of the general 
unfolding of the Great Basin flora and vegetation since the Tertiary and 
at the demonstration of the main evolutionary mechanisms involved. Re- 
sults will be published in a series under the general heading of which this 
is the first contribution. The main questions dealt with here are: (a) basic 
evolutionary mechanisms as elucidated by cytological behaviour; and 
(b) distribution of diploids and polyploids within the complex. 


Maprono, Vol. 16, No. 4, pp. 109-140. October 30, 1961. 


110 MADRONO [ Vol. 16 


MATERIALS AND METHODS 


Populations of various races of the Galium multiflorum complex studied 
in the field were designated by collection numbers. The italicized numbers 
in the text refer to the field collection numbers of the author and his 
various co-workers. Further data on these collections will be found in the 
detailed list of collections below. Transplant material was obtained and 
grown in the greenhouse of the Genetics Department, University of Cali- 
fornia, Davis, and later also in Orinda, California. Cultivation over long 
periods seems difficult, however, as requirements are quite specialized as 
to soil, temperature, etc. Carnoy solution (5 parts of 95 per cent alcohol: 
3 parts of chloroform: 1 part of concentrated acetic acid) was used for 
the fixation of young flower buds in the field and of fresh shoot apices 
from transplanted material. Fixations were stored in the refrigerator. 
Saturated aceto-carmine with iron was employed for staining: anther 
material was squashed after short and gentle heating in the stain, whereas 
vegetative tissues were boiled in aceto-carmine for two minutes. A Zeiss 
standard Series GFL microscope was used for cytological work, and the 
drawings were made with a camera lucida. Herbarium vouchers from 
plants with known chromosome number will be deposited in principal 
herbaria after conclusion of the present research project. 


GENERAL CYTOLOGY 


Mitosis, meiosis and pollen grain mitosis were studied in various races 
of the Galium multiflorum complex. As a main result the existence can be 
established of a polyploid series with diploids, tetraploids and a local 
hexaploid (see discussion), all having the base number x11. Apart from 
determination of chromosome numbers some general cytological observa- 
tions have been made. 

The dioecious sex differentiation in the group led to the question as to 
whether sex chromosomes were cytologically recognizable. Mitoses of 
shoot apices from male and female plants of the diploid G. halla were 
carefully compared. No obvious difference in number or shape of chromo- 
somes could be detected. One has to consider, though, that with relatively 
small and not very clearly differentiated chromosomes (fig. la), minor 
differences could easily escape notice. In various diploid and polyploid 
races meioses in pollen mother cells were scrutinized in order to find 
heteromorphic bivalents of sex chromosomes, but without success. Cyto- 
logical differentiation of males and females in the G. multiflorum com- 
plex, therefore, seems to be absent or cryptic. 

Comparisons were made of mitoses from shoot apices of very distantly 
related diploids, like G. hallii and G. hypotrichium subsp. tomentellum, 
in order to find out whether structural differentiation of chromosome sets 
had occurred (fig. la, b). Results show that there is much similarity. Ab- 
solute size differences apparent from the figures la and 1b may be due 
to the developmental stage of the individual cells. In both races the chrom- 
osome set consists of: A) one pair of SAT-chromosomes,! B) one pair 


1961 | EHRENDORFER: GALIUM 111 


O AO 20 


Jaa 


Fic. 1. Somatic mitosis in Galium. a, G. hallit (5901), b, G. hypotrichium subsp. 
tomentellum (5941), c, G. grayanum subsp. glabrescens (5963, tp. 2). 


of long and strongly heterobrachial chromosomes, C) four pairs of chrom- 
osomes with less conspicuously different arms, and D) five pairs of 
chromosomes with median or submedian centromere. Groups C and D 
are not always clearly separable. This general pattern is also maintained 
in the allotetraploid G. grayanum subsp. glabrescens (fig. 1c). In spite of 
these similarities, indications for some structural differentiation can be 
found, e.g., in the SAT-chromosomes, where relation between long and 
short arms as well as satellite size are clearly different in G. hall and 
G. hypotrichium subsp. tomentellum. 

In spite of some cell-to-cell variability in chromosome size, there is a 
clear hereditary diminution of chromosome size with polyploidy. This is 
demonstrated better by comparison of somatic mitoses (fig. 1) from dip- 
loids than by comparison of corresponding stages of pollen meioses (fig. 
2a, b) or pollen mitoses (fig. 2 1,m). Diminution of chromosome size is 
less obvious in the probably quite recent local hexaploid (fig. 2c). This 
reduction of chromosome size (and very likely nuclear size) in poly- 
ploids does not fully compensate the cell size increase correlated with 


1 The satellites often stick to the short arms, making it difficult to recognize the 
SAT-chromosomes. Such sticking seems to happen nearly always in pollen mitoses 
fig. 2 1,m). 


112 MADRONO [Vol. 16 


polyploidy, as is apparent from comparison of PMC’s and young pollen 
grains which are larger in polyploids than in diploids (fig. 2a-c, 1, m). 
Studies to determine the effect of polyploid level upon stomatal size are 
being carried on by Mrs. Dempster, with promising results for the use of 
stomatal measurement as an indication of ploidy when cytological evi- 
dence is lacking. 

In polyploids of the Galium multiflorum complex there is occasional 
intra-individual somatic instability of chromosome number. This has been 
established for G. gravanum subsp. glabrescens (5963, tp. 1) and it prob- 
ably occurs in G. munzit (5980, tp.c.). From excellent fixations resulting 
in exceptionally clear mitotic pictures, it is evident that the former plant 
has 2n=—45 as the most common number in young buds and shoot apices, 
but that occasional deviations occur, of which —44 —43 —42 -41 asa 
reductional series and —47 as an indication for accumulation were actu- 
ally observed. One plant (5980, tp.c.) seems to vary between 2n=42 
—44—-45. In a number of other polyploids, e.g., G. grayanum subsp. gla- 
brescens (5963, tp.2) and G. munzii var. kingstonense (5969), counts of 
numerous mitoses always gave the stable number 2n—44. Intra-individual 
instability of chromosome number may be due to unbalanced primary 
number and/or spindle defects. Unstable somatic chromosome numbers 
have not been found in diploids. 

Normal sexual reproduction is highly probable within the group. The 
course of pollen meiosis is remarkably normal in diploids and polyploids; 
male and female plants always coexist in the populations, often with 
an excess of males, and there is obvious morphological variability within 
the populations. 

Chiasma frequency is variable. In the diploids with their relatively 
large chromosomes, frequency of bivalents with two chiasmata is often 
higher than 50 per cent, the other bivalents having only one chiasma 
(fig. 2d, e). Unpaired chromosomes seem to be extremely rare in the 
diploids. In the tetraploids with reduced chromosome size, the number 
of chiasmata is decidedly lower. In G. hypotrichium from Alpine County, 
for instance, there are only 15 per cent bivalents with two chiasmata in 
a plant from the Ebbetts Pass series (5920) and 10.9 per cent in a plant 
from the Sonora Pass series (5917) [each calculated from 50 PMC’s]. 
Multivalents (trivalents with univalents and quadrivalents in chains and 
rings, fig. 2f) are relatively rare. In the above-mentioned plants there 
were 8 per cent PMC’s with ITI-+I and 28 per cent with IV (5920) and 
8 per cent PMC’s with III-++I and 22 per cent with IV (5917). Uni- 
valents lag in anaphase I (fig. 2h), and are prematurely divided, but 
their halves are unable to divide a second time in anaphase II (fig. 21) ; 
so they are eliminated or finally fuse with some of the telophase nuclei. 
As a result, pollen grains of polyploids occasionally contain deviating 
chromosome numbers (fig. 2m). This phenomenon must be basically re- 
sponsible for some polymorphism of 2n numbers in populations of poly- 
ploids. 


1961] EHRENDORFER: GALIUM 113 


ag = 


i k m 


oO 10 20 30 


Fic. 2. Division figures in Galium. a-c, PMC metaphase II in diploid, tetraploid 
and hexaploid plants: a, G. serpenticum—2x (5911-5913, fd. Ila); b, G. munzii-4x 
(5903, tp.); c, Galium (6x race), Warner Mountains (5909, 5910). d-i, PMC divi- 
sions: d, diakinesis, G. grayanum—2x (5933-5936) ; e, metaphase I, side view, G. ser- 
penticum—2x (5914); f, multivalents, III+I, IV chain, IV ring, G. hypotrichium 
subsp. hypotrichium—4x (5920); g, anaphase I with bridge and attached fragment, 
G. parishti-2x (5902) ; h, telophase I with lagging univalent, G. hypotrichium subsp. 
hy potrichium—4x (5917) ; i, telophase II with lagging univalent halves, G. hypotri- 
chium subsp. hypotrichium-4x (5920). k-m, first pollen mitosis: k, anaphase with 
free acentric fragment, G. hypotrichium subsp. hypotrichium-2x (5916); 1, meta- 
phase G. hypotrichium subsp. hypotrichium-2x (5916); m, metaphase, aberrant 
pollen grain with n=23, G. munzii * G. hypotrichium subsp. subalpinum—4x (5947). 


114 MADRONO [Vol. 16 


In 3 diploids, 1 tetraploid and 1 hexaploid, consequences of chromo- 
some aberrations have been seen: bridges with and without fragments in 


\. «© coloradoense (2x) 
& i 
one re pines serpenticum (2x) 
5 Nae Si 6) var. puberulum (2x) 


x3 0 grayanum (2x) 
©) @ subsp glabrescens (4x) 


hypotrichium 
subsp. hypotrichium (2x) 
?)subsp.tomentellum (2x) 
8)subsp. utahense (2x) 
9) subsp. scabriusculum(2x) 


10) subsp. hypotrichium 4) 
11) subsp. subalpinum (4x) 


i] 
4 
1 
Ly 
L 
uy 
y 
~ at—t-le 7, 
fie tn ‘“  |12)subsp. scabriusculum(4x 
\ N 
4 is 
ry \ 
! = 
J Aye  6x-race, Warner MTs. 
7 - I-{ 
7 x 
/ 
aa ‘ 
Yv a 
4 
z 5 
ay = 
\ T 
~ T 
v 
SiSiola jatar eveiaieceeiieds ee “Wn, 
Se Be ot le b ok cuca. > 4, 
xy d O 8 = aya “My, 
Ay x : i ty 
x oF x 3 % 
x~Q * + : % 
x” *& £ : 4 
eee Re 3 3 
x pi = = 
~Q 2 + O E E 
4 gy +. 9 2 2 
Bad 2 E 
5 ween 1 Oy % % 
~ aes ACN Eats “ny, = 
“Ne “ RAKXX 5 a i, 
GAPE yx KK KEEE ~ “, 
Nese Ny 


Oe 


x*7 \ N 


ai VT stellatum 
subsp. eremicum (2x) . 
£2: multiflorum (2x) ° 


<> 0 matthewsii (2x) S 
£3 magnifolium (2x) 
munzii 


subsp.munzii (4x) 
1) var. kingstonense (4x) 

Od 3)approaching matthewsii (4x) 
4)approaching magnifolium 4x) 
5) hybrids with hypotrichium 


subsp. subalpinum (4x) 
pets snaen eat ue x Cedros Island: stellatum subsp. stellatum 


Fic. 3. Distribution of the Galium multiflorum complex, shown by outlines. Loca- 
tions of populations examined cytologically are indicated by individual symbols. 


1961] EHRENDORFER: GALIUM Ns 


anaphase I (fig. 2g), and liberation of attached fragments in anaphase of 
pollen mitosis (fig. 2k). In the hexaploid there are occasional spindle 
defects, restitution nuclei, and formation of resultant dyads. 

Male sterility as a result of break-down of pollen development after 
normal meiosis has been observed in a diploid G. serpenticum plant from 
Mount Bidwell (5977). 


DISTRIBUTION OF DIPLOIDS AND POLYPLOIDS 


Determination of chromosome numbers and ploidy levels is one of the 
major prerequisites for the reconstruction of the evolutionary history of 
the Galium multiflorum complex. So far, chromosome counts are avail- 
able for most of the major species, subspecies, and hybrid populations 
presently recognized. These counts originate from 51 populations and 
more than 100 individuals. Populations with known chromosome numbers 
are shown on the map (fig. 3), as single symbols within the outlines of 
the total distributional area of their respective taxa. 

The following list contains further information on geographical origin 
and habitat of the populations cytologically studied. The abbreviation 
“fq.” indicates fixations made in the field, mostly comprising several indi- 
viduals; chromosome counts in such instances usually refer to more than 
one individual. The abbreviation “tp.” indicates fixations from single 
transplant individuals further designated by numbers or letters. Definite 
chromosome numbers are given after a minimum of several unquestion- 
able counts per plant; if only one or few counts are available the symbol 
“==” is used. The abbreviation “‘ca.” refers to approximate counts. Chrom- 
osome numbers are indicated in the diploid state (2n). They have been 
determined from vegetative tissues in flowers and shoot apices. Where 
counts have been carried out on pollen meiosis or pollen mitosis, 2n num- 
bers have been extrapolated and an asterisk is added. 

As is evident from the list, diploid populations are always uniform in 
respect to chromosome number (2n=22), while certain fluctuations of 
2n numbers around straight x multiples have been observed in some 
polyploid populations: G. munzi from Wildrose Canyon (2n=—44, 46), 
Lone Pine Creek (2n=42, 44), Zion National Park (2n=44, 45) and 
Grand Canyon (2n=42, 44). Polyploids are obviously less sensitive to 
aneuploidy, addition or loss of chromosomes, than diploids. The origin of 
aberrant types must be due to meiotic irregularities and deviating chrom- 
osome numbers in the gametes, or to intra-individual somatic instability, 
with similar consequences, situations which have already been referred to. 

In the following list of collections, only previously published names 
have been used, and the taxonomic scheme followed is not necessarily in 
entire agreement with the author’s present views. The taxonomic position 
of the p’ants in some of the populations represented is still uncertain, and 
their pl::cement in the list is conservatively based on published work of 
the past 


116 MADRONO [Vol. 16 


GALIUM COLORADOENSE Wight 

Utah, Dinosaur National Monument, 1 mile west of campground near Split Moun- 
tain, 5150 feet; Shinarump Quartz conglomerate; sandy and gravelly north slope, 
very open Pinyon-Juniper: Ehrendorfer & Stutz 5950. tp. 2n==22 


Utah, Uintah County, just south of Brush Creek on Highway 44, ca. 11 miles north 
northeast of Vernal, ca. 6000 feet; open bushy and herbaceous pioneer growth on 
steep Jurassic sandstone cliff: Ehrendorfer & Stutz 5951. tp.0' Zn 22 


GALIUM GRAYANUM Ehrendf. 

subsp. GRAYANUM 
California, Tehama County, North Coast Ranges, South Yollo Bolly Mountain, 
7700-8080 feet; gentle to steep talus slopes, metamorphic schists; subalpine pioneer 
vegetation among very open Abies magnifica: Ehrendorfer 5933, 5934, 5935, 5936. 


id.: 2n=—22* 
California, Lake-Colusa County, Goat Mountain, North Coast Ranges, 6000 feet: 
leg. G. L. Stebbins {d.: 2n=22 


California, Lassen National Park, Brokeoff Mountain, 9000 feet; gentle to steep talus 
slopes, volcanic andesite; subalpine pioneer vegetation: Ehrendorfer, Stebbins & 
Dempster 5964, 5965, 5966, 5967. tp.0 : <2n=22 

tp.1 : 2n=22 
California, Butte-Plumas County, Summit above Jonesville, ridge east of pass, 
6700-7000 feet; rocks and talus slopes, volcanic andesite; pioneer vegetation among 
subalpine chaparral, Abies magnifica and Pinus monticola: type locality, Ehren- 
dorfer 5937, 5938, 5939. fd.: 2n=22* 


California, Placer County, Sierra Nevada, Sugar Bowl resort southwest of Donner 
Summit, 7200 feet; steep northeast slope, andesite talus and rocky outcrops; pioneer 
vegetation among Quercus vaccinifolia, Abies magnifica and Pinus monticola: Ehren- 
dorfer & Stebbins 5905. fave 22 20 


subsp. GLABRESCENS Ehrendf. 
California, Siskiyou-Trinity County, west flank of Scott Mountain, ca. 6300 feet; 
ultrabasic intrusives; serpentine-peridotite; talus and gravel within very open Pinus 
jeffrevi and Abies concolor: Ehrendorfer & Stebbins 5915. fd.: 2n=44* 


California, Siskiyou County, 0.5 mile south-southeast of Castle Lake, southwest of 
Mount Shasta village, ca. 6000 feet; serpentine talus, pioneer vegetation among 
Arctostaphylos nevadensis, Pinus monticola, Abies magnifica: type locality, Ehren- 
dorfer, Stebbins & Dempster 5963. tp. 1 : 2n=41-42-43-44-45-47 
tp. 2 : 2n=44 (stable) 

GALIUM HALLII Munz & Jtn. 

California, Los Angeles County, northeast side of San Gabriel Mountains, Sawmill 
Canyon at west end of Swartout Valley, between Wrightwood and Lone Pine, 6600 
feet; on steep north slopes, in mineral soil (schist) among Quercus Kelloggii and 
Pinus ponderosa: Ehrendorfer 5901. fdvs.2n==22 


GALIUM HYPOTRICHIUM Gray 
subsp. HYPOTRICHIUM 

California, Alpine-Mono County, Sierra Nevada, ca. 0.5 mile south of Sonora Pass, 

ca. 10,200 feet; wind-exposed talus slope on crest line, volcanic rocks; very scat- 

tered pioneers: type locality, Ehrendorfer 5916. {de 2n=-22 7; 
fd)2n==22 

California, Alpine County, Sierra Nevada, ca. 2 miles west of Sonora Pass, ca. 9500 

feet; sheltered west slope, volcanic talus; Ribes bushes and herbs: Ehrendorfer 5917. 
fd.: 2n=44* 

California, Alpine County, Sierra Nevada, ca. 0.5 mile southeast of Ebbetts Pass, 

8950 feet; northwest slope, volcanic talus; Symphoricarpos, Artemisia, herbs: 

Ehrendorfer 5920. id:; 2n=44> 


1961 | EHRENDORFER: GALIUM 117 


subsp. SCABRIUSCULUM Ehrendf. (G. coloradoense var. scabriusculum Dempster) 
Utah, Carbon County, Castle Gate, side canyon on southwest of main valley, en- 
trance ca. 1 mile northwest of the town, ca. 6500 feet; gully with steep sandstone 
slopes; grassy and herbaceous cover among open Pinus ponderosa and Symphori- 
carpos: Ehrendorfer & Stutz 5954. (Oa eet 


Utah, Emery County, smali side canyons on either side of Buckhorn Wash, ca. 1 mile 
north of San Rafael River bridge, ca. 4500 feet; dry bottom of gullies with boulders 
and sand from sandstone; very open Pinyon-Juniper and scrub: near type locality, 
Ehrendorfer & Stutz 5952, 5953. tp. a: 2n=—44 
tp. b: 2n=+44 
. tp. c: 2n=ca. 44 
subsp. SUBALPINUM (Hilend & Howell) Ehrendf. 
California, Inyo County, Sierra Nevada, Cottonwood Lake Basin, slopes north and 
northwest above Muir Lake, ca. 11,200 feet; granitic talus; open Pinus balfouriana, 
Holodiscus and pioneer herbs: Ehrendorfer 5945, 5946. tp. 1: 2n==2644 
tp.2: 2n=+44 
subsp. TOMENTELLUM Ehrendf. 
California, Inyo County, Panamint Mountains, Telescope Peak, below top on north 
side, ca. 11,000 feet; talus of metamorphic schists; pioneers among very open Pinus 
aristata: type locality, Ehrendorfer 5941. fd: 2n==22 


subsp. UTAHENSE Ehrendf. 
Utah, Utah County, Wasatch Mountains, American Fork Canyon, steep north slopes 
along trail to Timpanogos Cave, 6400-6800 feet; limestone rocks and talus; bushes 
and herbaceous pioneers among scattered Pseudotsuga and Abies: type locality, 
Ehrendorfer & Stutz 5949. tp:: 2n=22 


Utah, Salt Lake County, Wasatch Mountains, steep north slopes, ca. 1 mile south- 
west of Alta, ca. 9000 feet; limestone rocks and crevices; vegetation similar to 5949; 
Ehrendorfer & Stutz 5955. tps 222 


GALIUM MAGNIFOLIUM (Dempster) Dempster 
Nevada, Clark County, Charleston Mountains, southwest of Las Vegas, near Cotton- 
wood Springs, canyon above Bar Nothing Ranch (= Wilson’s Ranch), ca. 4200 feet ; 
steep north slope along creek, alluvial material below mesozoic sandstone; open 
Pinyon with Artemisia, Yucca and Opuntia: type locality, Ehrendorfer & Dempster 
ey Ede CN tee 
ipy Zn" 22 
Utah, Washington County, ca. 1 mile northeast of Hurricane, steep slope above trib- 
utary of Virgin River, ca. 3000 feet; Jurassic sandstone and talus; with Ephedra, 
Artemisia, etc.; Ehrendorfer & Dempster 5979. fd:: 2n==22 


GALIUM MATTHEWSII Gray 

California, Inyo County, east side of Sierra Nevada, Big Pine Creek, slopes above 
the camp site and road head, 8800 feet; loose granitic talus and sand, among Arte- 
mista: Ehrendorfer 5922. fides 2-22" 


California, San Bernardino County, Kingston Mountains, steep slopes south of King- 
ston Pass, ca. 5500 feet; crevices of granite: Ehrendorfer & Dempster 5974. 
tDeal 222 
tp. bz 2n=22 
GALIUM MULTIFLORUM Kell. 
California, Modoc County, south of Eagleville, cliff above Lower Alkali Lake, 5000 
feet; in crevices and at the base of east-northeast-exposed basalt rocks: Ehrendorfer 
& Stebbins 5908. toe 2n==27 


California, Mono County, northwest shore of Mono Lake, on upper terrace, 6550 
feet; among Artemisia and Purshia on light pumice sand: Ehrendorfer 5918. 
tp.: 2n—=22 


118 MADRONO [Vol. 16 


Nevada, Storey County, Washoe Mountains, 2.2 miles north of Virginia City on 
Highway 17 to Reno, ca. 6600 feet; steep north slope, on volcanic talus among 
Artemisia, open Pinyon-Juniper along gully: type locality, Ehrendorfer & Stebbins 
5906. fdce2n=22 5 


forma HIRSUTUM (A. Gray) Ehrendf. 
California, Mono County, Sherwin Grade, dry wash southwest of Highway 6, 
ca. 6600 feet; volcanic rhyolite talus and rocks among Artemisia, open Pinyon: 
Ehrendorfer & Dempster 5921. tp.s 2Zn=—22 


GALIUM MuNzII Hilend & Howell 
var. KINGSTONENSE Dempster 
California, San Bernardino County, Kingston Mountains, steep slopes south of King- 
ston Pass, in gullies and toward the top, 5800-6000 feet; rocky ravines and steep 
slopes with Pinyon, granite: type locality, Ehrendorfer & Dempster 5969, 5971. 
tp.a: 2n=44 
tp.b: 2n=44 
subsp. MUNZII 
California, Inyo County, Lone Pine Creek just above its break-through into Owens 
Valley, ca. 5000 feet; steep north slopes with Pinyon on granitic sand: Ehrendorfer 
5927. tp.a (glabrous form): 2n=42 
tp.b (hirsute form): 2n=44 


California. Inyo County, Panamint Mountains, Wildrose Canyon, above charcoal 
kilns, ca. 7200 feet; metamorphic schists and sandstones; open Pinyon and Juniper, 
talus: Ehrendorfer & Dempster 5968. 


tp.a (hirsute form): 2n—44 
tp. b (hirsute form): 2n—44 
tp.c (glabrescent form): 2n= ca. 44 


tp.d (approaching G. mathewsiit 
= var. carneum Hilend & Howell, 1934): 2n=46 


California, Inyo County, Panamint Mountains, along trail from Mahogany Flat to 
Telescope Peak, ca. 8500 feet; metamorphic schists; talus, with Holodiscus, Arte- 
misia, etc.: Ehrendorfer 5943. fd.: 2n=44* 


California, San Bernardino County, San Bernardino Mountains, lower portion of 
Cushenbury Canyon, ca. 5000 feet; granitic talus, northeast slopes just above the 
wash, among open Pinyon-Juniper: Ehrendorfer 5903. id: (2n== 544 

tp.: 2n=44* 


probable hybrids with G. HyPOTRICHIUM subsp. SUBALPINUM 
California, Inyo County, east side of Sierra Nevada, Lone Pine Creek, 0.5 mile below 
Whitney Portal, steep ravine with northwest exposure, ca. 8370 feet; granitic boul- 
ders and talus; open Chrysothamnus and Artemisia: Ehrendorfer 5929. fd.: 2n=44* 
tp.: 2n=+44 


California, Inyo County, east side of Sierra Nevada, Little Cottonwood Creek above 
Lone Pine, ca. 8800 feet; granitic rock and sand; Cercocarpus, Holodiscus and Arte- 
misia: Ehrendorfer 5947. fd.:. 2n=44* 


approaching G. MAGNIFOLIUM 
Utah, Zion National Park, trail to The Narrows, ca. 4300 feet; sandy and gravelly 
talus below Navajo sandstone cliffs, eastern exposure; loose cover of herbs and scrub; 
Ehrendorfer & Dempster 5980. tp.a (hirsute form) : 2n=—44 
tp.b (hirsute form): Di==45 
tp.c (glabrous form): 2n=-42—44-45 
Arizona, Grand Canyon National Park, south rim, uppermost portion of Grand View 


1961] EHRENDORFER: GALIUM 119 


Trail, ca. 7000 feet ; arenaceous limestone; steep slope with herbs, among Cercocar pus, 
Amelanchier and Pinus edulis in northern exposure: Ehrendorfer & Dempster 5981. 
tp. a (somewhat hairy form): 2n=44 
tp. b (glabrous form): 2n— 42 


GALIUM PARISHII Hilend & Howell 
California, San Bernardino County, San Bernardino Mountains, top of San Gorgonio 
Mountain, 11,485 feet: leg. P. Raven 11,152. ide 2n==22+ 


California, Los Angeles County, San Gabriel Mountains, hills about 2 miles north 
of Big Pines, ca. 6500 feet; granitic talus in northwest exposure: open Artemisia with 
Pinus jeffryi: Ehrendorfer & Grant 5902. id. 2n=2 


California, San Bernardino County, Kingston Mountains, steep slopes south of King- 
ston Pass, ca. 5600 feet; granitic talus slopes and shady rock crevices: Ehrendorfer 
& Dempster 5972, 5973. fds. 222 

tps Zn = 22 


Nevada, Clark County, Charleston Mountains, Kyle Canyon, southwest slopes, south- 
east of Cathedral Rock camp site, ca. 7600 feet; limestone talus and rock crevices 
among open Pinus, Abies and Cercocarpus: Ehrendorfer & Dempster 5976. 

Ure =2n=22 


GALIUM SERPENTICUM Dempster [G. watsonit (Gray) Heller sensu Ehrendf.] 
Washington, Asotin County, Blue Mountains, below the crest overlooking Indian 
Tom Creek, 30 miles southwest of Asotin, ca. 5300 feet; basalt, rocks and fine talus 
below, open bushy and herbaceous pioneer growth: Ehrendorfer & Ownbey 5956, 5957. 
tp. ds 2n=—22 


California, Modoc County, Warner Mountains, Mount Bidwell, southeast side of 

plateau top, ca. 8000 feet ; basalt talus; bushy and herbaceous pioneer vegetation with 
Artemisia, grasses, etc.: Ehrendorfer & Stebbins 5911, 5912, 5913. 

Boye 2n—22 

2n=225 

tp. II-1: 2n=22 

id ic 2n 22 


California, Modoc County, Mount Bidwell, southwest side of plateau top, below rim, 
ca. 7600 feet; east slope, steep basaltic talus; similar vegetation: Ehrendorfer & Steb- 
bins 5914. fd... 2n—227, 


var. FUBERULUM (Piper) Dempster [G. watsonii (Gray) Heller subsp. puberu- 
lum Ehrendf. ] 
Washington, Kittitas County, Wenatchee Mountains, west-southwest slope in upper 
Beverley Creek, ca. 4500 feet; serpentine rock and talus, open pioneer vegetation: 
Ehrendorfer & Kruckeberg 5958, 5959. idi2*2n==22" 
tp. 1; 2n=22 


Washington, Kittitas County, Liberty, knoll above Boulder Creek, ca. 3500 feet; 
Eocene sandstone shale; sandy talus slope with pioneers: Ehrendorfer & Krucke- 
berg 5960. tp; 1s 2n==22 


Washington, Kittitas-Chelan County, south slope near Blewett Pass, ca. 4000 feet; 
sandy talus: Ehrendorfer & Kruckeberg 5962. tps 2022 


GALIUM STELLATUM Kell. subsp. EREMICUM (Hilend & Howell) Ehrendf. 

California, Inyo County, Darwin Falls, about 3 miles south of Lone Pine-Death 
Valley highway, 3000 feet; rock crevices in canyon walls, metamorphic schists; desert 
scrub and some cacti: Ehrendorfer & Dempster 5940. fd< -2n==22 


California, San Bernardino County, hills south of Highway 66, 7 miles northeast of 
Essex, ca. 1000 feet; steep rocky slope, gneiss; desert scrub (Larrea, etc.): Ehren- 
dorfer & Dempster 5982. tp.: 2n==22 


120 MADRONO [Vol. 16 


Nevada, Clark County, Valley of Fire State Park, east entrance opposite Elephant 
Rock, ca. 2000 feet; rocky north slope, Triassic sandstone; desert scrub: Ehrendorfer 
& Dempster 5978. tpi? 2n==22 


UNNAMED TAXON (6x race) 

California, Modoc County, Warner Mountains, Horse Mountain, south of and 
toward summit, ca. 8500 feet; basalt talus and rocks; grassy and shrubby pioneers: 
Ehrendorfer & Stebbins 5909, 5910. ids: 2n—=065 


GALIUM ROTHROCKII Gray subsp. ROTHROCKIZ (a monoecious member of section 
Lopho-Galium, not directly connected with the G. multiflorum complex). 
Arizona, Grand Canyon National Park, south rim, uppermost portion of Grand View 
Trail, ca. 7000 feet ; arenaceous limestone; steep slope with herbs, among Cercocarpus, 
4Amelanchier, and Pinus edulis'in northern exposure (together with G. munzii ap- 
proaching G. magnifolium) ; Ehrendorfer & Dempster 59814. tps: 2n=22 


DISCUSSION 


In spite of general dioecious sex differentiation, an eu-polyploid series 
2x—4x—6x with x11 has developed in the Galium multiflorum complex, 
just as in practically all of the hermaphrodite groups of this genus which 
have been checked cytologically so far. Polyploidization is less advanced 
than in some other groups with the majority of the races still diploid and 
only one very local hexaploid known. In the European section Lepto- 
Galium, for instance, polyploidization has proceeded to the 10x level, with 
the most widely spread and successful types (G. pumilum, G. rubrum, 
G. marchandit) on the 8x level (Ehrendorfer 1954). 

Sexual differentiation in the G. multiflorum complex exemplifies an 
evolutionary trend established in a number of other New World groups 
of the genus as well. No sex chromosomes have been recognized so far. 
Sex differentiation in the complex must be similar in character to that in 
Rumex subgenus Acetosella or Melandrium (Love and Sarkar 1956, and 
literature cited there). In these groups genetic sex determiners for the 
heterogametic sex are so strongly epistatic that polyploidization does 
not upset the 1:1 segregation mechanism (XY =~, XXXY= 4, 
XXXXXY = 7). 

The basic evolutionary differentiation of the G. multiflorum complex 
is accompanied by only very slight visible structural changes in chromo- 
somes. Occasional spontaneous aberrations give a clue as to the origin 
of these. | 

As a result of the total evolutionary differentiation within this species 
complex, “marginal” and ‘‘extreme” positions in respect to distribution, 
ecology, and morphology are taken by diploids. Known sympatric con- 
tacts between diploids are rare. Galium matthewsu and G. parishu grow 
in mixed populations in the Kingston Mountains of southeastern Cali- 
fornia, but there are no indications of hybridization. This must be owing 
to the development of internal barriers, possibly involving chromosome 
structure. In contrast with the diploids, the polyploids are intermediate 
in distribution, ecology, and many morphological characters. There is 
some additional evidence that they are of hybrid origin and hybridize 


1961] EHRENDORFER: GALIUM 121 


much more freely with each other than do the diploids (e.g., G. munzu 
and G. hypotrichium subsp. subalpinum on the eastern slope of the south- 
ern Sierra Nevada). The present cytological findings substantiate the hy- 
pothetical racial diagram and interpretation developed by the author in 
1956 (fig. 7): therein the polyploid G. grayanum subsp. glabrescens, 
G. munzii, and the partly polyploid G. hypotrichium form central “‘hot 
spots” of the complex, while all the marginal racial “‘cornerstones”’ are 
diploid. The general evolutionary situation therefore is very similar to 
that in other Galium groups, e.g. the section Lepto-Galium (Ehrendorfer 
1954, 1955). 

Details of cytological behaviour of the polyploid members of the CG. 
multiflorum complex are very much in conformity with facts already 
known from European species: diminution of chromosome size in poly- 
ploid, stabilization of chromosome pairing, possibly via some influence 
on chiasma frequency, occasional irregularities of chromosome distribu- 
tion into the gametes caused by formation of multi- and univalents and 
consequent appearance of biotypes with aberrant chromosome numbers. 
Intra-individual somatic instability of chromosome number has not been 
reported for Galzaum before, but the phenomenon seems not to be rare in 
polyploids (Gottschalk 1958, and literature cited there). 

The cytological data here set forth point the way to some revisions in 
the taxonomic treatment of the G. multiflorum complex. The tetraploid 
Galium grayanum subsp. glabrescens, for example, should perhaps be ac- 
corded specific status, but in other cases specific separation on the basis 
of different ploidy levels seems imposible and highly impractical as within 
G. hypotrichium where even subspecific separation of the very closely 
adjacent and very similar 2x and 4x populations on Sonora Pass (Sierra 
Nevada) is an extreme procedure. Obviously no generalized rules can be 
applied in diploid-polyploid racial pairs, as has been previously shown 
with European Lepto-Galium and other groups. Taxonomic questions con- 
cerning the Galium multiflorum complex will be dealt with in other papers 
of this series. 


SUMMARY 


1. The western North American Galium multiflorum complex consists 
of sexual, dioecious races. Chromosome numbers have been established 
for mest of the recognized taxa, including counts from 51 populations and 
more than 100 individuals. Primarily the chromosome numbers form an 
eu-ployploid series 2x—4x—6x with x11. The distribution of the various 
diploids and polyploids is mapped. 

2. Cell size (PMC’s, pollen) is generally increased in polyploids. 

3. Chromosome size and number of chiasmata are generally reduced in 
polyploids. 

4. In two tetraploid plants a certain intra-individual somatic instability 
of chromosome number has been established; this has not been observed 
in diploids. 


122 MADRONO [Vol. 16 


5. In polyploids there is a limited amount of multi- and univalent for- 
mation during PMC meiosis, with consequent irregularities of chromosome 
number in the gametes. 

6. In four tetraploid (but never in diploid) populations, individuals 
with different standard chromosome numbers (2n—=42—44—-45—46) have 
been observed. | 

7. No differences in the chromosome sets of male and female plants 
could be demonstrated. 

8. In PMC meioses and pollen mitoses some consequences of sponta- 
neous chromosome aberrations (bridges, fragments) have been found. 

9. Chromosome sets of various diploids and polyploids are quite similar, 
but there are certain differences (e.g. in the SAT-chromosomes) as a 
result of structural changes. 

10. Cytological findings are briefly discussed from comparative evolu- 
tionary and taxonomic viewpoints. 


Museum of Natural History 
Vienna, Austria 


LITERATURE CITED 


DempstTeER, L.T. 1959. A re-evaluation of Galium multiflorum and related taxa. 
Brittonia 11:105-122. 

EHRENDORFER, F. 1954. Phylogeny and evolutionary mechanisms in Lepto-Galium. 
Rapp. et Comm. VIII. Intern. Congr. Bot., Paris 1954, Sect. 4:82-84. 

. 1955. Hybridogene Merkmals-Introgression zwischen Galium rubrum L. 
s.str.und G. pumilum Murr. s. str. (Zur Phylogenie der Gattung Galium, IV.) 
Osterr. Bot. Zeitschr. 102:195—-234. 

. 1956. Survey of the Galium multiflorum complex in western North Amer- 
ica. Contr. Dudley Herb. 5:1-36. 

—. 1958. Ein Variabilitatszentrum als “fossiler” Hybrid-Komplex: Der ost- 
mediterrane Galium graecum L.-G. canum Req.-Formenkreis. Eine Mono- 
graphie. (Zur Phylogenie der Gattung Galium, VI.) Osterr. Bot. Zeitschr. 105: 
229-279. 

GottscHaALK, W. 1958. Uber Abregulierungsvorgange bei kiinstlich hergestellten 
hochpolyploiden Pflanzen. Zeitschr. Abst. Vererb. 89:204-215. 

Love, A. and N. Sarkar. 1956. Cytotaxonomy and sex determination of Rumex 
pauciflorus. Canad. Jour. Bot. 34:261-268. 


A NEW SPECIES OF LYCIUM IN NEVADA 
CoRNELIUS H. MULLER 


A unique endemic Lycium occurs in Nevada in the area of the Atomic 
Energy Commission Nevada Test Site on Frenchman Flat. The plant was 
first discovered by Dr. William H. Rickard who remarked its extremely 
viscid, 4-merous corolla, and who suspected that it represented an un- 
described species. It was collected in quantity by him, V. K. Carpenter, 
and Janice E. Beatley in the course of ecological investigations and sub- 
sequently by Dr. Beatley at my request. Material was submitted almost 


1961] MULLER: LYCIUM 123 


simultaneously to C. L. Hitchock and to me. Professor Hitchcock con- 
curred in the opinion that the plant was undescribed and offered an analy- 
sis of its position in the genus but generously disclaimed any desire to 
undertake its publication. To him and to Dr. Beatley I am indebted for 
the opportunity to study this interesting material. Dr. Philip Wells has 
gathered considerable information on the distribution of the species. He 
discovered a large population on the southeasterly bajada of the Spotted 
Range and in northwestern Clark County. I am indebted to him for guid- 
ance to these localities. 

The species typically grows on gravelly alluvium, predominantly lime- 
stone, in association with Atriplex confertifolia at the upper limit of Lar- 
rea divaricata and about the lower limit of Coleogyne ramosissima. It 
extends onto the playa clay on Frenchman Flat and onto quartzite beds 
on the lower slopes of the Spotted Range. 


Lycium rickardii sp. nov. Frutex 0.5 m. altus, glaber; ramis albis 
spinosis; foliis 3-12 vel 18 mm. longis, 1.5—3 vel 6 mm. latis, 4—-8-fascicu- 
latis, obovatis vel spatulatis, floribus solitariis, pedicellis 0.5 mm. longis; 
calyce campanulato, tubo 6 mm. longo, lobis 4, 3 mm. longis; corolla 
tubuliformi, tubo 8-14 mm. longo, extra et intra viscido, lobis 4, circa 
3 mm. longis; staminibus inclusis, corollae tubi partem supra mediam 
adhaerentibus, basi corollae intraque villosis; bacca subrotunda, 4-5 mm. 
longa, 2- vel 3-sperma, in calyce inclusa. 

Intricately branched shrub about 0.5 m. tall or less; branchlets spinose, 
their smooth bark strikingly glaucous, weathering gray and fissuring after 
2 or 3 years, the wood very soft and brittle; leaves 3 to 12 or even 18 mm. 
long, 1.5 to 3 or 6 mm. broad, in fascicles of 4 to 8, spatulate to obovate, 
the gradually narrowed base scarcely distinguishable from the blade, the 
apex broadly rounded, very thick and succulent, the midrib scarcely dis- 
cernible in dried leaves, slightly glaucous green, the epidermal cells almost 
vescicular, giving the false impression of puberulence upon drying; flow- 
ers usually solitary in the leaf fascicles, the pedicels less than 1 mm. long; 
calyx highly variable, accrescent during and after anthesis, very succulent, 
the tube 6 mm. long, about 4 mm. broad, the 4 lobes 1 to 3 mm. long, mere 
teeth or broadly deltate-ovate, obtuse, broadly spreading or rarely erect; 
corolla white, the throat and veins suffused with purple or green, strictly 
tubular or narrowly funnel-shaped (the basal portion shrinking strongly 
upon drying), 8 to 14 mm. long, 2.5 to 3.5 mm. broad, the lobes about 
3 mm. long, ovate, apically rounded, rotate or reflexed with age, 4-merous 
but a fifth lobe sometimes represented by a vascular bundle and an abor- 
tive petal, both outer and inner surfaces markedly viscid-glandular (this 
not apparent in dried material) ; stamens as many as the corolla lobes, an 
abortive petal sometimes carrying a full-sized staminode; filaments equal, 
plain, adnate about 34 to 44 the height of the tube, strikingly pubescent 
with long hairs in the basal 3 or % of their length; anthers included by 
the throat; gynoecium bilocular, glabrous, with thin yellow walls, on a 


124 MADRONO [Vol. 16 


J 


Fic. 1. Lyctum rickardu sp. nov.: A-C, flowers, x 3; D, interior of corolla, x 3; 
E, fruiting calyx, x 3; F, mature seed, & 3; G, longitudinal section of ovary at an- 
thesis parallel to the septum showing a pair of ovules on a single placenta, x 9; 
H, longitudinal section of ovary at anthesis at right angles to the septum showing one 
of each pair of ovules on each placenta, x 9; I, cross section of ovary, X 9; J, ma- 
ture fruit, x 3. Drawn by Isabelle Haller from fresh material (Beatley 2, 3, 4 and 
Muller 10945) except in the instances of F and J which were based on Rickard and 
Beatley, 2 May 1959. 


thickened bright red disc, each locule 2-ovulate, no abortive ovules and 
no lower cells developing in the disc; style at anthesis reaching about half 
the length of the corolla; stigma green, slightly 2-lobed, irregular; fruit 
subrotund, about 4 or 5 mm. long and broad, completely enclosed in the 
accrescent, urceolate calyx and surmounted by the constricted throat and 
divergent sepals, the exocarp cartilaginous, the disc remaining red and 
slightly fleshy but not enlarging with the fruit, 1 or 2 seeds maturing in 
each locule of the ovary, thus producing a 2-seeded or, more often, a 3- 


1961] MULLER: LYCIUM 125 


seeded fruit, the seeds flattened on the common face, about 3 or 4 mm. 
long, minutely pitted, the aborted ovule(s) always in the original position. 

NEVADA. Nye County: codominant in shadscale scrub at 4100 feet on south- 
facing bajada of the Spotted Range, 16.6 miles west of Indian Springs, 8 April 1961, 
Muller 10940, 10941, 10943, 10944, 10945 (holotype UCSB, sheet no. 8765), 10946, 
10947; codominant in shadscale scrub at 4200 feet in southerly foothills of the 
Spotted Range, 14 mile above the highway and 16.6 miles west of Indian Springs, 
8 April 1961, Muller 10948, 10949, 10950; “east of playa” on Frenchman Flat, 
2 May 1959, Rickard and Beatley s.n. (from which the fruit is described) ; “south of 
playa, near Lycium Plot 4;” “northwest of playa;” “near playa”:! all on French- 
man Flat, 2 April 1959, Rickard, Carpenter, and Beatley s.n.; with Larrea and Atri- 
plex south of playa at 3100 feet on Frenchman Flat, 11 April 1961, Beatley 2; with 
Larrea south of playa at 3100 feet on Frenchman Flat, 11 April 1961, Beatley 3; with 
Larrea east of playa at 3100 feet on Frenchman Flat, 11 April 1961, Beatley 4. Clark 
County: rare in shadscale scrub at edge of foothills 144 miles south of Indian 
Springs, 8 April 1961, Muller 10951. 

All specimens cited are deposited in the herbarium of the University 
of California, Santa Barbara, and duplicates are being distributed. 

Lycitum rickardii negotiates Hitchcock’s key (1932) past L. pallidum 
Miers (p. 202) but fits neither ““G. Fruit 2—4-seeded, with 1 or 2 fertile 
seeds in the top of each carpel, and abortive ovules in compartment below” 
nor “GG. Fruit not as above, with more than four seeds.”’ Rather, each 
locule contains two ovules, and there is no division of the locule into 
compartments as in L. macrodon and L. puberulum. If one ovule aborts, 
it appears on the same placenta and in the same locule with the matured 
seed. In this respect L. rickardii agrees with L. shockleyi Gray, an emend- 
ed description of which was published by Muller (1940). The Hitchcock 


(1932) key may be emended as follows: 


G. Fruit 2-4 seeded. 
H. Fruit with each carpel divided into two locules, the upper bearing 1 or 2 seeds 
and the lower locule bearing aborted ovules; calyx not enclosing fruit. 
L.macrodon and L. puberulum 


HH. Fruit with one locule to each carpel, the seeds or abortive ovules totalling 
2 in each locule; calyx enclosing fruit. 


I. Fruit with an irregular suture or fold on one or both sides, filaments adnate 


nearly full length, the anthers appearing almost sessile. . . L.shockleyi 

II. Fruit lacking a suture or fold, filaments free in upper % of 14 of their 
lengethSe).) a Sree eee to wees a ao a0 SDarterardy 

GG. Fruit with more than 4 seeds. ‘is Teste we (ee be 7 Lacoo peri et. seq: 


The relationship of L. rickardu to L. shockleyi is apparent in the num- 
ber of ovules in each carpel, the lack of a lower compartment with abor- 
tive ovules, and the 4-merous condition. However, the partially free fila- 
ments and lack of a suture or fold on the side of the fruit clearly distin- 
guish it from L. shockleyi. Its fruit and stamen characters suggest L. cali- 
fornicum Nutt. ex Gray, but in the latter species the corolla is much 
smaller with proportionately shorter lobes and each locule contains a 


1 The latter two collections bear the following notes: “Corollas 4-merous, shining 
viscid within and without; shrub less than 2 feet high.” 


126 MADRONO [Vol. 16 


single ovule. Professor Hitchock (personal letter to William H. Rickard, 
22 November 1960) pointed out the intermediacy of L. rickardii between 
L. californicum and “such species as L. macrodon and L. puberulum.” It 
might be added that both L. rickardu and L. shockleyi stand in this posi- 
tion with L. rickardi closer to L. californicum and L. shockleyi more simi- 
lar to L. macrodon Gray, L. puberulum Gray, and L. cooperi Gray. 

It is extremely likely that L. rickardu is somewhat more widely distrib- 
uted than at present known. In the rather copious material at hand there 
is no evidence that heavy doses of irradiation at the Test Site are in any 
way responsible for the characters of L.rickardu. The longevity of these 
plants insures their being older than the Test Site, and their essential uni- 
formity with those of the southerly and southeasterly range of the species 
makes it highly unlikely that the characters here noted might have arisen 
as a result of somatic mutation. 

Department of the Biological Sciences, 


University of California, Santa Barbara, 
Goleta, California. 


LITERATURE CITED 


Hitcucock, C.L. 1932. A monographic study of the genus Lycium of the Western 
Hemisphere. Ann. Missouri Bot. Gard. 19:179-374. 

Mutter, C. H. 1940. New and otherwise noteworthy plants of the Southwest. 
Madrono 5:152-158. 


SOME RECENT OBSERVATIONS ON PONDEROSA, JEFFREY 
AND WASHOE PINES IN NORTHEASTERN CALIFORNIA 


JoHN R. HALLER 


In an earlier paper (1959), I suggested that Pinus jeffreyi Grev. and 
Balf. is less susceptible to cold than is P. ponderosa Dougl. ex Lawson, 
and that for this reason P. jeffreyi replaces P. ponderosa at high altitudes 
in the mountains of California. Dr. Willis W. Wagener reported recently 
(1960), however, that established trees of P. ponderosa survived at least 
as well and occasionally better than P. jeffreyi following periods of severe 
cold in northeastern California. The purpose of the present paper is to 
present additional information on the pines of northeastern California 
which I believe will show that there is no discrepancy between Wagener’s 
observations and my own, and that, in fact, they even reinforce one 
another. 

In our respective papers, Dr. Wagener and I were discussing examples 
from different areas—his from northeastern California, mine mostly 
from cismontane California, that portion of the state lying to the west 
of the Sierra-Cascade crest. I deliberately omitted a discussion of the 


1961] HALLER: PINUS 127 


northeastern Californian pines because the great bulk of the Ponderosa 
Pine in California occurs in the cismontane portion of the state, and be- 
cause northeastern California is climatologically and floristically more 
closely related to the Great Basin region than to the rest of California. 
In addition, the ecological, genetic, and taxonomic relationships of the 
pines in northeastern California appear to be far more complex than in 
the remainder of the state. I am currently preparing a paper which will 
attempt to describe their relationships with each other and with the pines 
farther to the east. 

There is considerable evidence available which supports the idea that 
Pinus ponderosa from cismontane California is more susceptible to low 
temperatures than either P. jeffreyi or P. ponderosa from more interior 
localities. The precise altitudinal zonation on the western slope of the 
Sierra Nevada, with P. ponderosa occupying the lower elevations and P. 
jeffrevxi occupying the higher, is very suggestive of a difference in cold 
tolerance (Table 1). As I have indicated previously (1959), in the narrow 
zone where these species overlap, P. jeffreyi nearly always occupies the 
colder sites, such as canyon bottoms and the margins of low-lying 
meadows. Pearson (1931) reported that first-year seedlings of P. pon- 
derosa from the Sierra National Forest (on the western slope of the 
Sierra Nevada) planted near Flagstaff, Arizona, were killed by a Novem- 
ber freeze, whereas seedlings from several other western states planted 
at the same locality were not injured. Weidman (1939) reported that 
young trees of P. ponderosa from the vicinity of Weed, California (west 
of the Cascade crest), planted near Sandpoint in northern Idaho were 
killed at the age of 12 years when the temperature fell rapidly from 45°F 
to —12°F. Trees of approximately the same age from 19 localities in Ore- 
gon, Washington, Idaho, Montana, South Dakota, Colorado, Utah, New 
Mexico and Arizona, planted in the same site as the Californian trees, 
survived the cold. Temperatures lower than —12°F occasionally occur 
within the range of P. jeffreyi in the western Sierra Nevada and within 
the range of both P. ponderosa and P. jeffreyi in transmontane north- 
eastern California, but virtually never within the cismontane range of P. 
ponderosa (Table 1). The temperatures shown in Table 1 are the lowest 
that occurred in 1949, which was a year of record-breaking cold for 
many of the stations. This is the same year that Wagener observed fairly 
extensive damage to both P. ponderosa and P. jeffreyi in California. Table 
1 shows that P. ponderosa occurs in the warmer cismontane localities and 
P. jeffreyi in the colder, but in transmontane California the two species 
grow in equally cold localities which are frequently colder than the coldest 
cismontane P. jeffrey localities. 


1 Neither the temperature nor the year of its occurrence were given by Pearson. 
However, the coldest November temperature on record for Flagstaff for the years 
1906-1930 is —4°F. This is probably not sufficiently cold to kill mature P. ponderosa 
from cismontane California (see Table 1), but was cold enough to be fatal to the 
more susceptible seedlings. 


128 MADRONO [Vol. 16 


TABLE 1. MintmuM TEMPERATURES IN 1949 IN CALIFORNIA. 
Based on official records of the United States Weather Bureau* 


A. Cismontane Localities 


LocaLiITy County ELEV. TEMP. 
Ponderosa Zone 
Placerville Eldorado 1900 ft. 14°F 
Sierra City Sierra 4200 oy 
Calaveras Grove Calaveras 4800 5° 
Yosemite Valley Mariposa 4000 oe 
Mt. Shasta (town) Siskiyou 3500 1° 
South Entrance, Yosemite N. P. Mariposa 5100 —3° 
Ponderosa-Jeffrey Zone 
Giant Forest Tulare 6400 —4° 
Grant Grove Tulare 6700 —6° 
Lake Spaulding Nevada 5000 —8° 
Jeffrey Zone 
Manzanita Lake Shasta 5800 —3° 
Huntington Lake Fresno 7000 —10° 
Twin Lakes Alpine 7900 —24° 
Soda Springs Nevada 6700 —27° 


B. Transmontane Localities 
Ponderosa Zone 


Cedarville Modoc 4700 ft. —20°F 

Mount Hebron Siskiyou 4200 —22° 

Alturas Modoc 4300 —31° 
Ponderosa-Jeffrey Zone 

Truckee Nevada 6000 —19° 

Sierraville Sierra 5000 —25° 

Boca Nevada 5500 —41° 
Jeffrey Zone 

Woodfords Alpine 5600 —10° 

Bridgeport Mono 6400 —31° 


* In some of the localities listed above, the species concerned does not occur in 
the immediate vicinity of the weather station. However, discrepancies between ° 
weather station temperatures and those in the adjacent pine localities have been 
kept to a minimum by selecting stations in situations that are ecologically similar to 
the pine localities and never more than a few miles removed from them. Stations 
have also been selected to show the maximum temperature variation within each zone. 


The evidence given above indicates that Pinus ponderosa from north- 
eastern California is different physiologically from that on the cismontane 
slopes, since it survives temperatures lower than those that have killed 
cismontane P. ponderosa in experiments. This physiological difference is 
reflected in the relative distribution of P. ponderosa and P. jeffreyi in 
northeastern California, where these two species occur together over much 
more extensive areas than on the western slopes of the mountains. 
Furthermore, there is no tendency for P. jeffreyi to occupy the colder 


1961 | HALLER: PINUS 129 


sites within these extensive mixed stands, as occurs on the cismontane 
slopes. 

In addition to the physiological—distributional differences between 
cis- and transmontane Pinus ponderosa, there are morphological differ- 
ences. To begin with, the P. ponderosa from northeastern California is 
far more variable than that in the western Sierra Nevada, ranging from 
essentially identical to that farther west to something strikingly different. 
Because of this high variability, it is difficult to generalize about par- 
ticular character differences. However, one character, needle thickness, 
shows relatively consistent differences between the west and east sides 
of the Sierra-Cascade crest. In all of the cismontane localities where I 
have measured needle thickness in mixed stands of P. ponderosa and P. 
jeffrevi, the mean thickness is greater in P. jeffrey. In the typical ex- 
amples given in Table 2A, the needles of P. jeffrew average 0.19 mm. 
thicker than those of P. ponderosa, and the difference between the two 
species is highly significant at each of the three localities shown. Just the 
reverse is true on the east side of the Sierra-Cascade crest, where the 
needles of P. ponderosa average 0.12 mm. thicker than those of P. jeffreyi 
(Table 2B). However, east of the crest the differences between the species 
range from essentially nil at Sierraville to very pronounced at Hobart 
Mills. It is noteworthy that at Sierraville, where the needles of P. pon- 
derosa are thinnest, the population is not unusually variable and is in 
most respects very similar to cismontane P. ponderosa. On the other hand, 
the P. ponderosa near Hobart Mills, which has much thicker needles than 
P. jeffrevi, is tremendously variable and for the most part very different 
from cismontane P. ponderosa (Haller, 1957). This thick-needled Hobart 
Mills population is located just three miles from Boca, which frequently 
has the lowest winter temperatures of any station in California (Table 1), 
suggesting that thick needles may be adaptively advantageous in cold 
climates. 

There are at least three possible causes for the physiological and mor- 
phological differences between the cis- and transmontane Pinus ponder- 
osa in northern California: environmental modification, differential selec- 
tion from a heterozygous gene pool, and introgressive hybridization. A 
certain amount of environmental modification no doubt occurs in all 
populations of P. ponderosa, as the experiments of Weidman, the Insti- 
tute of Forest Genetics, and my own observations (1957 and in press) 
have shown. However, these same experiments and observations show 
that the greater proportion of all physiological and morphological traits 
is genetically determined and cannot be ascribed solely to modification. 
Selection from a heterozygous gene pool has probably been the principal 
mechanism that has enabled P. ponderosa to occupy so many diverse 
habitats in western North America and to differentiate into a number of 
geographical races or subspecies. The geographical pattern of variation 
shown by these races is, however, a subtle one, and I have found that the 


130 MADRONO [Vol. 16 


TABLE 2. MEAN NEEDLE THICKNESS IN PONDEROSA, JEFFREY AND WASHOE 
PINE POPULATIONS 


A. Cismontane Localities 


MEAN NEEDLE WIDTH SIGNIFI- 
LocaLity CouNTY ELEVATION SAMPLE CANCE OF 
(feet) SIZE PONDEROSA JEFFREY _DIFFERENCE 


Silver Fork, 
American River Eldorado 6400 10 132mm. 1.54mm. .004 
Ebbetts Pass 


Highway Calaveras 6300 24 ~=«:1.60 1.75 005 
Shasta Valley Siskiyou 4500 25) 1:65 1.83 <.001 
Mean of cismontane Ponderosa and Jeffrey 

populations: 1.52 e/a 
B. Transmontane Localities 
Sierraville Sierra 5000 25 1.78 LS dies 
Dixie Mtn. 

Game Refuge Plumas 5700 10-32 01 1.88 075 
Hobart Mills Nevada 5800 50 =. 2.10 1.87 <.00003 
Mean of transmontane Ponderosa and Jeffrey 

populations: 1.96 1.84 
WASHOE JEFFREY 
Mt. Rose, Nev. Washoe 7200 25 2.05mm. 1.88 .0006 
Warner Mtns. Modoc 7500 142.13 2.024 a0 
Mean of transmontane Washoe and Jeffrey 
populations: 2.09 1.95 


* The Warner Mountain Jeffrey Pine population is not sympatric with the Washoe 
Pine population, but is located a few miles away. 


overall variability of the populations is usually about the same from one 
locality to the next. As already stated, many of the P. ponderosa popula- 
tions in northeastern California differ strikingly from nearby popula- 
tions and also display great variability. Such a pattern would be expected 
if hybridization were taking place. 

The probable introgressant of Pinus ponderosa in northeastern Cali- 
fornia is P. washoensis Mason and Stockwell. One of the more outstand- 
ing characters of P. washoensis is its thick needles, which also characterize 
the variable P. ponderosa populations at Hobart Mills and Dixie Moun- 
tain Game Refuge (Table 2B). In addition there are other characters of 
P. washoensis, such as compact cones and short needles, which are preva- 
lent in these variable P. ponderosa populations. I am still in the process 
of analyzing data from P. ponderosa, P. washoensis, and P. jeffreyi in 
northeastern California, but I am reasonably certain that the hybridiza- 
tion suggested here will be confirmed by further investigation. 

The only published record of Pinus washoensis is from the type lo- 
cality, on the watershed of Galena Creek, Mount Rose, Washoe County, 
Nevada (Mason and Stockwell, 1945). This locality is about 14 miles 
east of the Hobart Mills P. ponderosa population. A much more extensive 


1961] HALLER: PINUS 131 


stand of P. washoensis occurs in the Warner Mountains, Modoc County, 
California. The best stands, which include many trees that are four feet 
in diameter, are located in the southern part of the range, in the general 
vicinity of the Patterson Ranger Station. Specimens from this area have 
been deposited in the herbarium of the University of California at Santa 
Barbara. Both the Mount Rose and Warner Mountain stands of P. 
washoensis occur above the 7000 foot elevation, apparently too high for 
P. ponderosa. The Mount Rose stand occurs sympatrically with P. 
jeffrevi, whereas the Warner Mountain stand occurs almost entirely above 
a narrow zone of P. jeffreyi. Additional stands of typical P. washoensis 
might well occur on other sufficiently high peaks in northeastern Cali- 
fornia. Pinus washoensis also occurs sporadically at lower elevations, for 
example in the variable “P. ponderosa” population near Hobart Mills. 
Very few individuals at this locality are “good” P. washoensis, but many 
of the trees in this apparent hybrid swarm are more similar to P. 
washoensis than they are to typical P. ponderosa (Haller, 1957). 

The factors which limit the distribution of individuals of taxa such as 
Pinus ponderosa, P. jeffreyi, and P. washoensts are extremely difficult to 
circumscribe exactly. However, some idea of the relative cold suscepti- 
bilities of members of these taxa might be obtained from a series of experi- 
ments. For example, seeds of P. ponderosa and P. jeffreyi from the same 
site in the western Sierra Nevada and from northeastern California could 
be grown under uniform conditions, and the seedlings subjected to in- 
creasing intensities of cold. If a sufficient number of experiments were 
made, it would be apparent whether P. ponderosa is ever more susceptible 
to cold than P. jeffreyi, or if either or both species vary from one locality 
to another in their relative susceptibility. The results of any such experi- 
ments would have to be interpreted with caution, however. For example, 
P. ponderosa and P. jeffreyt from the same site in the western Sierra 
Nevada (near the upper altitudinal limit of the former and the lower 
limit of the latter) might be found to have an identical tolerance for 
cold. The upward migration of P. ponderosa could nonetheless be checked 
at this point by cold, because the species has exhausted its genetic poten- 
tial for cold tolerance. Pinus jeffreyi, on the other hand, could have a 
much greater potential cold tolerance, which might be expressed only at 
higher altitudes, where it would be favored by natural selection. 

The possibility also exists that the upward migration of Pinus ponder- 
osa is not checked by low winter temperatures, but by insufficiently high 
temperatures during the growing season. Pearson (1931) stated that low 
summer temperatures are the principal deterrent to the success of P. 
ponderosa when it is planted at elevations above its normal range in the 
San Francisco Mountains of Arizona. In this region, P. ponderosa is 
limited to the valleys and lower slopes of the mountains where summer 
maximum temperatures are higher but winter minima are lower than in 
the Douglas Fir zone immediately above. 


132 MADRONO [Vol. 16 


Evidence has been submitted in this paper that Pinus ponderosa from 
cismontane California is more susceptible to cold than is P. jeffreyi, that 
P. ponderosa from northeastern California is at least as tolerant of cold as 
is P. jeffreyi and that the spread of P. ponderosa to higher elevations is 
checked in Arizona by low summer temperatures rather than by extremes 
of cold in winter. Most of this evidence is indirect, and a more precise 
determination as to the factor or factors which limit P. ponderosa in its 
many different habitats will have to await the outcome of future experi- 
ments. For the present, it appears reasonable to postulate that low tem- 
peratures, whether in the form of low winter minima or low summer 
maxima, play an important role in limiting the distribution of P. 
ponderosa. 

Department of Biological Sciences 


University of California, Santa Barbara 
Goleta, California 


LITERATURE CITED 


Hatter, J. R. 1957. Taxonomy, hybridization and evolution in Pinus ponderosa and 
P. jeffreyi. Doctoral dissertation, Univ. Calif., Los Angeles. 

—-———. 1959. Factors affecting the distribution of P. ponderosa and P. jeffreyi in 
California. Madrono 15:65-71. 

Mason, H. L., anp P. STOCKWELL. 1945. A new pine from Mount Rose, Nevada. 
Madrono 8:62. uA 

Pearson, G. A. 1931. Forest types in the Southwest as determined by climate and 
soil. U.S. Dep. Agr. Tech. Bull. 247:1-144. 

WAGENER, W. W. 1960. A comment on the cold susceptibility of Ponderosa and 
Jeffrey Pines. Madrono 15:217-219. 

WEIDMAN, R. H. 1939. Evidence of racial influence in a 25-year test of Ponderosa 
Pine. Jour. Agr. Res. 59:855-887. 


INFLUENCE OF TEMPERATURE AND OTHER FACTORS 
ON CEANOTHUS MEGACARPUS SEED GERMINATION 


ELMER BurRTON HapDLey! 


One of the striking characteristics of chaparral is the absence of any 
kind of seedlings beneath mature, undisturbed stands. After such dis- 
turbances as bulldozing or fire, however, an abundance of seedlings 
appears, suggesting that scarification or heat make the germination 
possible (Cooper 1922, Went et al. 1952, Horton & Kraebel 1955, Quick 
1959). 

The density and dryness of this chaparral brush cover in California, 
the accumulation of large quantities of dry litter beneath the brush, and 
the Mediterranean type climate, all combine to create an extreme fire 


1 The author wishes to express his thanks to the University of California at Santa 
Barbara for extending the full use of its facilities and to Dr. L. C. Bliss and Dr. C. H. 
Muller for generous aid and advice in the course of this study. 


1961] HADLEY: CEANOTHUS 133 


hazard during the long summer droughts. Many chaparral species are so 
specifically adapted to the resulting periodic fires as to indicate a long 
history of subjection to recurrent fires in the geologic past (Jepson 1925). 
Horton (1945) found that Ceanothus species in general have a life span 
of about forty years. In areas unburned for a period of at least forty 
years, however, other species have, to a large extent, replaced Ceanothus 
because there has been no regeneration by Ceanothus. 

Two fire responses are very common among chaparral species: re- 
sprouting from burls (underground root crowns) that are not killed by 
fire, and heat induced seed germination due to cracking of impervious 
seed coats. Arctostaphylos glandulosa is an example of a species that 
shows the first type of fire response. Almost no seedlings of this “stump 
sprouting” species are found following fires. Ceanothus megacar pus, 
wherein the entire shrub is usually killed outright by fires, represents the 
second type of response—regeneration by seed following fire. Other 
shrubs, such as Adenostoma fasciculatum, exhibit both types of fire 
responses. 

In addition to the resprouting shrubs and seedlings of chaparral species, 
a recently burned-over area contains many annuals and _ short-lived 
perennial herbs and subshrubs as well as weeds. Eventually the chaparral 
vegetation dominates and the under vegetation perishes. 

Fire or heat induced increases in germination were first investigated by 
Wright (1931), who found that oven heating greatly increased the ger- 
mination of a number of chaparral species, including Ceanothus mega- 
carpus. Sampson (1944), and Went e¢ al. (1952) have also investigated 
increased germination of chaparral species due to fire. Stone & Juhren 
(1951), investigating germination of seeds of Rhus ovata, found that 
temperatures of 80°C induced the rupturing of impervious seed coats and 
thus permitted water to reach the embryos. Quick (1935), using seeds of 
several species of Ceanothus, also found that heat cracked the impervious 
seed coats. Stratification, following the heat treatment, resulted in further 
germination increases in these species. 

In none of these investigations, however, were the possible modifying 
effects of natural field conditions on heat induced germination considered. 
In the present investigation an attempt was made to determine the effect 
of temperature, of mechanical injury, and of accumulated litter on the 
germination of seed of Ceanothus megacar pus. 


METHODS 


Except for field observations and collecting the seeds, all other phases 
of this investigation were carried out in the greenhouse of the University 
of California at Santa Barbara. All results obtained must, therefore, be 
considered no more than suggestive of what might occur under field 
conditions. 


2 Nomenclature is that of Munz (1959). 


134 MADRONO [Vol. 16 


Large quantities of Ceanothus megacarpus seeds were gathered from 
the Santa Ynez Mountains above Santa Barbara, California, in June, 
1959, and air dried for two months in the laboratory. Voucher specimens 
are on deposit at the herbarium of the University of California at Santa 
Barbara. 

All experiments were conducted in controlled temperature boxes, using 
sterilized petri dishes with moistened filter paper. All seeds were treated 
with the fungicide, Semesan. Seeds were germinated at a temperature 
regime of 26°C and 17°C (alternating 12-hour periods at each tempera- 
ture). Each experiment was conducted for a thirty-day period, and ger- 
mination in all cases was defined as emergence of the radicle. 

In order to determine the effect of heating or cutting of the seed coat on 
Ceanothus megacar pus seed germination, a first experiment was run using 
three lots of seeds. The first lot was subjected to a temperature of 100°C 
for 5 minutes in an electric oven, the second to mechanical rupturing of 
the seed coat at the micropilar end with a razor blade, and the third lot 
served as controls (no heating or cutting). Each of the above three treat- 
ments consisted of three replicates using 25 seeds per dish. These seeds 
were moistened with distilled water and germinated as described in the 
previous paragraph. 

In order to determine the possible effects of leaf litter on germination, 
a second series of experiments was set up using actual leaf material on 
top of the seeds in the petri dishes. Sets of seeds were prepared, each 
with three replicates as in the previous experiment, i.e., heated seeds, cut 
seeds, and controls. Fresh leaves, duff (dead fallen leaves not yet decayed 
beyond recognition), and ashed duff (7 grams of duff ashed at 700°C in 
a muffle furnace for 45 minutes) of Adenostoma fasciculatum were used 
in equal quantities on each of the first three sets of seeds, while a fourth 
set of three petri dishes of seeds was left as a control. Adenostoma fascicu- 
latum was chosen because it is one of the most abundant and cosmopolitan 
species comprising the chaparral community. Except for the presence of 
leaf material, the seeds were germinated with distilled water as in the 
previous experiment. To test the hypothesis that any stimulation of seed 
germination due to duff is really a mineral effect, another part of this 
second experiment was run using a modified Hoagland’s solution contain- 
ing trace elements (Hoagland and Arnon 1950) in place of the distilled 
water. 

In the third experiment, the effect of using leached duff (partial re- 
moval of minerals) was investigated. The duff was leached in eight 
changes of distilled water for 96 hours before being placed over the seeds. 

All data were subjected to an analysis of variance using the individual 
degrees of freedom technique (Snedecor 1956). Space does not permit its 
inclusion, but a complete analysis of variance for the data may be found 
in Hadley (1960). 


1961 | HADLEY: CEANOTHUS 135 
TABLE 1. EFFECT OF TEMPERATURE, MECHANICAL RUPTURING OF SEED COAT, AND 
ADENOSTOMA FASCICULATUM LEAF LITTER ON CEANOTHUS MEGACARPUS 


SEED GERMINATION. 


Percentage Germination* 


Treatment Cut seed Heated seed Untreated seed 
(100°C for 5 min.) 
I. Distilled water 86 23 0 
II. ‘Distilled water + 
Adenostoma duff 87 hs 1 
Adenostoma duff ashed 41 24 0 
Adenostoma fresh leaves 24 13 @) 
Control 83 DS 0 
Hoaglands solution + 
Adenostoma duff 95 oi 0 
Adenostoma duff ashed 31 29 0 
Adenostoma fresh leaves 21 ZN ) 
Control 88 77 1 
III. Leached duff 85 41 1 
Unleached duff 87 80 4 


* All experiments run in replicates of 3 with 25 seeds in each replicate. 
Experiment I—Effect of heating and cutting of seed coat on Ceanothus megacarpus 
seed germination. 
Experiment I]—Influence of Adenostoma fasciculatum leaf litter and/or Hoagland’s 
~ solution on the germination percentage of Ceanothus megacarpus seeds. 
Experiment I]I—Effectiveness of leached Adenostoma fasciculatum duff vs. unleached 
duff in stimulating C. megacarpus seed germination. 


RESULTS 


Under the conditions of this investigation, germination of Ceanothus 
megacarpus seeds is facilitated by either heating or cutting the seed coats. 
As shown in Table 1, however, mechanical rupturing is the more effective 
treatment. 

Presence of the various Adenostoma leaf material did not significantly 
affect the germination of the untreated controls, but did significantly 
affect the germination of the heated and cut seed (Table 1). Presence of 
duff over the heat treated seeds significantly stimulated the germination 
percentage of these seeds, resulting in a four-fold increase over the con- 
trols. This increase was shown by later experiments to be attributable to 
increased minerals made available by the decayed duff. Since cut seeds 
displayed maximum germination with or without duff being added, the 
effect of adding duff could not be measured accurately in the case of the 
cut seed. 

Presence of fresh leaf material resulted in a significant reduction of 
cut seed germination, possibly due to the presence of an inhibitor in the 
fresh leaves (Naveh 1960). Application of ashed duff also caused a sig- 
nificant reduction in the germination of cut seed. 

Substitution of Hoagland’s solution in place of distilled water re- 
sulted in little change in the germination per cent of seed treated with 


136 MADRONO [Vol. 16 


either ashed duff or fresh leaves (Table 1). These results show that 
fresh leaf material or ashed duff have the same inhibiting effect, whether 
distilled water or Hoagland’s solution is used. Germination per cent of 
heated seeds is very similar, whether treated with distilled water plus 
Adenostoma duff or only with Hoagland’s solution. Leaching of this duff 
(partial removal of minerals) significantly reduced the effectiveness of 
duff in stimulating germination of the heat treated seeds (Table 1). 


DISCUSSION AND CONCLUSIONS 


Ceanothus megacarpus seed germination percentage was increased by 
heating these seeds for 5 minutes at 100°C. Mechanical rupturing of the 
seed coat was found to have an effect similar to heating, but to a greater 
degree. This would suggest that the stimulatory effect of heating involved 
in this species is primarily one of rupturing a previously impervious seed 
coat, thus allowing water to reach the embryo. The smaller increases in 
germination in the case of the heated seed may be due to injury to some 
of the embryos due to heat, to a random cracking of the seed coat away 
from the micropilar end which might hamper radicle emergence, or to the 
variability in seed coat thickness (some of the seed coats may not be 
cracked by this particular temperature). 

Application of leaf material of another chaparral species, Adenostoma 
fasctculatum, has a definite effect on the germination percentage of Cea- 
nothus megacarpus. Adenostoma duff enhances the germination of heat 
treated C. megacarpus seeds. Since a similar effect was obtained when 
Hoagland’s solution was substituted in place of the distilled water and 
duff, this stimulation of germination can possibly be attributed to increas- 
ing mineral concentration provided by the decaying duff. The conclusion 
that germination was stimulated by available minerals in the duff is sup- 
ported by the fact that there was marked reduction in percentage of 
seed germination when the seeds were topped by leached duff. 

The apparent inhibition of germination by fresh leaves of Adenostoma 
may be due to the presence of an inhibitor or inhibitor complex in these 
leaves (Naveh 1960). The reduced germination of cut seed in the pres- 
ence of ashed duff may be due to increased pH. Sampson (1944), using 
several grass species, has noted this ash inhibition, which was attributed 
to increased pH. The germination percentage of heated seeds in the pres- 
ence of ashed duff, remained similar to that of non-treated heated seeds. 
This would suggest that ashed duff did not have an inhibitory effect on 
heated seed. 

What part heating, mechanical rupturing, and plant litter actually play 
in the field can only be suggested, for in the field the situation created in 
the laboratory does not exist. Obviously, rupturing of the seed coat due 
to mechanical injury can be of only minor ecological significance in the 
field except where bulldozing, sharp deer hoofs, or some other agent may 
crack the seed coats. Accidental rupturing may therefore account for at 


1961] HADLEY: CEANOTHUS 137 


least a portion of the few young seedlings that are sometimes found in 
disturbed but unburned areas. 

The extremely low percentage of germination noted for untreated seed 
may serve to explain the field observation that young seedlings of 
Ceanothus megacar pus are not found under undisturbed, mature chapar- 
ral in which this species is a constituent. Heat treatment of Ceanothus 
megacar pus seeds by fires should be of tremendous importance in the re- 
population of burned areas. This increase in germination percentage due 
to cracking of the seed coat by heat could account for the abundance of 
Ceanothus seedlings found immediately following a fire. 

It must be remembered that all chaparral fires are not alike; they 
differ in intensity, duration, and temperatures reached during the fire. 
Some fires consume both shrub crowns and litter; others are principally 
confined to the shrub crowns leaving pockets of litter unconsumed. There- 
fore a differential destruction of duff by fire is noted in the field. Some 
fires could easily provide the required temperatures for the duration of 
time necessary to crack the seed coats and yet not burn away all of the 
duff that would be present. Other fires, even though they might burn 
away most or all of the duff and seeds, would still provide temperatures 
necessary to crack the seed coats of those seeds which were buried in and 
therefore protected by the soil. Thus breaking of the seed coat by heat 
would account for the Ceanothus seedlings that Quick (1959) and others 
have encountered after fires in the chaparral. 

Only Adenostoma fasciculatum leaf litter was used in these experi- 
ments. It is possible that the duff and fresh leaves of many of the other 
chaparral species might exhibit similar effects on Ceanothus megacar pus 
seeds and those of several other chaparral species. This is a subject that 
would indeed be worth further investigation. 


Department of Botany 
University of Illinois 
Urbana, Illinois 


LITERATURE CITED 


Cooper, W. S. 1922. The broad-sclerophyll vegetation of California. Carnegie Inst. 
Wash. Publ. 319. 

Haptey, E. B. 1960. Influence of fire and other factors on Ceanothus megacarpus 
seed germination. Unpublished Master’s thesis. University of Illinois Library, 
Urbana. 

Hoacianp, D. R., and D. I. Arnon. 1950. The water-culture method for growing 
plants without soil. Univ. Calif. Agr. Exp. Sta. Bull. 347. 

Horton, J. S. 1945. Vegetation of the San Bernardino Forest. Ms. Oct. 3, 1945. (as 
cited by Shantz, H. L. 1947. The use of fire as a tool in the management of the 
brush ranges of California. Calif. Div. Forestry, Calif. State Board of Forestry, 
Sacramento). 

, and C. J. Krarper. 1955. Development of vegetation after fire in the 
chamise chaparral of southern California. Ecology 36:244-262. 

Jerson, W. L. 1925. A manual of the flowering plants of California. Assoc. Students’ 
Store, Univ. Calif., Berkeley. 

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


138 MADRONO [Vol. 16 


NaveH, Z. 1960. The ecology of Chamise as affected by its toxic leachates. Bull. Ecol. 
Soc. Amer. 41:56—57. 
Quick, C. R. 1935. Notes on the germination of Ceanothus seeds. Madrono 3:135-140. 
. 1959. Ceanothus seeds and seedlings on burns. Madrono 15:79-81. 
Sampson, A. W. 1944. Plant succession on burned chaparral lands in northern Cali- 
fornia Univ. Calif. Agr. Exp. Sta. Bull. 685. 

SNEDECOR, G. W. 1956. Statistical Methods (5th Ed.). Iowa State Coll. Press, Ames. 

STONE, E. C., and G. JUHREN. 1951. The effect of fire on the germination of the seeds 
of Rhus ovata Wats. Am. Jour. Bot. 38:368-372. 

WENT, F. W., G. JUHREN, and M. C. JuHREN. 1952. Fire and biotic factors affecting 
germination. Ecology 33:351-364. 

Wricnut, E. 1931. The effect of high temperature on seed germination. Jour. Forest. 
29:679-687. 


REVIEWS 


Flora of the Santa Cruz Mountains of California. A manual of the vascular 
plants. By John Hunter Thomas. viii + 434 pages, 249 figs. and 16 photos, 1 map. 
Stanford University Press, Stanford, California. 1961. $8.50. 

The first impression, upon taking up the “Flora of the Santa Cruz Mountains,” 
is of an attractive, well-designed book with clear typography, generous spacing, 
indented keys, good illustrations, and with an adequate binding. The Stanford 
University Press is to be congratulated upon producing a volume of exceptionally 
fine appearance. 

Although the book is entitled ‘“‘Flora of the Santa Cruz Mountains,” it encom- 
passes the whole San Francisco peninsula, and thence southward to the Pajaro River 
and from the ocean east to the middle of the Santa Clara Valley. Coverage is com- 
prehensive, including both native and introduced plants. The number of kinds of 
introduced plants occurring spontaneously is amazing; 31 per cent of the 1799 taxa 
listed fall into this category. 

For each of the species listed, Thomas gives the scientific name, common name, 
habitat, localities in the area, time of blooming, and place of origin for introduced 
species. Sometimes elevation is given and, occasionally, associated species. Brief 
comments, often on taxonomic problems, are made for some species. Synonyms are 
included only for convenience in referring to the same taxon in other regional and 
sectional floras. Specimens are not cited except in a few instances. There are no new 
names or combinations. 

The flora is written for the serious beginner as well as the trained botanist. The 
beginner, especially, will appreciate the 250 line drawings which are from the “Illus- 
trated Flora of the Pacific States.” As a result of better spacing and better paper 
they are clearer and more attractive than many of the original reproductions in the 
Illustrated Flora. The common names also appear to correspond to those used by 
Abrams. Possibly the influence of the latter flora may be responsible in part for the 
recognition of certain families, for example, Melanthaceae, Parnassiaceae, Hydrangea- 
ceae, Grossulariaceae, Amygdalaceae, Malaceae, Mimosaceae, Caesalpinaceae, Mono- 
tropaceae, Pyrolaceae, Vacciniaceae, Convallariaceae, Amaryllidaceae, all of these 
segregated from the Liliaceae, Saxifragaceae, Rosaceae, Fabaceae, and Ericaceae. 
Nevertheless, Thomas’ taxonomic concepts are, in general, conservative. For example, 
Berberis rather than Mahonia is recognized; Montia exigua is considered as synony- 
mous with M. spathulata; ssp. decurrens of Eriogonum nudum is not recognized. 
However, Dudleya (not Echeveria) and Horkelia are used; Allium breweri is consid- 
ered distinct from A. falcifolium; and all the forms of Arctostaphylos in the Santa 
Cruz Mountains are accorded specific status. The varietal designation is usually em- 
ployed rather than the subspecific except when the latter designation has been used 


1961] REVIEWS 139 


in a recent monograph. The arrangement of families in general follows the sequence 
proposed by Engler and Prantl. 

Part I consists of 33 pages of introductory material and 13 pages of keys to the 
divisions, classes, subclasses, and families. A map shows place names and supplements 
the description of the area. The geology (4 pages including a stratigraphic profile) 
is discussed by Dr. Earl E. Brabb. Monthly and yearly average temperatures and 
average rainfall are given for seven stations. Classification of the vegetation follows 
Munz and Keck insofar as it may be applied to the Santa Cruz Mountains. The more 
characteristic plants are listed for each of the plant communities, and photographs 
illustrate most of them. 

Ten pages are devoted to a discussion of the composition and relationships of the 
flora. The number of native species is approximately 1246 in the Santa Cruz Moun- 
tain area of 1386 square miles, compared to 1004 in Marin County (529 square miles), 
700 in the Mount Hamilton Range (1500 square miles), and 530 on Mount Diablo 
(55 square miles). Five distributional patterns are recognized, whereas Campbell and 
Wiggins recognized 16 for the whole state. Endemic in the area are 10 species, 3 sub- 
species, 11 varieties, 2 forms, and 1 hybrid. Some are closely restricted to certain geo- 
logical formations. 

Lists, together with localities, are given: 1) of taxa reaching their southern limits 
of distribution in the area, 2) of taxa reaching their northern limit, 3) of those with 
affinities with the inner Coast Ranges, 4) of the more obligate serpentine taxa, and 
5) of plants with a disjunct distribution to the north. Plants of sandhills and marshes 
are also discussed. The list of taxa with “their northern limits of distribution in the 
Coast Range in the Santa Cruz Mountains” applies to the “Outer Coast Ranges” only, 
as seven species are included which occur somewhat farther north in the Inner Coast 
Range, on Mount Diablo. These are Anemopsis californica, Malacothamnus halli 
(Sphaeralcea fasciculata), Osmorrhiza brachypoda, Linanthus ambiguus, Pholistoma 
membranaceum, and Salvia mellifera. 

Of the 34 taxa occurring in the Santa Cruz Mountains, but regarded by Thomas 
as being typically species of the Inner Coast Ranges, 24 grow in the Mount Hamilton 
Range. Only 19 are on Mount Diablo, 13 of which are common to Mount Diablo and 
the Mount Hamilton Range. Whether Helianthella castanea should be considered a 
plant of the inner ranges is debatable. 

Thomas’ list of “the more obligate local serpentine plants” (p.31) caused the 
writer considerable surprise, as a number of plants which are common and wide- 
spread on Mount Diablo are included. The following species are not associated in the 
writer’s mind with serpentine although the records indicate that they may, at times, 
grow on serpentine: Festuca pacifica, F. reflexa, Koeleria macrantha (K. gracilis, K. 
cristata), Calochortus venustus, Allium serratum, Lewisia rediviva, Astragalus gam- 
bellianus, Sanicula bipinnatifida, and Rigiopappus leptocladus. Of the remaining 
taxa, more than half commonly or often grow on serpentine, but are by no means 
limited to it. Ten or fewer may be truly obligate serpentine plants. Perhaps there is 
here a difference of opinion as to the interpretation of the phrase “more obligate.” 
However, the list does include species of rather widely differing ranges of tolerance 
with respect to the substratum and soils. Further observations on the relation of 
serpentine tolerant species to their substratum need to be stimulated. 

Part I closes with a brief résumé of the history of botanical collecting in the 
area. Photographs of six collectors are presented. The annotated catalogue of vas- 
cular plants comprises Part II. Part III consists of a list of 34 general references and 
a glossary of technical terms. Part IV consists of an index of place names, an index 
of common names, and an index of scientific names. 

The “Flora of the Santa Cruz Mountains of California” presents a synthesis of 
the present knowledge of the flora of the Santa Cruz Mountain area based on many 
collections by others as well as by Dr. Thomas. It represents a great deal of work 
and is a worthy volume which does credit to its author and will be useful to many. 
In conclusion, we will all, I am sure, concur wholeheartedly with the author’s wish 


140 MADRONO [Vol. 16 


that, by acquainting more people with the plants around them, this volume will serve 
as “a stimulus, however slight, toward more permanent protection of our environ- 
ment.”’—Mary L. Bowerman, Department of Botany, University of California, 
Berkeley, California. 


Principles of Plant Breeding. By R. W. ALLarp. xi + 485 pp. John Wiley & Sons, 
Inc. New York and London. 1960. $9.00. 


In the Preface, the author states that “Principles of Plant Breeding” is designed 
primarily to serve as an undergraduate text for students in agriculture. The aim of 
the book is to stress principles, and to illustrate them with appropriate examples. 
This task has been accomplished with a high degree of competence. Allard writes 
with clarity, precision and force. For this reason it should not be difficult for an 
undergraduate with some training in biometry, and a semester course in genetics, to 
follow his closely reasoned explanations and interpretations. The entire book is ar- 
ranged to serve as a text for a two-semester course, but it is conveniently segmented 
so it can be adapted to the needs of a one-semester or one-quarter course. In addi- 
tion to its pedagogical function, this book can be studied with profit by the profes- 
sional plant breeder. It will serve to broaden his outlook and invigorate his research. 


The material used to illustrate the principles is slanted to some extent towards 
cereal and forage crops, but this is not unnatural. More thorough information about 
plant breeding techniques and procedures is available for this group of crops than for 
fruit, vegetable, fiber or ornamental crops. A few more examples could, however, 
have been drawn from cotton and possibly other crops. 


As one could anticipate, knowing his interests, the author is particularly sure- 
footed and lucid in chapters concerned with quantitative genetics, population genetics, 
systems of mating and heterosis. But other sections, for example, ‘Breeding methods 
with cross-pollinated crops,” “Breeding for disease resistance,” and “Polyploidy,” are 
also discussed with equal skill. 


This reviewer can suggest only one feature that would perhaps increase the useful- 
ness of the book. A set of carefully composed questions and problems at the end of 
each chapter might serve as a source of understanding and stimulation. This has been 
done to seme extent by inserting questions in the legends of a few figures. More 
complete development of this aspect might add to the teaching value of the book. 


The references are not copious, but adequate for the purpose. The book is notable 
for an unusually low incidence of typographical errors. A glossary of terms used in 
plant breeding and a good index add to its serviceability. 


It has taken time for plant breeding to bridge the gap between art and science. 
“Principles of Plant Breeding” is likely to be marked as a significant milestone in 
establishing plant breeding as a full-fledged scientific discipline —THomas W. WuirT- 
AKER, U.S. Horticultural Field Station, La Jolla, California. 


NOTES AND NEWS 


The Smithsonian Institution is reprinting Paul C. Standley’s Trees and Shrubs 
of Mexico, Contr. U.S. National Herbarium, vol. 23, 1920-26, Parts 1 (pp. xviii + 
1-169), 2 (xxxvii + 171-515), 3 (pp. xxviii + 517-848), and 5 (ii + 1313-1721), in 
2 paper-bound volumes containing pts. 1-3 and pt. 5, respectively. The price of these 
4 parts is $20, postpaid. Part 4 (pp. xxxiv + 849-1312), which is available in the 
original 1924 edition published by the U.S. National Museum, will be enclosed free 
of charge. Orders should be accompanied by check and should be addressed to: 
Publications Distribution Section, Smithsonian Institution, Washington 25, D.C. 


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 


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search, observation, and history. Back volumes may be obtained 
from the Secretary at the following rates: 


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The subscription price of MApDRONo is $6.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
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ONO 


VOLUME 16, NUMBER 5 JANUARY, 1962 


Contents 


PAGE 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF 
THE GENUS MIMULUS (SCROPHULARIACEAE). V. 
THE CHROMOSOMAL HOMOLOGIES OF M. GUTTATUS 
AND ITS ALLIED SPECIES AND VARIETIES, Barid B. 
Mukherjee and Robert K. Vickery, Jr. 141 


Mito S. BAKER (1868-1961), Herbert L. Mason 155 


CYTOLOGICAL OBSERVATIONS ON ADIANTUM 
x Tracyi C. C. HALL, Warren H. Wagner, Jr. 158 


TAXONOMIC AND NOMENCLATURAL NOTES ON 
PLATYDESMA (HAWAII) AND A NEW NAME FOR A 
MELICOPE (SOLOMON ISLANDS), Benjamin C. Stone 161 


A NEw SPECIES OF GALIUM IN CALIFORNIA, 
Lauramay T. Dempster 166 


A NEw SPECIES OF CRYPTANTHA (SECTION CIRCUM- 
SCISSAE) FROM CALIFORNIA AND Two RECOMBINA- 
TIONS (SECTION CIRCUMSCISSAE AND SECTION 
ANGUSTIFOLIAE), Kunjamma Mathew and 
Peter H. Raven 168 


Review: C. L. Porter, Taxonomy of Flowering 
Plants (John Mooring) 1a 


A WEST AMERICAN JOURNAL OF BOTANY > SPaRe 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


| 
| 
i} 


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 
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN Benson, Pomona College, Claremont, California 

HERBERT F. CoPpELAND, Sacramento College, Sacramento, California 

Joun F. Davinson, University of Nebraska, Lincoln 
MItprep E. Martuias, University of California, Los Angeles 24 
Marion OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
IrA L. WiccINs, Stanford University, Stanford, California 


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


Business Manager and Treasurer—JoHN H. THomAS 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Roxana S. Ferris, Dudley Herbarium, Stanford University, Stanford, 
California. First Vice-President: Edward C. Stone, School of Forestry, University 
of California, Berkeley. Second Vice-President: Thomas C. Fuller, Department of 
Agriculture, State of California, Sacramento. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding Sec- 
retary: Margaret Bergseng, Department of Botany, University of California, Berke- 
ley. Treasurer: John H. Thomas, Dudley Herbarium, Stanford University, Stanford, 


California. 


1962 | MUKHERJEE & VICKERY: MIMULUS 141 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF THE 
GENUS MIMULUS (SCROPHULARIACEAE). V. THE 
CHROMOSOMAL HOMOLOGIES OF M. GUTTATUS 
AND ITS ALLIED SPECIES AND VARIETIES 


Barip B. MUKHERJEE AND ROBERT K. VICKERY, JR. 


The purpose of this study! was to investigate the chromosomal homol- 
ogies of Mimulus guttatus and its allied species and varieties. This was 
done by observing the pairing behavior of the chromosomes in the pollen 
mother cells of F, and a few F. hybrid plants obtained from crossing 
various members of the M. guttatus complex and its related taxa. This 
large, highly polymorphic group (Grant, 1924; Pennell, 1951) of gay- 
looking, yellow-flowered plants consists of a vast number of typically 
isolated populations of various sizes, of differing combinations of morpho- 
logical characteristics, and of assorted taxonomic ranks. Its populations 
grow by springs and streams from the Aleutian Islands to southern Mex- 
ico and from the Pacific coast to the Mississippi River in North America, 
and in the Andes and their foothills in South America. 

Of these populations, thirty-seven which exhibited much of the mor- 
phological variation and much of the diversity of geographical origin of 
the entire group were sampled for this investigation. The thirty-seven 
cultures which were grown from these populations represented at least 
eighteen different species and varieties (table 1). They included all of 
the most common forms of the group as well as several rare ones. These 
representative cultures were crossed in all possible combinations (Vick- 
ery, 1956a, 1956b) and most of the resulting seeds were sown. Some of 
the combinations failed to produce flowering hybrids due to the presence 
of crossing barriers of various strengths (Vickery, 1956a, 1956b, 1959). 
Consequently the cytological analysis was limited to the successful hy- 
brids (table 2), which were chiefly combinations of /. guttatus with each 
of the related species plus a few combinations among the latter. 

The method of fixing the buds was the same as that employed in the 
previous investigations (Mukherjee and Vickery, 1959, 1960); i.e., fix- 
ation in 2 parts absolute ethanol to 1 part glacial acetic acid saturated 
with ferric acetate. After 24 hours in the fixative, the buds were trans- 
ferred to 70% ethanol if they were to be stored for later study. In prepar- 
ing the slides, the anthers were dissected from the buds and then squashed 
in a drop of iron-aceto-carmine stain. In many cases the most interesting 
hybrids produced only one to several flowers which in turn might or might 
not yield one or more cells suitable for cytological examination. Conse- 


1 This investigation was supported by the National Science Foundation and the 
University of Utah Research Fund. Most of these results form a portion of the dis- 
sertation of the senior author submitted to the Faculty of the University of Utah 
in partial fulfillment of the Ph.D. requirements. 


Maprono, Vol. 16, No. 5, pp. 141-172. January 31, 1962. 


142 MADRONO LVol. 16 


quently many of our counts are based on suboptimum numbers of cells. 
Nevertheless certain trends are clearly apparent in the results. Most of the 
cells analyzed were drawn with the aid of a camera lucida and many were 
photographed. In addition, numerous F, hybrid plants were pressed, 
mounted, and deposited for future reference in the Garrett Herbarium 
of the University of Utah. 

The chromosomes of the different species and varieties ranged in size 
from dots as small as one-half micron in diameter to ovals as large as one 
micron wide by two micra long (see figure 1 and the previous papers of 
this series). Despite this variation in size, which probably was due in 
part to differing orientations of the chromosomes in the cells, the chromo- 
somes were so similar in general appearance that rarely could we identify 
the individual chromosomes contributed by each parent to a particular 
F, hybrid. Therefore, our analysis of chromosomal homologies was car- 
ried out at the genome level rather than at that of the individual chrom- 
osomes. We observed the amount and regularity of chromosome pairing 
in as many pollen mother cells as possible in over 60 different interspecific 
and intervarietal hybrids (table 2). 

In our cytological examinations of the parental species and varieties 
(Vickery, 1955; Mukherjee, Wiens, and Vickery, 1957; Mukherjee and 
Vickery, 1959, 1960), we found no indication of true autosyndesis in any 
of the cultures. However, under the fixation schedule employed, several 
of the annual races of M. guttatus exhibited chromosome stickiness which 
simulated autosyndesis and secondary chromosome associations (Vick- 
ery, 1959). This difficulty was overcome by techniques suggested by 
Doctors Harlan Lewis and Henry J. Thompson of the University of Cali- 
fornia at Los Angeles. 

According to our findings, the basic genome in the group appears to be 
that of the diploid species, specifically, of the type of M. guttatus with its 
n—14 chromosomes. Possibly M. guttatus and/or the other diploid 
species such as M. nasutus, M. glabratus var. utahensis, M. tilingu, etc. 
(see table 1) may be ancient tetraploids inasmuch as a distantly related 
species, M. mohavensis Lemmon, has n=7 chromosomes (Carlquist, 
1953). However, at the present time there is no evidence for this hypoth- 
esis. Therefore, tentatively we may consider the whole group, M. guttatus 
and its relatives, to consist of species and varieties at the diploid (n=13, 
14,15), tetraploid (n=26, 30, 31,32), and hexaploid (n=45, 46) chrom- 
osomal levels, with one to several examples of aneuploidy at each level. 
The data at hand suggest to us that this polyploid series is built up on a 
base number of x15 which presumably is an aneuploid derivative of 
the basic genome of n=14 chromosomes so commonly found in this group 
of species. 

Despite the tremendous range of morphological and physiological vari- 
ation within 1. guttatus itself (Grant, 1924), all of its populations thus 
far counted have n=14 chromosomes. The chromosomes of the inter- 
population F, hybrids exhibited normal chromosome pairing (table 2) 


1962 | MUKHERJEE & VICKERY: MIMULUS 143 


f?, 14 Il 


5001, M. guttatus, n = 14 


5001, M. guttatus, n= 14 
(subsp. guttatus). 


5001, M. guttatus, n= 14 
(subsp. guttatus). 


(subsp. guttatus), 


Xx x X 


5834, M. guttatus, n = 14 


5003, M. guttatus, n = 14 
(var. puberulus). 


5004, M. guttatus, n= 14 
(subsp. litoralis). 


(subsp. guttatus). 


14 Il 


@ 13 II, 21 y, 

P) 
7 eos op, 
. wd \ ‘ 4 


5001, M. guttatus, n= 14 


= 14 5004, M. guttatus, n = 14 
(subsp. guttatus). 


5001, M. guttatus, n = 
(subsp. guttatus). (subsp. guttatus). 


x x x 


5346, M. guttatus, n= 14 


5007, M. guttatus, n= 14 5010, M. guttatus, n- 147 1,2 
(M. arvensis). 


(M. lyratus). (M. laxus). 


Pd wD 14 II on 

- %. > 5 4 48 % J 
yy & 

apes anys 1 2 Mee 


5017, M. guttatus, n= 14 


5017, M. guttatus, n= 14 5052, M. guttatus, n = 14 
(M. cordatus). 


(M. cordatus). (subsp. guttatus). 


x x x 


5064, M. laciniatus, n = 14 5339, M. laciniatus, n = 14 5044, M. nasutus, n = 14 


6 10 20 micra 


Fic. 1. Meiotic chromosomes of intraspecific Fy hybrids of the Mimulus guttatus 
complex as defined in this article. All configurations at or near first metaphase. Camera 
lucida drawings at an original magnification of * 2,520, reduced to X 1,260. 


144 MADRONO [Vol. 16 


except in a few cases which were probably the result of factors of tech- 
nique. One exception turned up in the lone intra-guttatus Fy hybrid plant 
analyzed (5346 & 5839), which had an extra chromosome. 

Due in part to this generally pervasive cytological homogeneity, M. 
guttatus has been treated in this article in the broad sense of Grant 
(1924). Except for M. platycalyx, the various species segregated from 
M. guttatus by Pennell (1951) have been included in it (table 1). Several 
of our cultures could be assigned to these segregate species, specifically, 
culture 5007 to M. lyratus, 5017 to M. cordatus |so identified by F. W. 
Pennell (see Alexander & Kellogg 2844, UC 696,020, from which our 
seeds came) |, 5010 to MW. daxus, and 5346 to M. arvensis (table 1.) How- 
ever, these species intergrade morphologically with each other and with 
M. guttatus. Cytologically, they all appear to possess the same genome 
(figure 1). Genetically, they are fully interfertile or else separated by 
no stronger barriers than those that occur within M. guttatus in the strict 
sense (Vickery, 1959). Therefore, with these facts in mind, we have 
treated M. lyratus, M. cordatus, M. laxus, and M. arvensis as synonyms 
of M. guttatus. 

Two of the M. guttatus cultures, 5009 and 5010, from Mather, Cali- 
fornia, were known to be aberrant in that they occasionally produced 
microspores with n=13, 15, or 16 chromosomes instead of the usual n—=14 
(Mukherjee and Vickery, 1959). The present investigation showed that 
at least some of these aneuploid microspores were functional as can be 
observed in the chromosome complements of their F, hybrids (figure 2 
and table 2). A comparable situation was observed in M. luteus (table 2) 
and in M. glabratus var. fremontu (figure 4). These facts are significant 
because they indicate a likely mechanism for the production of aneuploid 
plants. Possibly the already-mentioned intra-guttatus Fy plant (5346 x 
5839) with the extra chromosome arose in this manner. Such aneuploid 
plants might in turn lead to the establishment of aneuploid populations 
and even, eventually, of aneuploid varieties and species such as common- 
ly occur in the M. guttatus complex and its relatives (table 1). 

Mimulus guttatus hybridized readily with M. laciniatus, with the n=14 
form of M. nasutus, and with M. glaucescens. In the first two cases the 
hybrids were fertile and their pollen mother cells exhibited regular chrom- 
osome pairing, although the regularity of chromosome pairing was con- 
siderably decreased in the F., individuals studied. The F, hybrids of 
M. guttatus * M. glaucescens were nearly sterile, and their chromosomes 
showed reduced pairing (figure 2 and table 2). Probably M. laciniatus 
and M. nasutus should be considered simply as well-marked varieties of 
M. guttatus, whereas M. glaucescens should be treated as a nearly distinct 
species. 

Mimulus platycalyx (n=15) and the n=13 form of M. nasutus both 
hybridized with M. guttatus, but the F, hybrids produced were partially 
sterile (Vickery, 1956b). The chromosomes in the pollen mother cells 
of the hybrids showed regular pairing of 13II and 11 for M. guttatus 


1962 | MUKHERJEE & VICKERY: MIMULUS 145 


V2” AT o@. 
3 ae eo 
ele 7 de Lf 
4 yes fey), Sop at a 
= Per S 
5327, M. nasutus, n = 13, 5014, M. guttatus, n= 14, 5017, M. guttatus, n = 14, 
x x Xx 
5003, M. guttatus, n = 14, 5653, M. glaucescens, n= 14, 5653, M. glaucescens, n = 14, 
13 1l, J1 
* 
14 II - 
ee 
iy agin 
sty = 6 0 10 20 
¢ »> = micra 
5752, M. platycalyx, n = 15, 5339, M. laciniatus, n = 14, 
x x 
5010, M. guttatus, n= 14% 1,2. 5327, M. nasutus, n = 13, 


Fic. 2. Meiotic chromosomes of interspecific Fy hybrids of Mimulus guttatus com- 
plex. All configurations at or near first metaphase. Camera lucida drawings at an 
original magnification of * 2,520, reduced to x 1,260. 


< M.nasutus and 1411 and 1I for M. guttatus « platycalyx except where 
the aberrant culture 5010 was a parent (figure 2 and table 2). Despite the 
aneuploidy and genetic differentiation of these species, both their genomes 
appear to be basically homologous to that of M. guttatus, just as the 
genomes of the various aneuploid species of section Alatae of Nicotiana 
are homologous (Goodspeed, 1954). Clearly, both species are an integral 
part of the M. guttatus complex although their accurate specific desig- 
nation must await a detailed study of the relevant literature and type 
specimens. 

One of the F, hybrids of M. nasutus (5751; n= 14), & M. nasutus 
(5327; n= 13), which would be expected from the foregoing results to 
show marked sterility was highly fertile instead (Vickery, 1956b). It set 
an average of 50 seeds per capsule whereas the hybrid resulting from the 
reciprocal combination averaged only 3 seeds per capsule. The cytological 
analysis of several pollen mother cells of one of the F. hybrids of the 
highly fertile cross provided the explanation. The fertile hybrid was an 
amphiploid with n=27 chromosomes (table 2). 

The alpine species M. tilingi did not hybridize readily with MW. gut- 
tatus. The hybrids that were formed with the exception of two possible 
amphiploids, produced sterile flowers if they flowered at all. However, the 
pollen mother cells of these F,; hybrids of M. guttatus « M. tilingii ex- 
hibited regular chromosome pairing in three of the four cells available for 
study (figure 3 and table 2). These cells came from hybrids involving 


146 MADRONO . [Vol. 16 


both the n=14 and n=15 races of M. tilingu var. tilingu. Therefore, the 
basic, or what we may call the M. guttatus genome of chromosomes, prob- 
ably is present in both the chromosomal races of M. tilingi var. tilingu 
also, although the crossing barriers between this species and the M. gut- 
tatus complex are so nearly complete as to warrant its exclusion from 
the complex. In fact, /. tilingit is itself the main species of another com- 
plex of related species and varieties. 

Mimulus tiling var. corallinus (n=24) forms completely sterile hy- 
brids with M. guttatus and with M. tilingu var. tilingu. In the majority 
of the pollen mother cells examined in the F, hybrids of M. guttatus « 
M. tiling var. corallinus, the chromosomes showed 14II and 10I (figure 3 
and table 2). In a few cases trivalent chromosome associations were ob- 
served which suggest the presence of at least a few residual homologies 
between some of the additional ten chromosomes of MV. tilingiu var. coral- 
linus and the basic genome. The extra ten chromosomes constitute a sec- 
ond genome which appears to be incomplete on the basis of the other 
known chromosome numbers in the group (table 1). Possibly it is a highly 
modified derivative of the basic genome. However, its origin and relation- 
ships have yet to be determined precisely. Mimulus tilingu var. coral- 
linus warrants specific rank, but its accurate designation must also, as 
with the members of the M. guttatus complex, await an opportunity to 
study the literature and type specimens involved. 

Mimulus guttatus will hybridize with South American M. luteus 
(n=30, 31, or 32), but the hybrids are completely sterile. The chromo- 
somes in the pollen mother cells of the hybrids show considerable pairing 
(figure 3 and table 2). In some cases the number of pairs exceeds that of 
the basic genome, which must mean that M. luteus chromosomes are, at 
least occasionally, pairing with each other. However, inasmuch as there 
was no indication of autosyndesis in M. luteus itself (Mukherjee and 
Vickery, 1960), probably most of the paired chromosomes are homologues 
coming from M. guttatus or M. tilingit on the one hand and from M. luteus 
on the other. Therefore the basic genome appears to be present in M. 
luteus though in slightly modified form. The second genome of M. luteus 
may be a drastically modified form of the basic genome, but its true origin 
and relationship is not clearly demonstrated by the available data. 

Mimulus guttatus formed nearly sterile hybrids with /. glabratus var. 
utahensis (n=14). Typically the chromosomes of the pollen mother cells 
of these hybrids exhibited 13 bivalent and 2 univalent chromosome con- 
figurations at the first metaphase stage of meiosis (figure 4 and table 2). 
Apparently the two genomes are essentially homologous, but one pair of 
chromosomes has become so modified as to synapse only rarely. There- 
fore, in view of the sterility of the F, hybrids and the slight cytological 
differentiation of these species and despite the morphological similarity, 
M. glabratus var. utahensis should not be included in the M. guttatus 
complex of species. It is an integral part of the large, widespread, and 
varied M. glabratus complex. 


1962 | MUKHERJEE & VICKERY: MIMULUS 147 


5017, M. guttatus, n= 14. 5967, M. tilingii var. 5689, M. tilingii var. 
tilingii, n = 15. tilingii, n = 14. 
xX 
x xX 
5012, M. tilingii var. 
tilingii, mn = 14. 5052, M. guttatus, n= 14. 5012, M. tilingii var. 
tilingii, n = 14, 


@ 14, 101 _) 1411, 18 I 
e @ 
@ 141], 101 
e 
&e e 6 ° as e A 
\ Ay Lugs f Ale te ALN 
ais ° °° hg 
Te ya% a” ‘ - 
" C6 ” Pp’ 
e@ eo 
oe 
® ® 
5011, M. tilingii var. 5011, M. tilingii var. 5052, M. guttatus, n= 14, 
corallinus, n = 24. corallinus, n = 24. 
xX 
x x 
5043, M. luteus, n = 32, 
5052, M. guttatus, n= 14. 5007, M. guttatus, n= 14, 
. 
® 
WS e 
ay 
we ° 
A \) 17, 107 
a= ef e bee ENR [So CSET fi 
PY) e ie) 10 2.0 micra 


5012, M. tilingii, n= 14. 
x 


5043, M. luteus, n = 32. 


Fic. 3. Meiotic chromosomes of interspecific Fy hybrids of Mimulus guttatus com- 
plex with M. tilingit and M. luteus complexes, etc. All configurations at or near first 
metaphase. Camera lucida drawings at an original magnification of < 2,520, reduced 
Lou 1.200; 


The pollen mother cells of the F; hybrids of M. tilingi var. tilingu < 
M. glabratus var. utahensis (5012 * 5747) frequently exhibited a small 
extra chromosome and hence were n=15. The extra chromosome was 
probably a B chromosome from culture 5747, because it was not observed 


148 MADRONO [Vol. 16 


in culture 5012 (culture 5747 has yet to be studied cytologically). Fur- 
thermore, both parental forms are known to contain other populations 
with n=15 chromosomes (Mukherjee, Wiens, and Vickery, 1957; Muk- 
herjee and Vickery, 1959). 

As with the preceding variety, M. guttatus formed nearly completely 
sterile F,; hybrids with M. glabratus var. fremonti (n=30, 31). However, 
the pollen mother cells of these hybrids displayed much variation in the 
pairing behavior of their chromosomes (table 2). They showed the least 
amount of consistent pairing of any of the hybrids studied. They averaged 


TABLE 1. ORIGIN OF CULTURES USED IN THE CYTOGENETIC INVESTIGATION OF THE 
RELATIONSHIP OF MIMULUS GUTTATUS AND ITS SPECIES 


Species, culture, and 

chromosome number Origin and Collector 
M. guttatus DC. 

(M. guttatus DC. subsp. guttatus) 


5001, n=14 Pacific Grove, Monterey County, California, 
altitude 5 feet, Vickery 1 (UT). 

5004, n=14 Chew’s Ridge, Monterey County, California, 
altitude 4,500 feet, Vickery 3 (UT). 

5015, n=-14 Mono Inn, Mono County, California, alti- 
tude 6,450 feet, Clausen 2043 (UT). 

5052, n==14 Mt. Diablo, Contra Costa County, California, 


altitude 1,000 feet, Stebbins 703 (UT). 
(M. guttatus subsp. litoralis Pennell) 
5003, n=14 Pescadero, San Mateo County, California, 
altitude 30 feet, Clausen 2083 (UT). 
(M. guttatus var. puberulus [Greene] Grant) 
5006, n=14 Yosemite Junction (rocky creek), Tuolumne 
County, California, altitude 1,300 feet, 
Hiesey 560 (UT). 
5009, n=14--F 1 or 2 Mather (Hog Ranch meadow), Tuolumne 
County, California, altitude 4,600 feet, 
Hiesey 571 (UT). 


5014, n=14 Lee Vining Canyon, Mono County, California, 
altitude 8,000 feet, Clausen 2039 (UT). 

Saf hSyoume) taxa U) Stanislaus River, Tuolumne County, California, 
altitude and collector uncertain. 

5834, n=14 Salt Lake City, Salt Lake County, Utah, 
altitude 4,400 feet, Vickery 330 (UT). 

5835, n=14 Centerville, Davis County, Utah, altitude 4,360 
feet, Vickery 331 (UT). 

5S6 i Ma=14 Fish Haven, Bear Lake County, Idaho, alti- 
tude 6,100 feet, Vickery 322 (UT). 

5839, n=14 Big Cottonwood Canyon, Salt Lake County, 
Utah, altitude 7,100 feet, Vickery 334 (UT). 

5864, n=14 Skaggs Springs, Sonoma County, California, 


altitude ca. 50 feet, R. W. Holm, 
Spring 1951, unmounted. 


1962 | MUKHERJEE & VICKERY: MIMULUS 149 


Species, culture, and 
chromosome number Origin and Collector 


(M. lyratus Bentham) 
5007, n=14 Yosemite Junction (marsh), Tuolumne County, 
California, altitude 1,350 feet, 
Hiesey 559 (UT). 
(Af. laxus Pennell) 
SOV; ma 14725 Teor 2 Mather (Hog Ranch spring area), Tuolumne 
County, California, altitude 4,800 feet, 
Hiesey 569 (UT). 
(M. cordatus Greene) 


5017, n=14 Darwin Falls, Inyo County, California, altitude 
2,500 feet, Alexander & Kellogg 2844 (UC). 


(M. arvensis Greene) 


5346, n=14 Mount Oso, Stanislaus County, California, alti- 
tude 1,000 feet, Vickery 190 (UT). 


M. laciniatus Gray 


5064, n=—14 The Dardanelles, Tuolumne County, California, 
altitude 5,775 feet, Alexander & Kellogg 3746 
(UC): 

5630 hmne= 4. Lake Eleanor Road, Tuolumne County, Cali- 


fornia, altitude 4,200 feet, Vickery 179 (UT). 
M. glaucescens Greene 


50534 n==14 Richardson Springs, Butte County, California, 
altitude 600 feet, Pennell & Heller 25,667 
(UT). 


M. platycalyx Pennell 


5/52. 15 Crystal Lakes Reservoir, San Mateo County, 
California, altitude 800 feet, G. T. Ober- 
lander, April 1951 (UT). 


M. nasutus Greene 


5044, n=14 Hastings Reservation, Monterey County, Cali- 
fornia, altitude 1,500 feet, Stebbins 701 (UT). 

562 /e mo—13 West of Yosemite Junction, Tuolumne County, 
California, altitude 475 feet, Vickery 168 
(UT). 

Al. tilingii Regel var. tilingi 

5012, n==14 Slate Creek, Mono County, California, altitude 
10,000 feet, Clausen 2075 (UT). 

5689, n=14 Dana Plateau, Mono County, California, alti- 
tude 11,300 feet, C.W.Sharsmith, Aug. 21, 
1950. 

5690, n=14 Budd Lake, Tuolumne County, California, 


altitude 10,250 feet, C. W. Sharsmith, 
Sept. 13, 1950. 


5960/7,.n—=1L5 Mount Timpanogos, Utah County, Utah, alti- 
tude 7,800 feet, Del Wiens, Aug. 6, 1956 
(UT). 
M. tilingit var. corallinus (Greene) Grant 
5011, n==25 Porcupine Flat, Tuolumne County, California, 


altitude 8,000 feet, Hzesey 576 (UT). 


150 MADRONO [Vol. 16 


Species, culture, and 


chromosome number Origin and Collector 
M. luteus L. 
5042, n==32 Ilapel, Coquimbo, Chile, altitude 6,200 feet, 


U.S.D.A. Plant Introduction number 
144,535 (UT). 
5043, n=30 + 0, 1 or 2 Illapel, Coquimbo, Chile, altitude 2,000 feet, 
U.S.D.A. Plant Introduction number 
144,536 (UT). 
M. glabratus var. utahensis Pennell 


5048, n=14 Mono Lake, Mono County, California, alti- 
tude 6,440 feet, Stebbins 714 (UT). 
5747, n=14+ 0, or 1B Pilot Cone, Mineral County, Nevada, altitude 
chromosome* 5,550 feet, J. Figg-Hoblein, July 4, 1950. 
M. glabratus var. fremontii (Bentham) Grant 
5063, n=30 Black Meadow, Black Metal Wash, Whipple 


Mountains, San Bernardino County, Cali- 
fornia, altitude ca. 1,200 feet, ? collector 
(UC). 
5373, -n==30, (31*) Kakernot Springs, Alpine Creek, Brewster 
County, Texas, Cory 53,186 (UT). 
M. glabratus var. parviflorus (Lindley) Grant 
5041, n=45 Illapel, Coquimbo, Chile, altitude 4,000 feet, 
U.S.D.A. Plant Introduction number 
144,534 (UT). 
M. pilosiusculus HBK. 
5320, n=46 Botanic Garden, Copenhagen, Denmark 
(Wild in Argentina, Chile, and Peru). 
U.S.D.A. Plant Introduction number 
181,130 (UT). 


* Chromosome number based on counts in Fy hybrids involving this culture 
(see table 2). 


about 9 pairs per cell. Probably M. glabratus var. fremonti contains the 
basic genome, but in definitely modified form. 

Mimulus guttatus forms nearly sterile hybrids with M. glabratus var. 
parviflorus (n=45) and its closely allied species M. pilosiusculus 
(n=46). The chromosomes of the pollen mother cells of these F, hy- 
brids exhibited essentially regular pairing of 14II, 30I and 14II and 31], 
respectively. These forms contain the basic genome plus two additional 
genomes. One of the additional genomes is probably homologous to the 
second genome of M. glabratus var. fremonti as shown by three some- 
what ambiguous counts (see table 2). This hybrid, M. glabratus var. 
parviflorus (5041) & M. glabratus var. fremonti (5373), was hard to 


EXPLANATION OF FIGURE 4 
Meiotic chromosomes of interspecific F; hybrids of Mimulus guttatus complex 
with the M. glabratus complex, etc. All configurations at or near first metaphase. 
Camera lucida drawings at an original magnification of 2,520, reduced to X 1,260. 


1962 } 


5004, M. guttatus, n= 14. 
Xx 


5747, M. glabratus var 


utahensis, n = 14. 


5346, M. guttatus, n 


Xx 


5063, M. glabratus var. 
fremontii, n = 30. 


2 Ill, 13 II, 271 


5041, M. glabratus var, 
parviflorus, n = 45. 


x 


5339, M. laciniatus, 
n= 14, 


Fic. 4. Meiotic chromosomes, Mimulus guttatus complex and relatives, F; hybrids. 


5052, M. guttatus, n = 


MUKHERJEE & VICKERY: MIMULUS 


5010, M. guttatus, n= 14, 5747, M. glabratus var. 


utahensis, ne» 1470, 1 
x 
x 
5048, M. glabratus var. 
utahensis, n = 14. 5012, M. tilingii var. 
tilingii, n= 14 
Ps 16 Il, ~" 
°"p 131 14, 311° @ -“» 
. e 
.) ee@ 
~ a m ey 
e 
e > 7 & > ¢ 
. + $ Ly ~ 
ro 04 @. Laan y eo” 
C] 
e 4e,° 
& e e 
@ 
5015, M. guttatus, n= 14. 5041, M. glabratus var. 
parviflorus, n = 45. 
x 
x 
5373, M. glabratus var. 


fremontii, n = 30, 31. 


5010, M. guttatus, n= 14. 


5320, M. pilosiusculus, 
n = 46. 


5041, M. glabratus var. 
parviflorus, n = 45 

Xx Xx 

= 14. 5373, M. glabratus var. 
fremontii, n = 30, 31 


eR (SEES ATT | 


0 10 20 micra 


152 MADRONO [Vol. 16 


make and even harder to analyze cytologically. The chromosome num- 
bers are too low for it to be a spontaneous autododecaploid instead of the 
true hybrid which it appeared to be on morphological grounds. We do not 
know how to explain the extra chromosomes, but the large number of pairs 
suggests to us that M. glabratus var. fremontu and M. glabratus var. 
parviflorus have two genomes in common. The affinities of the third gen- 
ome in the South American form are not apparent from the data at hand. 

The basic M. guttatus genome appears to be little modified in these 
South American forms, whereas it was slightly modified in M. glabratus 
var. utahensis from the Great Basin and greatly modified in M. glabratus 
var. fremontiu from the southwestern United States. These North and 
South American entities of the W/. glabratus complex probably are not as 
closely related as their current taxonomic status suggests (Grant, 1924; 
Fassett, 1939; Pennell, 1947). 

In conclusion, despite the low number of pollen mother cells analyzed, 
the basic or M. guttatus genome of 14 chromosomes appears to be present 


TABLE 2. PAIRING BEHAVIOR OF MEIOTIC CHROMOSOMES IN Fy AND A FEW Fo HYBRIDS OF 
MIMULUS GUTTATUS AND ITS RELATIVES. 


Number of PMC’s 
Combinations of parental Culture numbers examined and 
species and varieties of the parents pairing behavior 
F, HYBRIDS 
guttatus « guttatus 5001 * 5003 1-14II 
na 34 na=14 5001 « 5004 4-14II 
5001 « 5006 12-14II* 
5001 x 5007 2-1411; 1-13], 21 
5001 x 5009 2-141If 
5001 * 5010 5-14IIt 
5001 « 5052 1-14II 
5001 « 5346 3-14II 
5001 <5 753 4-14II 
5001 X 5834 1-14II; 1-13II, 21 
1-12II,, 41 
1-11II, 61 
5003 X 5839 3-14II 
5004 x 5006 8-14I1* 
5004 x 5010 1-141If 
5006 & 5834 3-14I1* 
5009 & 5010 1-14IIf 
5014 x 5834 2-14II1 
5052 & 5006 10-14II* 
5052 & 5837 1-14II 
5753 < 5001 7-14II 
5835 & 5834 3-14I1 
guttatus < laciniatus 5017 « 5064 3-141I 
n=—14 n=14 5017 >< $339 2-14II1 
5052 * 5339 4-14IT 


5064 x 5017 2-1411 


1962 | MUKHERJEE & VICKERY: MIMULUS 


Combinations of parental 
species and varieties 


Culture numbers 
of the parents 


Number of PMC’s 
examined and 
pairing behavior 


guttatus < glaucescens 
n=14 n=—14 


guttatus « platycalyx 
n=—14 n=—=15 


guttatus x nasutus 


n=—14 n=14 
guttatus K nasutus 
n—14 n=13 


guttatus X< tillingii var. tilingii 
n= ne 14 


guttatus < tilingi var. corallinus 
n=—14 n=24 


guttatus < luteus 
n=—14 n=30,31,32 


guttatus < glabratus var. utahensis 
n—14 n=14 


guttatus « glabratus var. fremontii 
n=14 n=30, 31 


laciniatus X nasutus 


n=14 n——13 
glaucescens < platycalyx 
n=14 15 
tilingit var. tilingit « guttatus 
n=—15 n=—14 
tilingit var. tilingit * tilingit var. 
n=14 tilingit 
n=14 


luteus X tilingii var. tilingii 
N== 505/31, 32 n=14 


luteus X tilingii var. tilingit 
N==32Z n==14 


5014 & 5653 


5017 & 5653 
5837 < 5653 


S017 K 57/52 
5752 X 5010 


5017 5044 


5017 < 5327 
5327 X 5003 


S012, 5052 
5017 X 5012 


5010 X 5011 
5011 5007 
SOIL XK S052 


5017 « 5043 
5052 X 5043 
5004 < 5747 
5010 & 5048 


5017 X 5747 
5837 X S747 


5014 X 5373 
S015y x 53573 


5346 & 5063 
9339 X 5327 
5603 KX S702 
5967 5052 


5689 < 5012 
5690 X 5012 


5043 x 5012 


5042 « 5690 


1-12II, 41 
2= 1168 
4-1411 
4-14II, 1-11I1I, 61 
S-14ly 21 
1-141; 1-13II, 21; 
1-121], 41+ 
3-141 


2-13II, 11 

2-13II, 11 

1-3II, 221 

1-14II 

1-14II, 10It 

7-1411,101 

1—4III, 101], 61; 
2-3III, 12II, 51; 
1-141], 101 

1-1III, 1111, 191; 
1-10II, 251 

1-16II, 121; 
1-15II, 141; 
3-141], 181 

2-141I 


11-131, 21f 


S311, 21 
4-14II 
1-441 
1-16II, 131; 
1-9II, 261 
1-151I, 141; 
1-7II, 301 
SIGN 11 


6-14II, 11 
2-14II, 11 


3-14I1 
1-14I1 


1-17II, 101; 
2-15II, 141; 
1-14II, 161 

2-141I, 181 


153 


154 MADRONO [Vol. 16 


Number of PMC’s 


Combinations of parental Culture numbers examined and 
species and varieties of the parents pairing behavior 

glabratus var. utahensis 5747 xX 5012 7-14II, 11; 

tilingi var. tilingii 1-14II 

n=14+0,1 n=14 

glabratus var. parviflorus X guttatus 5041 « 5010 3-14II, 311f 
n—=45 n=—14 

glabratus var. parviflorus < laciniatus 5041 5339 1-21T], 131], 271; 
n=45 n=14 1-14], 311 

glabratus var. parviflorus & glabratus 5041 x 5373 1-24II, 311; 
m==45 var. fremonti 1-30II, 231 

n=—30, 31 (and one M,, cell 


containing config- 
uration of 31 and 


40-++-chromosomes) 

pilosiusculus < guttatus 5320 & 5052 2-14II, 321 

n—46 n—14 5320 5864 2-15II, 301* 
Fo HYBRIDS 
guttatus x guttatus 5346 & 5839 7-14I], iI 

n—14 n—14 
guttatus X laciniatus 5052 & 5339 1-14II; 2-13], 21; 

n=—14 n=14 1-12II, 41; 

1-9II, 101 

nasutus XK nasutus 5751 < 5327 1-12I11,301 


n—14 n=13 


* Culture 5006 and 5864 and their hybrids were subject to chromosome stickiness 
due to too slow fixation. 
t In culture 5009 and 5010, n=14 + 1 or 2. 


in all 18 species and varieties of the M. guttatus complex and its rela- 
tives in section Simiolus studied in this investigation. In several cases, 
e.g., M. nasutus, M. platycalyx, M. tilingi, the basic genome has been 
changed in number by aneuploidy. In other cases, e.g., M. glaucescens, 
M. luteus, M. glabratus var. utahensis and particularly in M. glabratus 
var. fremontii, it has been modified by mutations, as indicated by a de- 
crease in the regularity of chromosome pairing in the F, hybrids. The 
second genome of MW. glabratus var. fremonti (n=30, 31) appears to be 
homologous to the second genome of M. glabratus var. parviflorus (n= 
45), but its further relationships are not known. The homologies of the 
additional genomes present in the various tetraploid and hexaploid species 
have yet to be fully determined. 


Department of Obstetrics and Gynecology 
Columbia University, New York City, N.Y. 


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


1962 | MASON: BAKER CIS) 


LITERATURE CITED 

CarRLQuisT, S. 1953. In Documented chromosome numbers of plants. Madrono 
12:31, 

Fassett, N.C. 1939. Notes from the herbarium of the University of Wisconsin— 
XVIII. Rhodora 41:524:529. 

GoopsPEED, T. H. 1954. The genus Nicotiana. Chronica Botanica Co., Waltham, 
Mass. 536 pp. 

Grant, A.L. 1924. A monograph of the genus Mimulus. Ann. Missouri Bot. Gard. 
11:99-389. 

MUKHERJEE, B. B. and R. K. VICKERY, JR. 1959. Chromosome counts in the section 
Simiolus of the genus Mimulus (Scrophulariaceae). IIJ. Madrono 15:57-62. 

. 1960. Chromosome counts in the section Simiolus of the genus Mimulus 
(Scrophulariaceae). IV. Madrono 15:239-245. 

MUKHERJEE, B. B., D. Wiens, and R. K. VicKErRy, JR. 1957. Chromosome counts 
in the section Simiolus of the genus Mimulus (Scrophulariaceae). IJ. Madrono 
14:128-131. 

PENNELL, F.W. 1947. Some hitherto undescribed Scrophulariaceae of the Pacific 
states. Proc. Acad. Phil. 99:155-199. 

. 1951. in Illustrated flora of the Pacific states by Leroy Abrams. Stanford 
Univ. Press. Vol. III, pp. 688-731. 

Vickery, R. K., Jr. 1955. Chromosome counts in the section Simiolus of the genus 
Mimulus (Scrophulariaceae). Madrono 13:107-110. 

. 1956a. Data on interracial hybridizations in Mimulus guttatus (Scrophu- 

lariaceae). Proc. Utah Acad. 33:37-43. 

—. 1956b. Data on interracial and interspecific hybridizations in the section 

Simiolus of the genus Mimulus (Scrophulariaceae). Proc. Utah Acad. 33:45-64. 
. 1959. Barriers to gene exchange within Mimulus guttatus (Scrophularia- 

ceae). Evolution 13:300—-310. 


MILO S. BAKER (1868-1961) 


On January 4, 1961, the career of Milo S. Baker came to an end in his 
92nd year. His was a role that closes the second dynasty of California 
botanists, namely those botanists who were direct career descendants of 
the colorful pioneers, many of whom he knew personally. His career as 
a plant collector of the California flora opened with the close of the last 
century and continued well over half of the current century, for he was 
very active to the end. 

Born in Strawberry Point in Iowa on July 19, 1868, he came to Cali- 
fornia with his parents in 1875 to settle in Oak Run, Tehama County. At 
the age of twelve he was taken to San Jose, where he completed high 
school and entered what was then San Jose Normal School. At the end 
of one year he was admitted by examination to the teaching profession 
in the public schools of Santa Clara County. In 1887 he went to Modoc 
County to teach in the elementary schools. To reach his school, he walked 
from Redding to Bieber, a distance of almost 100 miles. He collected 
plants in this general area, and corresponded about them with Pro- 


156 MADRONO [ Vol. 16 


fessor E. L. Greene of the University of California. Much of the flora 
of eastern Shasta and Lassen counties was first made known through his 
work. Noteworthy is Cupressus Bakeri, discovered by him in the lava beds 
and named in his honor by Professor Willis Linn Jepson in the first 
volume of the Flora of California. In 1894, Milo Baker and F. P. Nutting, 
a like-minded field botanist, spent six weeks collecting together in Lassen 
and Modoc counties. Their collections were widely distributed in herbaria. 
At the close of the century he came to the University of California, where 
he majored in chemistry and took courses in botany. On completion of 
his work he taught at Lowell High School in San Francisco from 1901 
to 1906. Somewhere in his travels he contracted malaria and decided to 
leave the teaching profession in the interests of his health. He purchased 
a ranch in Kenwood, Sonoma County, which he named “The Maples,” 
a name which appears on some of his collections; it is situated at the 
entrance of Adobe Canyon, where Sonoma Creek enters Sonoma Valley. 
I suspect he was led to this particular ranch because of its botanical 
assets rather than its agricultural promise. Its rocks are of basalt, Sonoma 
tuff, and serpentine, and they selected their flora accordingly. His career 
as a rancher was not a great success, and it is not surprising that in 1922 
he began teaching in Santa Rosa High School and later in Santa Rosa 
Junior College, where he had a distinguished career. Few high school or 
junior college teachers of botany have inspired so many students to 
follow plant science in some form as a career. Few teachers have aroused 
a greater interest in botany among the laymen of their community. 
Under his generalship, the annual wild flower shows in Sonoma County 
attracted the attention of all of central California. They were outstanding 
in their representation of the flora as well as in the inspired participation 
of the community in making them a success. As a result Santa Rosa is 
outstanding in its botanically informed populace. 

His research interests were twofold. First and foremost was the genus 
Viola, upon which he published several dissertations and which he studied 
through a living collection at his home. Second was his interest in the 
flora of the North Coast Ranges of California. He published an annotated 
list of the plants of this area in mimeograph form and kept it up-to-date 
in several editions. 

Baker was a man whose scientific ambition was always afire. Although 
a robust man, in his later years his ambition far exceeded his physical 
capacity. He literally refused to accept old age and at the age of 91 he 
spoke frequently of his plan to collect violets on Mt. McKinley in Alaska 
and sought companions to accompany him. He was not easily dissuaded. 

He built an excellent herbarium of the North Coast Range counties. It 
is now fittingly housed at Santa Rosa Junior College and stands as a 
monument to his inspired teaching. His collection of violets is now housed 
at the University of California at Berkeley and is a marvelous research 
collection of this group of plants. 


1962] MASON: BAKER 157 


A photograph of Milo S. Baker and a dedication to him form the 
frontispiece of Volume XIII of MaproNo. 

I wish to express my appreciation to Mrs. Avis Stopple, Librarian at 
Santa Rosa Junior College, for assistance in the preparation of the an- 
notated bibliography which follows—HeERBErT L. Mason, Department 
of Botany, University of California, Berkeley. 


PUBLISHED WRITINGS OF Mito S. BAKER 


1929. Field notes on certain Brodiaea species in Humboldt County. Madrofio 1: 
199-200. 

1932. A new species of Arctostaphylos. Leafl. West. Bot. 1:31-32. 

——. A partial list of seed plants and a few spore plants growing spontaneously in 
Sonoma, Napa, Lake, and Mendocino counties. Santa Rosa Jun. Coll. [undated, 
received at Univ. Calif. May 8, 1932; mimeographed; arranged alphabetically 
by genera]. 

1934. Pitkin Marsh, a floral island at Vine Hill, Sonoma County. Leafl. West. Bot. 1: 
103. 

-——. A partial list of seed plants of the North Bay counties. Santa Rosa Jun. Coll., 
December 9, 1934 [mimeographed; revised list containing corrections and addi- 
tions to the 1932 list; arranged alphabetically by genera]. 

1935. Studies in western violets—I. Madrofio 3:51-57. 

1936. Studies in western violets—II. Madronio 3:232-239. 

1937. List A. A partial list of seed plants of the North Bay counties. Santa Rosa 
Jun. Coll., March 20, 1937 [mimeographed; revised list containing corrections 
and additions to the 1934 list; arranged alphabetically by genera]. 

——. List B. A partial list of seed plants of the North Bay counties arranged alpha- 
betically under their families. Santa Rosa Jun. Coll. [undated, received at Univ. 
Calif. 1937; mimeographed; the same as 1937 List A except for arrangement]. 

1938. An undescribed species of Viola from Utah. Madronio 4:194-196. 

1940. Supplementary list of seed plants of the North Bay counties. Santa Rosa Jun. 
Coll. [undated, received at Univ. Calif. January 11, 1940; mimeographed; sup- 
plement to 1937 List A]. 

——. Studies in western violets—III. Madrono 5:218-231. 

1941. List A. A partial list of seed plants of the North Bay counties of California 
arranged alphabetically under their botanical names. Santa Rosa Jun. Coll., 
May, 1941 [mimeographed; revised list containing corrections and additions to 
the 1937 List A]. 

——. List B. A partial list of seed plants of the North Bay counties of California 
arranged alphabetically under their families. Santa Rosa Jun. Coll., May, 1941 
[mimeographed; revised list containing corrections and additions to the 1937 
List B]. 

1943. 1943 supplement to the partial list of seed plants (1941) of the North Bay 
counties of California. Santa Rosa Jun. Coll. [mimeographed]. 

1945. A partial list of seed plants of the North Bay counties of California. Santa 
Rosa Jun. Coll., May, 1945 [mimeographed; arranged alphabetically by genera; 
revision of 1941 List A]. 

1946. Supplement to the 1945 check list of the seed plants of the North Bay counties. 
Santa Rosa Jun. Coll., August, 1946 [mimeographed ]. 

1947. A partial list of seed plants of the North Bay counties of California arranged 
alphabetically under their families. Santa Rosa Jun. Coll., May, 1947 [mimeo- 
graphed; revised list containing corrections and additions to the 1945 list]. 

—. A new violet from Mexico. Madrono 9:131-133. 

1948. A new western violet. Leafl. West. Bot. 5:101-102. 


158 MADRONO LVol. 16 


1949. Studies in western violets—IV. Leafl. West. Bot. 5:141-147. 

——. Studies in western violets—V. Leafl. West. Bot. 5:173-177. 

——. Studies in western violets—VI. Madrofio 10:110—-128. 

1951. A partial list of seed plants of the North Coast counties of California arranged 
alphabetically under their families. Santa Rosa Jun. Coll., June, 1951 [mimeo- 
graphed]. 

1953. Studies in western violets—VII. Madronio 12:8-18. 

——. A correction in the status of Viola macloskeyi. Madrono 12:60. 

1954. A partial list of seed plants of the North Coast Ranges by the North Coast 
Herbarium. Santa Rosa Jun. Coll., June, 1954 [mimeographed; arranged alpha- 
betically by families]. 

1957. Studies in western violets—VIII. The Nuttal[llianae continued. Brittonia 
9:217-230. 

1958. Supplement to 1954 partial list of seed plants of North Coast Ranges of Cali- 
fornia by the North Coast Herbarium. Santa Rosa Jun. Coll., 1958 [mimeo- 
graphed]. 

1960. Studies in western violets—IX. Miscellaneous species in the sections Nomimi- 
um and Chamaemelanium. Madronio 15:199-204. 


CYTOLOGICAL OBSERVATIONS ON ADIANTUM xX TRACYI 
C. C. HALL? 


WarrEN H. WAGNER, JR. 


The California maidenhair fern, Adiantum jordani C. Muell. (syn. 
A, emarginatum D. C. Eaton) is one of the endemic pteridophytes of the 
California Floral Province (Howell 1960). In the North Coast Ranges 
where it comes into association with the wide-ranging A. pedatum L., 
there has occasionally been found an intermediate plant, A. x tracy 
C. C. Hall, which combines the characteristics of these sharply different 
species (Wagner 1956). A single plant of the intermediate fern was dis- 
covered as early as 1895 along the Eel River near Pepperwood, Humboldt 
County, by Mr. J. P. Tracy, and the observations to be recorded here 
are based on a propagated descendant of that plant. Other naturally 
occurring specimens of A. * tracyi have been found in Sonoma and 
Marin counties. Easily propagated from rhizomes, this fern has proved 
a decorative and hardy garden plant. 

Adiantum  tracyi has been interpreted as an interspecific hybrid 
because of its morphological intermediacy in a number of obvious fea- 
tures; its sporadic distribution, and occurrence where the putative par- 
ents grow nearby; and the irregularity of its spores (Wagner, ibid.). The 
facts to be reported below tend to supply additional evidence for con- 
sidering that this fern is a natural hybrid. To obtain cytological observa- 
tions, the immature sori of Adiantum < tracyi were fixed in Newcomer’s 
Fixing Fluid (Newcomer, 1953). Collections were made in May, June, 
and July 1960 from plants growing at the University of Michigan Botani- 
cal Gardens. 


1 Research in connection with National Science Foundation Grant G10846. 


1962 | WAGNER: ADIANTUM 159 


N\\ w 


Fic. 1. Adiantum xX tracyi: A, meiotic metaphase showing 59 univalents; B, 
sporocytes extruded from a single sporangium and squashed, showing different stages 
and unassimilated chromosomes; and C, sporangium forced open with alcohol and 
diaphane, showing part of spore complement; note small wrinkled spores and large 
smooth spores. (Camera lucida drawings, based on material obtained from descendant 
of the original plant.) 


160 MADRONO [Vol. 16 


The proper stages of meiotic division were found by selecting pinnules 
which had reached full size and upon which the sori were of approxi- 
mately mature dimensions but pale greenish-white in color. Because of 
the leathery false indusium it was difficult to scrape out the young spo- 
rangia. The entire indusium was removed, therefore, placed on a micro- 
scope slide in acetocarmine stain, and broken apart, after heating, by 
tapping briskly with the point of a dissecting needle on the cover-slip. 
This broke apart the false indusium and the sporangia (and sometimes, 
unfortunately, the cover-slip), but the spore mother cells became suf- 
ficiently separated so that they could be properly squashed and studied 
under the microscope. Camera lucida drawings were made of good prep- 
arations, and the slides were made permanent. 

The observations of meiosis were interesting for two reasons. First, 
there was no pairing at all between the chromosomes. At metaphase, the 
chromosomes become very short and oblong in outline, and in not one of 
the numerous figures observed were there any indications of pairing. 
This fact suggests that there is a lack of homology between the two 
genomes that make up the chromosome complement, and that they very 
likely came from different species. 

The second interesting observation was that the number of chromo- 
somes is 59. Such a number seems at first unusual for a presumably di- 
ploid plant, and it suggests at least two explanations: either the plant is 
one in which there has been the loss or addition of a chromosome, or it 
is a hybrid between parents with different chromosome numbers, one of 
them with an odd number and the other with an even number. 

The distribution of chromosomes at metaphase is irregular, and tetrad 
formation is characterized by three to five daughter nuclei plus a varying 
number of excluded chromosomes, as shown in figure 1, B. There is some 
lack of synchrony in the meiotic divisions of the sixteen spore mother 
cells, so that at one time it is possible to find several different stages in 
tetrad formation in the same sporangium. This is unlike the situation 
ordinarily observed in normal leptosporangiate fern species, where 
sporogenesis proceeds approximately simultaneously in all sixteen spore 
mother cells of a sporangium. As would be expected and as was reported 
earlier (Wagner, 1956), the spores are abortive and irregular. Even at 
an early stage in the spore maturation their irregularity is evident. Many 
of them are small and become corrugated or folded, and in the same 
sporangia others are very large and smooth. Figure 1, C, shows a mature 
sporangium forced open by alcohol and diaphane and with part of the 
variable spore complement present. 

That Adiantum X tracyi may actually have arisen from parents with 
different chromosome numbers is suggested by the fact that in four 
genera of Adiantaceae (Adiantum, Cheilanthes, Aleuritopteris, and 
Saffordia) two numbers are known, viz. n = 29 and n = 30, among the 
species of each. The other adiantaceous genera, so far as is known at 


1962 | STONE: PLATYDESMA AND MELICOPE 161 


present, have only single numbers among their species, the number of 
each genus being either n = 29 or n = 30 (Manton 1959). Adiantum 
pedatum, which is one of the presumed parents of A. X tracyi, has been 
observed in material from two regions (Vancouver: Manton 1959; and 
Ontario: Britton 1953) to have n = 29. If it can be assumed that this 
number is characteristic of A. pedatum everywhere, then we may sug- 
gest that the other presumed parent, the endemic Californian A. jordani, 
which has not yet been examined cytologically, will probably prove to 


have n = 30 chromosomes. 
Department of Botany 
University of Michigan 
Ann Arbor, Michigan 


LITERATURE CITED 


Britton, D. M. 1953. Chromosome studies on ferns. Am. Jour. Bot. 40:575-583. 

Howe LL, JouN THomas. 1960. The endemic pteridophytes of the California Floral 
Province. Am. Fern Jour. 50 (no. 1):15-25. 

Manton, I. 1959. Chromosomes and fern phylogeny with special reference to “Pteri- 
daceae.” Jour. Linn. Soc. Bot. 56:73-91. 

Newcomer, E. H. 1953. A new cytological and histological fixing fluid. Science 
118:166. 

WAGNER, WARREN H., Jr. 1956. A natural hybrid, « Adiantum tracyi C. C. Hall. 
Madrono 13:195-205. 


TAXONOMIC AND NOMENCLATURAL NOTES ON 
PLATYDESMA (HAWAII) AND A NEW NAME FOR A 
MELICOPE (SOLOMON ISLANDS)! 


BENJAMIN C. STONE ” 


The genus Platydesma was proposed by Horace Mann, Jr. (1866) to 
include one species, P. campanulatum (—a), which had been collected by 
Mann and W. T. Brigham “on the mountains behind Honolulu.” A 
slightly expanded description is found in Mann (1869). Two species were 
added to the genus by Hillebrand (1888) in his “Flora of the Hawaiian 
Islands”: P. cornutum (—a), from the island of Oahu, and P. rostratum 
(—a), from the island of Kauai. Hillebrand (op. cit.) also transferred 
to Platydesma a species described by Asa Gray as Pelea auriculaefolia 
(1854, p. 343; 1857, pl. 36), but this was an error, as Rock (1913, 1918) 
has shown, for Gray’s original placement is correct. Although Hillebrand 


1 Studies in the Hawaiian Rutaceae, I. This paper is the first in a series of studies 
concerned primarily with the Hawaiian Rutaceae, of which the second and third 
papers are now in press. 

2 This work was carried out while the writer was Research Assistant, Botany 
Department, University of Hawaii, Honolulu. It is an outgrowth of studies for a 
monograph of the genus Platydesma, now in press. 


162 MADRONO 


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Fic. 1. Platydesma spathulatum (Gray) B. C. Stone. Holotype of Melicope spa- 
thulata Gray (US), collected by United States Exploring Expedition of 1838-42 on 
mountains of Kauai. (Photo courtesy U.S. Nat. Herb.) 


1962 | STONE: PLATYDESMA AND MELICOPE 163 


cited the type specimens of Pelea auriculaefolia Gray (United States Ex- 
ploring Expedition of 1838-1842, Mauna Kea, Hawaii), it was perhaps 
not seen by him. He apparently based his conclusion for the transfer of 
this species to Platydesma upon specimens collected by himself in the 
Kohala Mountains and on the island of Hawaii and also on a specimen 
collected by Reverend John Lydgate near Laupahoehoe, Hawaii. These 
Hillebrand and Lydgate specimens are in fact representatives of Platy- 
desma and not of Pelea, and may be referred to the distinct species Platy- 
desma Remyi (Sherff) Degener, Sherff, & Stone (Degener, 1961), to 
which P. campanulatum var. sessifolium (—a) Rock (1913) may also be 
referred. Rock’s description is typified by one of his collections (Rock 
4222). Platydesma Remyi is based on a collection in the Museum d’His- 
toire Naturelle of Paris collected by Jules Remy in 1853. 

Hector Léveillé (1911) described two species of Platydesma, P. oahu- 
ense (-1s) and P. Fauriei, but both names are later homonyms, as pointed 
out by Rock (1914). The first is referable to Mann’s original species; 
the second is not Platydesma and is not even rutaceous, but is referable 
to the solanaceous Nothocestrum longifolium Gray. 

The species which has been known as Platydesma campanulatum is 
relatively common in the Hawaiian rain-forest, and is met with much 
more frequently than are the other two species, P. cornutum and P. ros- 
tratum. It is also of wider distribution, at least as presently known, since 
P. cornutum is endemic to Oahu, while P. rostratum is found only on 
Kauai. It is always or nearly always accompanied by such characteristic 
rain-forest plants as species of Pelea, Fagara, Straussia, and Gouldia. It 
is reasonably well represented in herbaria (much better than the other 
two species), and thus it is rather unfortunate that the specific epithet 
must be changed. 

The name Platydesma campanulatum (—a) Mann was not the first for 
this species. Some years earlier, Asa Gray (1854) had described it as two 
different species, Melicope spathulata (p. 352) and M. grandifolia (p. 
354). This fact, suspected by Rock (1918) and later by Skottsberg (1936, 
although he still used the name Platydesma campanulatum in 1944), has 
not received the formal recognition it requires under the International 
Code of Botanical Nomenclature. It is necessary, therefore, to choose 
between Gray’s two specific names. The type specimens of both of Gray’s 
species are sterile or nearly sterile. The first, Melicope spathulata, is a 
specimen from Kauai which is in bud (fig. 1). The second, M. grandi- 
folia, is a specimen from Hawaii which is altogether sterile (fig. 2). 
Although the foliage in this second specimen is sufficient for specific 
placement, it is not entirely reliable for infraspecific placement; none- 
theless, this specimen certainly belongs to the same species as the first 
(M. spathulata). Because the type specimen of M. spathulata is at least 
not entirely sterile, being in bud, it seems preferable to use M. spathulata 
rather than M. grandifolia as the basionym in the following combination: 


164 MADRONO [Vol. 16 


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Fic. 2. Platydesma spathulatum (Gray) B. C. Stone. Holotype of Melicope? 
grandifolia A. Gray (US), collected by United States Exploring Expedition of 1838-42 
on Mauna Kea, Hawaii. (Photo courtesy U. S. Nat. Herb.) 


1962] STONE: PLATYDESMA AND MELICOPE 165 


Platydesma spathulatum (A. Gray) B. C. Stone, comb. nov.* Meli- 
cope spathulata A. Gray, Bot. U.S. Expl. Exped. 15:354. 1854. M.? 
grandifolia A. Gray, loc. cit. Platydesma campanulata H. Mann, Proc. 
Boston Soc. Nat. Hist. 10:317. 1866. Hillebrand, Fl. Haw. Ids. 71 (as 
companulata). 1888. Rock Indig. Trees Haw Ids. 241. 1913. Heller, Minn. 
Bot. Stud. 1(9):841. 1897. Skottsberg, Acta Horti Gothob. 10:120. 1935; 
15:388. 1944. P. campanulata var. macrophylla Hillebrand, F1. Haw. Ids. 
72. 1888. P. campanulatum f. coriaceum Rock. Indig. Trees Haw. Ids. 
243. 1913. P. oahuensis Léveillé in Fedde, Rep. Sp. Nov. 10:153. 1911. 

In addition to the above transfer of specific epithet, there are two 
varieties requiring transfer, as follows: 

Platydesma spathulatum var. pallidum (Hillebr.) B.C. Stone, comb. 
nov. P. campanulata var. pallida Hillebrand, op. cit. 

Platydesma spathulatum var. pubescens (Skottsb.) B.C. Stone, comb. 
nov. P. campanulata var. pubescens Skottsberg, Acta Horti Gothob. 
15:388. 1944. 

The type of the genus is now to be called Platydesma spathulatum, 
and P. campanulata becomes a synonym. However, the ultimate type of 
the genus is the type specimen of P. campanulata (Mann & Brigham 94, 
CU; isotypes, 94 or 94-bis, at K, BISH), rather than the type of Gray’s 
Melicope spathulata (US). 

A species from Bougainville, Solomon Islands, given the name Meli- 
cope grandifolia by B. L. Burtt in 1935, bears a later homonym since 
Gray’s M. grandifolia preémpts that epithet. It is thus necessary to pro- 
pose the following new name: 

Melicope Burttiana B. C. Stone, nom. nov. M. grandifolia B. L. Burtt, 
Kew Bull. 1935: 300, non A. Gray, 1854. 

Type. Solomon Islands: Bougainville, Waterhouse B.227 in 1930-31 
(US). 

College of Guam 
Agana, Guam 


LITERATURE CITED 


DEGENER, OTTO. 1961. Flora Hawaiiensis. Unpaged. Oahu, Hawaii. 

Gray, AsA. 1854. Botany. Phanerogamia. in U. S. Expl. Exped... . under . . . Charles 
Wilkes ... 1:1-777. 

————-, 1857. Atlas. Botany. Phanerogamia. in U. S. Expl. Exped. ... under. . 
Charles Wilkes ... 1: plates 1-100. 

HILLEBRAND, WILLIAM. 1888. Flora of the Hawaiian Islands. Carl Winter, Heidelberg ; 
B. Westermann & Co., New York. pp. 673. 

LEVEILLE, HEcTor. 1911. Platydesma oahuensis and P. Fauriei, in Fedde, Rep. Sp. 
Nov. 10:153-154. 

Mann, Horace, Jr. 1866. Revision of Schiedea and the Hawaiian Rutaceae. Proc. 
Boston Soc. Nat. Hist. 10:310-319. 


® Mann used the generic name Platydesma as if it were of feminine gender, and was 
followed in this by several later authors, but the name is one of several in Greek 
(such as Geniostoma) which, though ending in —a, takes a neuter modifier. 


166 MADRONO [Vol. 16 


. 1869. Alsinidendron, Platydesma, and Brighamia. Mem. Boston Soc. Nat. 
Hist. 1:529-531. 
Rock, JOSEPH F. 1913. The indigenous trees of the Hawaiian Islands. Honolulu. T.H., 
privately printed. pp. 518. 
. 1914. Revisio plantarum hawaiiensium a Léveillé descriptarum, in Fedde, 
Rep. Sp. Nov. 13:352-361. 
. 1918. Pelea and Platydesma. Bot. Gaz. 65:261-267. 
SKOTTSBERG, CARL. 1936. Vascular plants from the Hawaiian Islands. II. Acta Horti 
Gothob. 10:97-193. 
. 1944. Vascular plants from the Hawaiian Islands. IV. Acta Horti Gothob. 
¥52275=53 5. 


A NEW SPECIES OF GALIUM IN CALIFORNIA 
LAURAMAY T. DEMPSTER 1 


A completely new and radically different species of Galizum has been 
discovered by Mrs. Clare Hardham of Paso Robles. Mrs. Hardham, who 
has been making a study of the flora of the Santa Lucia Mountains, has 
found the new Galium in six separate localities, almost invariably asso- 
clated with Cupressus sargentii Jepson, which is a well-known indicator 
of serpentine soils. The new species is diploid (2n = 22), highly uniform, 
and almost certainly primitive and residual. 

Its nearest relative would seem to be Galium clementis Eastwood, 
which is another endemic occurring a little farther north on Cone and 
Junipero Serra (Santa Lucia) peaks. More distantly, the new species is 
certainly related to G. californicum H.&A., G.nuttallu Gray, G. bolanden 
Gray, G. sparsiflorum Wight, etc., all of which it resembles in its dioecism 
and in the possession of fleshy fruits. It differs sharply from all of those 
species, however, in having six leaves to a node instead of four. Galium 
clementis has generally four leaves to a node, but occasionally six, and 
stands uniquely, therefore, in an intermediate position between the new 
species, described herein as G. hardhamae, and all of our other berry- 
fruited species. Galium hardhamae differs additionally from G. clementis 
in having a less compact habit, fewer shorter hairs (being consequently 
of a much darker green), slightly smaller leaves and flowers, and in the 
succulence of its leaves. 

Galium hardhamae is a dioecious perennial growing usually under or 
near Cupressus sargentu, on humous serpentine soil. The plants are low 
and matted, rooting at the nodes, the flowering branches ascending or 
weakly clambering, the hispid internodes 1% to 2% cm. long, mostly much 
longer than the leaves. The leaves (invariably 6 to a node) are bright 
green, sparsely hispid, lanceolate, acute, and each tipped with a stout 
hair. When living, they are fleshy, round above and plane beneath. In the 


1 Contributions from the Jepson Herbarium, number 5. 


1962 | DEMPSTER: GALIUM 167 


Fic. 1. A, single flowering branch (staminate) x 34; B, cross section of leaf blade, 
x 15; C, single node with leaves, x 2%; D, leaf bases of a single whorl, x 15; 
E, staminate flower, < 10; F, pistillate flower, < 10; G, nearly ripe fruit, x 2%. 
The difference in the hairs of the two flowers illustrated is not a sexual, but an 
individual difference. 


desiccated condition, however, they curl downward laterally, owing to 
shrinkage of the spongy parenchyma. Apart from this lateral curling, the 
leaves are more reflexed or less, depending upon the condition of moisture, 
being sharply turned back when dried naturally, and spreading hori- 
zontally when wet. This activity is apparently a function of the color- 
less petioles, the green leaf-blades being quite rigid because of their suc- 
culence. The inflorescence is long and slender, the branchlets being mostly 
less than 1 cm. long. The corollas are rotate, about 2 mm. across, yellow 
or green, sparsely hispid externally. The ovaries are glabrous or nearly 
so, and the fruit is a didymous berry, or sometimes it is globose because 
of the abortion of one seed. Fully ripe and dehydrated fruits are black 
and wrinkled. 

Galium hardhamae Dempster, sp. nov. Herba dioica perennis, ramis 
decumbentibus vel rare scandentibus. Caules hispidi, internodiis longis 
cm. 14--2'%, quam foliis saepe multo longioribus. Folia sena plus minusve 
reflexa, mm. 2—4 longa, ovata, acuta, ambis paginis hispida, in vita plano- 
convexa, in herbario deorsum lateraliter volvata. Inflorescentia longa 


168 MADRONO [Vol. 16 


angustaque, ramis brevibus plus minusve 1 mm. Corollae rotatae, di- 
ametro circa mm. 2, flavae vel virides, extra sparse hispidae. Ovaria 
glabra. Fructus carnosi didymi vel saepe orbiculares, laeves. 

Type. Seaward slope of Santa Lucia Mountains in grove of Sargent 
Cypress, south ultimate fork of Alder Creek, southwestern Monterey 
County, California, at 2200-2500 feet, Clare Hardham 5650 (JEPS). 

Other collections. Monterey County: upper reaches of Alder Creek, Dempster & 
Hardham 1406 (JEPS) ; Villa Creek south of Lion Den Spring, Hardham 6065. San 
Luis Obispo County: Waterdog Creek, Hardham 6380 (JEPS); Cypress Swamp 
just northeast of Cypress Mountain, Hardham & Dempster 5703 (JEPS), Hardham 
3962 (JEPS) ; Spanish Cabin Creek, Hardham 5963; Tobacco Creek, Hardham 5967 ; 
headwaters of Chris Flood Creek (San Carpoforo), Hardman 6145. Numbers other- 
wise undesignated are in the private collection of Mrs. Hardham at Paso Robles, 
California. 

Jepson Herbarium, Department of Botany 
University of California, Berkeley 


A NEW SPECIES OF CRYPTANTHA (SECTION 
CIRCUMSCISSAE) FROM CALIFORNIA AND TWO 
RECOMBINATIONS (SECTION CIRCUMSCISSAE AND 
SECTION ANGUSTIFOLIAE)! 


KUNJAMMA MATHEW AND PETER H. RAVEN 


Cryptantha circumscissa (H. & A.) I. Johnston is an annual herb that 
occurs over a wide area of western North America, from Wyoming and 
central Washington to Arizona and northern Baja California. In one 
population, supposed to be referable to this species and located north of 
Adelanto on the Mojave Desert of San Bernardino County, California, 
three distinct entities were found, differing modally from one another in 
corolla size. The largest-flowered group of plants had corollas 4-6 mm. 
in diameter and pollen grains 5.5—6.5 w long. These measurements are 
beyond the range of variation characteristic of C. circumscissa. With 
further exploration it was found that plants with such measurements 
comprised a distinctive series of populations from a limited area in south- 
ern California, and they are described below as a new species. 

Cryptantha similis Mathew & Raven, sp. nov. Herba annua sectionis 
Circumscissae, a C. circumscissa persimilie, at differt: corolla 4-6 mm. 
lato; granis pollinis 5.5—6.5 py longo. 

Slender or bushy, bristly-pubescent annual herb, cymosely branched 
from the base, with ascending branches to 10 cm. long; leaves linear to 
narrowly oblanceolate, 3-10 mm. long, inconspicuously pustulate, with 


1 Thanks are due Professor Harlan Lewis for pointing out the variation pattern 
leading to this study and for a critical review of this manuscript, and to the curators 
of the following herbaria for permission to examine material in their care: British 
Museum (Natural History), California Academy of Sciences, University of California 
(Berkeley), Dudley Herbarium (Stanford University), Pomona College, Rancho 
Santa Ana Botanic Garden, and Royal Botanic Gardens (Kew). 


1962 | MATHEW & RAVEN: CRYPTANTHA 169 


no apparent veins, well distributed but congested just below the short 
inflorescences; inflorescences to 1.5 cm. long, the flowers axillary and 
very crowded; corollas conspicuous, 4—6 mm. in diameter, white, yellow- 
ish at throat; pollen 5.5—6.5 p» long, oblong; fruiting calyx 2.5-4 mm. 
long, bristly pubescent, the sepals united to near the middle, with a cir- 
cumscission just below the sinuses, the upper half falling away at ma- 
turity, the persistent portion cupulate; pedicels less than 0.5 mm. long; 
nutlets 4, homomorphous, lance-ovate, 1.2-1.5 mm. long; gynobase 
about two-thirds the height of the nutlets, slender-pyramidal; style equal 
to or barely exceeded by the nutlets. Gametic chromosome number, n=6. 
Type. 8.7 miles north of Adelanto on United States Highway 395, San 
Bernardino County, California, Lewis & Mathew 1113A (RSA). 
Additional specimens examined. California. Kern County: Red Rock Canyon, 
Howell 4925 (with C. circumscissa); Mojave, Lemmon in 1881 (with C. circum- 
scissa) ; 12 miles southeast of Mojave, Crum 1798. San Bernardino County: Hesperia, 
Spencer 387 (with C. circumscissa) ; north of Hinkley, Anderson 6853; near Victor- 
ville, Jones in 1926 (with C. circumscissa), Lee 8545, Wilson 31; Randsburg road, 
Weston 621; 10 miles north of Adelanto, Kirby 1116; Barstow, Munz 2544 (with 
C. circumscissa) ; between Hesperia and Adelanto, Munz 4474; 1.7 miles northeast 
of Helendale, Raven 11951; Swartout Canyon, Hall 1537; Swartout Valley, Munz 
4618; Cajon Pass, Johnston in 1920, Munz et al. 4672, Parish 11832. Los Angeles 
County: above Littlerock, Ray 931; between Big Rock Creek and Little Rock Creek 


on State Highway 138, Abrams 13946; Arraster Creek, San Gabriel Mountains, 
Peirson 1001. 


Cryptantha similis and C. circumscissa comprise the section Circum- 
scissae I. Johnston (Contr. Gray Herb. 74:40. 1925). Cryptantha similis 
has both larger corollas and larger pollen grains, the corollas of C. cir- 
cumscissa being only from 1—4 mm. in diameter and the pollen grains 
from 7—9 vy. in length (these measurements based on an examination of 
more than 100 collections from throughout the range of C. circumscissa 
and including the type: Snake Country, “California,” Tolmie, K). 

The gametic chromosome number of C. s¢milis is n=6, whereas that of 
C. circumscissa is usually n=12, although one plant of C. circumscissa 
from the locality north of Adelanto had n=18 (table 1). As already 
stated, three distinct morphological entities were found at the location 
north of Adelanto. One of these was the large-flowered C. similis just 
described. The remaining plants at this locality fell into two groups 
with respect to corolla size, both conforming, however, to the size limits 
given above for C. circumscissa. One of the plants of the smaller-flowered 
group was counted and was found to have a chromosome number of 
n=18 (hexaploid; table 1), whereas the plant which was counted from 
the group with medium-sized flowers had n=12 (tetraploid), the same 
chromosome number that was found in plants of C. circumscissa from 
other populations. The difference in pollen size between these two groups 
was not significant. The n= 18 plant from north of Adelanto might have 
had an allohexaplaid origin, with tetraploid C. circumscissa (n=12) and 
diploid C. stmilis (n=6) as the probable parents. Some tetraploid plants 
of C. circumscissa that were examined from other localities, however, had 


170 MADRONO [Vol. 16 


TABLE 1. CHROMOSOME NUMBERS OF CRYPTANTHA SPECIES* 


C. circumscissa (H. & A.) I. Johnston subsp. circumscissa 
n=12. Nevada. Washoe County: 6.1 miles north-east of Sparks, Raven 14287. 
Esmeralda County: Lida Pass, Raven 15476. 
California. Mono County: Sherwin Grade, Raven 14295, 14296; Paradise 
Camp, Raven 14262. Inyo County: Bishop Creek, Raven 14302, 14303. 
Kern County: Walker Pass, Raven 13992. San Bernardino County: Len- 
wood, Raven 13902; 8.7 miles north of Adelanto, Lewis & Mathew 
1113B. 
n=18. California. San Bernardino County: 8.7 miles north of Adelanto, Lewis 
& Mathew 1113C. 
C. micrantha (Torr.) I. Johnston subsp. lepida (Gray) Mathew & Raven 
n=12. California. Riverside County: San Jacinto Mountains, Raven 14241. 
C. micrantha subsp. micrantha 
n=12. California. San Diego County: Borrego Valley, Raven 14847. 
C. similis Mathew & Raven 
n=6. California. San Bernardino County: 8.7 miles north of Adelanto, Lewis 
& Mathew 1113A. 


* Vouchers deposited at Rancho Santa Ana Botanic Garden, Claremont, California. 


corollas as small as those of the Adelanto hexaploid. For example, at two 
California localities outside the range of C. similis (Sherwin Creek, Mono 
County; Bishop Creek, Inyo County), two groups of plants were found 
growing together that corresponded in corolla size with the tetraploid 
and the hexaploid (n=18) from north of Adelanto. In both of these 
latter cases, however, all plants examined were tetraploid (n=12). 
Thus there appears to be no basis for the taxonomic segregation of the 
hexaploid. As for the difference in corolla size between the diploid and 
tetraploid, it is possible that the diploid may be allogamous to a greater 
extent than is the teraploid, and thus follow the sort of correlation between 
polyploidy and autogamy discussed in some detail by Grant (Am. Nat. 
90:319-322. 1956). Although the chromosomes are small and difficult to 
observe, there is no suggestion of multivalent formation in the tetraploid, 
and hence it may have had an alloploid origin. Cryptantha similis, how- 
ever, appears to be the only diploid closely enough related to C. circum- 
scissa to have participated in its origin. 

In view of these results, it is pertinent to discuss the patterns of morpho- 
logical variation of C. circumscissa. In the southern Sierra Nevada of 
California at high elevations there occurs a distinctive geographical entity, 
Cryptantha circumscissa subsp. rosulata Mathew & Raven, comb. nov. 
(C. circumscissa var. rosulata J. T. Howell, Leafl. West. Bot. 6:104. 
1951). Based on measurements of its pollen, C. circumscissa subsp. 
rosulata is probably tetraploid like subsp. cércumscissa. Another variant 
involves hispid plants from the eastern slopes of the Sierra Nevada which 
have been separated as Krynitzkia dichotoma Greene or as Cryptantha 
circumscissa var. hispida (Macbride) I. Johnston (Contr. Gray Herb. 
74:1-114. 1925), but inasmuch as they occur intermingled with plants 


1962 | REVIEW 171 


typical of subsp. circumscissa in this area and less commonly elsewhere 
and do not appear to be sharply distinct morphologically, we prefer not 
to recognize them taxonomically. These patterns of variability in C. 
circumscissa are apparently analogous with that concerned with corolla 
size, which has already been discussed, and, like it, they are doubtless 
reinforced by autogamy. 

Since the detection of Cryptantha similis became possible following 
the determination of its chromosome number, we also investigated an 
apparently analogous pair of taxa in the related section Angustifoliae, 
Cryptantha micrantha (Torr.) I. Johnston subsp. micrantha, with very 
small flowers, and another entity with larger flowers, Cryptantha micran- 
tha subsp. lepida Mathew & Raven, comb. nov. (Eritrichium micrantha 
var. lepidum Gray, Syn. Fl. 2:193. 1878). In this case, however, both 
taxa were found to have the same gametic chromosome number, n = 12 
(table 1). It should also be noted that the large- and small-flowered taxa 
have not been found growing together. They appear to be largely geo- 
graphical entities best recognized as subspecies. 


University of California, Los Angeles 


Rancho Santa Ana Botanic Garden 
Claremont, California 


REVIEW 


Taxonomy of Flowering Plants. By C. L. Porter. viii + 452 pp., W. H. Freeman 
San Francisco. 1959. $6.75. 

“Taxonomy of Flowering Plants” is one in the series of high-quality biology texts 
published by Freeman and Company. Dr. Porter states that he hopes it will help to 
fill the gap he sees existing between “texts that are really reference books for ad- 
vanced students and much abbreviated texts that have had much of the meat of the 
subject deleted from them.” This rather effective compromise is a work of some 450 
pages, suitable for introductory courses of either one or two semester’s length. It is 
divided into three principal parts: History, Principles, and Methods; Selected Orders 
and Families of Monocotyledons; and Selected Orders and Families of Dicotyledons. 
A 16-page glossary precedes the Index. 

Part I, History, Principles, and Methods, is rather abbreviated. It should be 
entirely satisfactory for many introductory courses, but will require some supple- 
mentation in courses where a substantial portion of the students requires more de- 
tailed information. An exposition of aims, history, literature, field and herbarium 
methods, nomenclature, concepts of taxa, construction and use of keys, phytography 
and terminology, and phylogeny and classification of angiosperms which can be en- 
compassed within 140 pages and yet prove entirely satisfactory for a wide variety of 
taxonomy courses is probably impossible. Dr. Porter intended to produce a concise 
treatment, and he is no doubt aware that some will find his work excessively synoptic 
in places. The reviewer believes that the material on field methods and the chapter 
entitled “Concepts of Taxa” are cases in point. In addition, instructors who stress 
nomenclature will find his chapter of the same name very brief indeed. However, 
the author generally has been remarkably successful within his self-imposed space 
limitations. The chapter on phytography and terminology is a good one. It is four 
times as long as the average chapter in this section, and replete with illustrations. 


172 MADRONO [Vol. 16 


The student will appreciate the detail in this chapter, and the instructor will find that 
the drawings can eliminate many blackboard drawings or pencil sketches on his part. 
The illustrations are excellent. The drawings are mainly the work of Evan Gillespie, 
but a few are from Gray’s “Lessons.” This chapter also introduces the symbols for 
floral parts which are used extensively in the last two portions of the book. These 
symbols are simple ones that make it possible to portray the characteristics of a fam- 
ily in an easily grasped floral diagram. Every chapter in Part I except the one on 
the construction and use of keys has a list of references pertaining to the subject 
matter of the chapter. Some of these lists are more than two pages long. These refer- 
ences add considerably to the merits of the book. They should encourage the student 
to follow up subjects that especially interest him. 

Parts II and III, dealing with slightly more than one hundred families of flowering 
plants frequently encountered in the North American flora, are very good. Porter’s 
treatment should help the student to see the classificatory function of taxonomy as 
other than an arbitrary system of pigeon-holing. He has maintained a balance be- 
tween convenience and progress toward more truly phylogenetic systems. His treat- 
ment of the monocots (Part II) is basically that of Hutchinson, whereas that of the 
dicots (Part III) is a modified Englerian sequence. These classes are divided into 
subclasses, and these in turn into orders and families. A few of the families are further 
divided into sub-families or tribes. The intent here is not to construct an elaborate 
system of hierarchies, but rather to show in a natural way that taxonomy is a science 
that categorizes living things upon the basis of similarities in form and function. 
Porter’s treatment of the identificatory function of taxonomy is excellent. There is a 
succinct description, a mention of representative genera, a floral diagram, and draw- 
ings for almost all of the families treated. In addition, there are many good photo- 
graphs of representatives of the larger families. These illustrations are designed to 
teach the student to recognize, without recourse to a book, the important families of 
flowering plants. The floral diagrams accurately symbolize the characteristics of a 
given family, yet are easy to comprehend. The drawings, in addition to their sight- 
recognition function, frequently show (and label) features that elicit questions from 
students using dichotomous keys. For example, the involucre, bracts, glands, and uni- 
sexual flowers of the Euphorbiaceae are labeled, as are the involucre, involucel, car- 
pophore, mericarp, ribs, oil tubes, and stylopodium of the Umbelliferae. The special- 
ized structures characteristic of the Cyperaceae, Gramineae, and Compositae are 
treated in the same way. The advantages of these drawings over verbal descriptions 
or blackboard sketches or projected slides are obvious. It is difficult to imagine a 
more teachable method than Porter’s combination of text, floral diagrams, drawings, 
and photographs. 

“Taxonomy of Flowering Plants” impresses this reviewer as being the best avail- 
able text for many introductory taxonomy courses, especially one-semester or one- 
quarter ones. It is generally free from errors and objectionable features. However, 
it should be noted that the tuft of hair on Epilobium seeds is better described as 
comose, rather than comatose (p. 342). Also, from a realistic point of view, conserved 
family names probably should be used in an elementary text rather than the permis- 
sible but relatively unused Apiaceae, Lamiaceae, etc. Gillespie’s illustrations are, as 
usual, very good, and Porter’s prose makes the text a readable one. The relative 
brevity of Part I (History, Principles, and Methods) poses certain difficulties, but 
this material can be supplemented by the instructor if he feels this is necessary. How- 
ever, the feature that distinguishes this book from other contemporary texts is the 
material on plant structures and family characteristics. These sections are perhaps 
the nearest approach to a do-it-yourself method of instruction in plant identifica- 
tion that can be devised. It should be welcomed by instructors and students in the 
classroom, and also by persons who wish to learn how to identify plants without 
having to take a course to do so. JoHN Moorinc, Department of Botany, Washing- 
ton State University, Pullman, Washington. 


INFORMATION FOR CONTRIBUTORS 


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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 
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holon 3 


MADRONO 


VOLUME 16, NUMBER 6 APRIL, 1962 


Contents 


PAGE 
THE SANTA LUCIA CUPRESSUS SARGENTII GROVES AND 
“ THEIR ASSOCIATED NORTHERN HyYDROPHILOUS AND 

ENDEMIC SPECIES, Clare B. Hardham 173 


CALIFORNIA BOTANICAL EXPLORERS—XII. 
Jouwn Mitton BicELow, Willis Linn Jepson 179 - 


A SUBARBORESCENT NEW ERIODICTYON (HYDROPHYLL- 
ACEAE) FROM SAN LuIS OBISPO CoUNTY, CALIFOR- 


NIA, Philip V. Wells 184 
A NEw SPECIES OF QUERCUS FROM BAJA CALIFORNIA, 

Mexico, Cornelius H. Muller 186 
PARASITISM IN PEDICULARIS, Elizabeth F. Sprague 192 


NOMENCLATURE, LIFE HISTORIES, AND RECORDS OF 
NorTH AMERICAN UREDINALES, George B. Cummins 
and John W. Baxter 201 


REvIEW: Victoria Padilla, Southern California Gardens, 
An Illustrated History (Elizabeth McClintock). 204 


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 
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont, California 

HERBERT F. CoPpELAND, Sacramento College, Sacramento, California 

Joun F. Davinson, University of Nebraska, Lincoln 
Mitprep E. Marutas, University of California, Los Angeles 24 
Marion OwWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
Tra L. Wiccins, Stanford University, Stanford, California 


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


Business Manager and Treasurer—JoHN H. THOMAS 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Roxana S. Ferris, Dudley Herbarium, Stanford University, Stanford, 
California. First Vice-President: Edward C. Stone, School of Forestry, University 
of California, Berkeley. Second Vice-President: Thomas C. Fuller, Department of 
Agriculture, State of California, Sacramento. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding Sec- 
retary: Margaret Bergseng, Department of Botany, University of California, Berke- 
ley. Treasurer: John H. Thomas, Dudley Herbarium, Stanford University, Stanford, - 
California. 


1962 | HARDHAM: CUPRESSUS SARGENTII 173 


THE SANTA LUCIA CUPRESSUS SARGENTII GROVES 
AND THEIR 
ASSOCIATED NORTHERN HYDROPHILOUS AND 
ENDEMIC SPECIES 


CLARE B. HARDHAM 


Cupressus sargentit Jepson occurs in the Coast Ranges of California 
from extreme northern Mendocino County in the North Coast Ranges to 
Santa Barbara County in the South Coast Ranges, where its most south- 
ern locality is an isolated outpost in the San Rafael Mountains. Through- 
out its range it is known as a species which is almost always restricted 
to soils of serpentine content. It is common over large areas in the North 
Coast Ranges, but occurs much more sparingly and interruptedly in the 
South Coast Ranges. In the Santa Lucia Mountains of the South Coast 
Ranges, it is restricted to a few small areas in Monterey and San Luis 
Obispo counties. Several local endemics, with only minor exceptions, are 
restricted to the same habitat. In addition, the dampest and most shaded 
groves provide ecological niches where several northern hydrophilous 
species persist. 

The groves of C. sargentu in the Santa Lucia Mountains occur on 
metamorphosed serpentine which was formed at various times during the 
Jurassic Age. In the Santa Lucias the exposed serpentine extends for 
nearly forty miles along the major fault zones of the range from south 
of the Cuesta Grade in San Luis Obispo County to Plaskett Creek in 
Monterey County. It is often slightly oxidized and owes much of its red 
color to superficial deposits of water-borne iron. It is also highly faulted, 
metamorphosed, and intruded with volcanics. Basalt, rhyolite, radi- 
olarian chert, and local concentrations of various minerals are often found 
in conjunction with it. Gold, chrome, and quicksilver have all been mined 
near one or another of the cypress groves, and there may be commercial 
deposits of manganese present. The cypress grove south of Lion Den 
Spring is the only grove which occurs on a substratum obviously different 
from the usual form of serpentine. Here the serpentine is thoroughly im- 
pergnated with aluminum, and the resultant soil is white. Although geolo- 
gists suspect that the cypress groves may indicate the presence of some 
mineral besides serpentine, no such correlation has yet been established 
for the Santa Lucia cypress groves. 

Actually the cypress groves occupy only a tiny fraction of the total 
area of serpentine in the Santa Lucia Mountains. The combined areas 
of the C. sargentiu groves can scarcely exceed four or five square miles, 
whereas the serpentine formations of the Santa Lucias have a total area 
of roughly 400 square miles. The cypress groves are found along either 
the King City or the Pine Mountain faults, regions of numerous springs, 


Mapro..0, Vol. 16, No. 6, pp. 173-204, April 18, 1962. 


174 MADRONO [Vol..16 


and the groves occur at the heads of the streams which are, in all but the 
very dryest years, permanent or semipermanent. The cypresses extend 
down the mountains along the streambeds, especially on the northeast 
slopes of the range, and isolated trees frequently grow in the streambeds 
below the groves. 

Most of the cypress groves are located along the main ridge of the 
Santa Lucias at altitudes of about 2500 feet. There are also some cypress 
groves near Bryson on the ridge formed by the King City Fault, the 
largest of which is on Waterdog Creek at elevations of 2000 to 2500 
feet. A few miles south of this grove there used to be “cedars in swamps,” 
but the trees never reached maturity after a fire in 1953, and they were 
again burned in 1960, so it is doubtful if these cedar swamps will ever 
again exist. There are many scattered trees on sandstone in the Los 
Burros Creek drainage at low elevations. Although they are occasionally 
as much as a mile distant from the serpentine ridges to the west, pre- 
sumably there are serpentine ions in the groundwater, and, since there 
are few competing species, the cypresses have succeeded in establishing 
themselves. I have found only one small grove on the serpentine above 
these scattered trees in the Los Burros drainage. 


As with many other species that occur in mesic habitats, some of the 
cypresses have established themselves on the rocky ridges that sometimes 
mark the upper borders of the cypress groves, but if one has the courage 
to explore, it is apparent that most of the trees are growing in relatively 
damp places and even, occasionally, in small swamps. A few of the 
cypress groves, including the easily accessible grove just north of San 
Luis Obispo, west of the Cuesta Grade, appear to be quite dry. Despite 
appearances, however, on the foggy morning of September 22, 1958, the 
ground beneath the cypress trees in this grove was wet down to the zone 
of permanent moisture. 

The only Forest Service station in the entire area that keeps moisture 
records is on Rocky Butte at 3590 feet, about 1000 feet higher than 
the cypress groves. On numerous mornings which are described as ‘‘clear”’ 
on their records, the relative humidity readings are recorded as being as 
high or higher than they are on mornings when fog is specified at that 
altitude. This corroborates my impression that the coastal fogs rise to 
a considerable altitude in the Santa Lucia Mountains and that the cypress 
groves probably receive an appreciable amount of moisture from summer 
fogs. In wet years some of the groves must receive a surprising amount 
of rain. At the 7X Ranch at the foot of Cypress Mountain near the head- 
waters of Las Tablas Creek, the rainfall varies from 40 to 80 inches 
annually (San Luis Obispo County Water Survey). The average at the 
Krenkel Ranch near Alder Creek is 60 inches. In a wet year at the Ocean 
View Mine near Burnett Peak, the rainfall was 117 inches, and in the 
dry winter of 1960-61, the total rainfall was more than 36 inches, the 
rain gauge having overflowed in one storm. 


1962 | HARDHAM: CUPRESSUS SARGENTII 15 


cf aS 
{ QO. 
ek, <. OEE 
YAS S CAP 
“\ fila wrros 7 3, 
! : BS 
Ne Salmon 
~ Cr. 
MONTEREY CO. : 
coho rere... = SAN LUIS OBISPO CO. RY 
Wsanf & Sn 1s ga By 
Burnett, PkA * AnD: 
) 5B 4 
\Arraro = S 
va = Gy Ck- 2 
Pine Mt. 9 RY 
A : =a 
Rocky Butte 5 
< a 
— , San Simeon ~ 
Senet Ch ie 
Cypress Mt 2". 7X Ranch 
Roso ‘eo 
Cambria\m 2227 ‘fe 


Pacific Ocean 


Cerro Alto 4 


San Luis Obispo 


Fic. 1. Distribution map of Cupressus sargenti in the Santa Lucia Mountains. 


Typically the boundaries of the cypress groves are sharp, with their 
drier edges bordered in the northern part of the range by mixed Ouercus- 
Pinus-Pseudotsuga woodland, and in the southern part by Quercus-Pinus 
woodland. 

On the eastern edge of the Santa Lucia Mountains there are two cy- 
press groves, one large grove on a northeast-facing slope at the headwaters 
of Waterdog Creek and a small grove in a swampy ravine at the head of 
the main branch of Los Burros Creek. These groves occur along the King 
City Fault. On the main ridge, along the Pine Mountain Fault, there are 
approximately fourteen groves. The largest and most southerly grove 
occurs west of the Cuesta Grade five miles north of San Luis Obispo 
and includes the Chorro Creek grove described by Carl B. Wolf in “The 
New World Cypresses” (EI Aliso 1:1—-444. 1948). The cypresses extend 


176 MADRONO [Vol. 16 


down the north side of the mountain as well as the south, and there are 
a few trees in Tassajara Creek. The next grove to the north is also large 
and is located on Cypress Mountain. Cypress Swamp is an area of springs 
in the extension of this grove down Delaganna Creek, a tributary of 
Las Tablas Creek. There are many groves near Burnett Peak where the 
tributaries of Arroyo de la Cruz, San Carpoforo (Chris Flood Creek), 
Tobacco and Little Burnett creeks have their origins on the main ridge 
of the Santa Lucias. Many of these groves are small, as well as inacces- 
sible, and I was not able to explore them before they were burned in the 
fire of 1960 and sprayed with 2-4-D in 1961. I am not sure how much 
of this area is included in Dr. Wolf’s description of the ‘“‘Pine Mountain 
Grove.” There are actually many small well-defined groves and not one 
large one as he implies. Also the grove at the head of Chris Flood Creek 
could not have been included since it was virtually impossible to reach 
until a few years ago. There are two groves at the head of Villa Creek 
in Monterey County, one north and one south of Lion Den Spring. The 
most northerly grove in the Santa Lucia Mountains is on the small tribu- 
tary of Alder Creek that joins the main stream about half a mile below 
the public camp ground. North of Alder Creek the serpentine formation 
dips down and disappears beneath the ocean at Plaskett Creek. 

Some of the groves have a thick layer of leaves covering the ground, 
and in these groves there are few herbaceous species except along the 
streambeds or rock outcrops. In other groves there has been a great deal 
of sheet erosion and a number of species grow in crevices in the rocks and 
on the relatively level areas. In spite of the great diversity in appearance 
of the different cypress groves, the associated flora is surprisingly uni- 
form, and one can expect to find many or all of the species listed below 
in any but the smallest cypress groves. Although I have not attempted 
a complete list of cypress forest endemics and associated northern species, 
I have included those which are characteristic of the majority of the 
groves. 

PINUS SABINIANA Dougl. 

P. COULTERI D. Don 

DENDROMECON RIGIDA Benth. 

ERIODICTYON CALIFORNICUM (H. & A.) Torr. As far south as Cypress Mountain. 

QUERCUS DURATA Jepson 

CEANOTHUS spp. 

ARCTOSTAPHYLOS spp. Typically A. obispoensis Eastw. 

LOMATIUM PARVIFOLIUM (H. & A.) Jepson 

ZIGADENUS FREMONTI Torr. 

BRODIAEA LUTEA (Lindl.) Mort. In the southern Santa Lucias, this species is not 
common and occurs only near springs and beside streams. 

FRITILLARIA LANCEOLATA Pursh 

POLYGALA CALIFORNICA Nutt. 

ERIOPHYLLUM CONFERTIFLORUM (DC.) Gray var. LAXIFLORUM Gray. Occasional 
plants of what appears to be E. lanatum (Pursh) Forbes as well as hybrids with 
FE. confertiflorum occur on nearby serpentine outcrops. 

CHEILANTHES SILIQUOSA Maxon. Occasionally C. californica (Hook.) Mett. or 
C. carlotta-halliae Wagner & Gilbert also grow in the cypress groves. 


1962 | HARDHAM: CUPRESSUS SARGENTII 177 


CHORIZANTHE VORTREIDII Bdg. As far south as Burnett Peak. This local endemic 
is rare on serpentine at high altitudes in the Santa Lucias but common on sandstone, 
especially at lower elevations. 

GALIUM HARDHAMAE Dempster. A Santa Lucia cypress forest endemic occurring 
as far south as Cypress Mountain. 

GALIUM CALIFORNICUM H. & A. Galium nuttallii Gray and G. andrewsii Gray also 
occur in one cypress grove or another. 

MOoNARDELLA PALMERI Gray. A Santa Lucia cypress forest endemic which occurs 
occasionally elsewhere on serpentine (Rinconada Mine east of Santa Margarita, 
Hoover 6146, and the west side of Cerro Alto). When it occurs in a cypress grove, the 
leaves are long, narrow and shiny. Outside the cypress forests the leaves are relatively 
dull and reminiscent of whatever variety of M. villosa Benth. grows nearby. In other 
characters as well, the plants appear to be intermediate. Monardella villosa does not 
grow in the cypress groves. 

In addition to the species in the above list, almost every cypress grove 
has some species of Carex. In groves with much litter covering the soil, 
the Carex is generally C. globosa Boott, which also occurs in the nearby 
oak-conifer woodland. Carex serratodens W. Boott grows beside the 
permanent streams and semipermanent streams. Carex senta Boott occurs 
in Cypress Swamp. Of the other three species of Carex, discussed below, 
one is a local endemic and the other two are northern species. 

Carex obispoensis Stacey, a San Luis Obispo County endemic, occurs 
along the intermittent streams and in low or level places along the top 
of the ridge in the Cuesta cypress grove and in openings in the surround- 
ing chaparral. It also grows on a rocky east-facing slope almost through- 
out one of the Burnett Peak groves which has a small semipermanent 
stream. In the dry year of 1960, only the plants in the obviously much 
damper lower half of the grove flowered. It is possible that C. obispoensts 
is primarily a plant of moderately dry serpentine soils and that its occa- 
sional occurrence at low elevations in boggy spots along streams that drain 
cyress groves (Stenner Creek,! Eastwood & Howell 2271 and Arroyo de 
la Cruz, Hoover 6684, 7951) should be considered exceptional. It has also 
been collected on “dry serpentine” outside of a cypress forest at Rin- 
conada Mine (Hoover 6115, 7208). 

Carex mendocinensis Olney has been found in three cypress groves. At 
Alder Creek the cypresses occur on several acres of rocky northwest slope 
as well as along the course of a semipermanent stream. Here C. mendoci- 
nensis is common, especially beside the stream. It is also common in the 
Chris Flood grove, where it grows both beside the intermittent streams 
and the permanent streams and on a northeast-facing slope that is prob- 
ably covered with hidden springs since the herbaceous cover is unusually 
luxuriant for a cypress grove. The small swampy grove at the head of 
Los Burros Creek also has an abundant growth of C. mendocinensis. In 
Cypress Swamp C. luzulina Olney grows beside the small permanent 


1Stenner Creek, according to the United States Geological Survey maps, is the 
correct name for this creek, though on many maps it is called Steiner Creek. It is 
Steiner Creek on the Eastwood and Howell labels. The tributary of San Simeon Creek 
that has its headwaters on Cypress Mountain is called Steiner Creek. 


178 MADRONO [Vol. 16 


streams that flow from the springs. I have not found C. mendocinensis or 
C. luzulina in the streambeds below the cypress groves. 

Many of the usual streamside plants of the Santa Lucias grow in the 
Swampy areas and beside the streams in those cypress groves that have 
permanent and semipermanent streams, for example, Mimulus guttatus 
DC., Lilium pardalinum Kell., and, at Alder Creek, Epipactis gigantea 
Dougl. In Cypress Swamp there are two northern hydrophilous species 
as well, Habenaria dilatata (Pursh) Hook. var. leucostachys (Lindl.) 
Ames and Parnassia palustris L. var. californica Gray. Both of these 
plants have been collected elsewhere in this region, the Parnassia on 
Santa Lucia Peak by M. S. Clemens in 1921, and the Habenaria in a 
swamp at Arroyo Grande, Hoover 7338. 

Until recently most of the groves had never been thoroughly explored 
and it was thought that the cypresses were restricted to the dry serpentine 
outcrops. In reality they are most abundant on mesic, north-facing slopes 
where they grow in the vicinity of streams and in swampy areas produced 
by springs. In several places the cypresses do not occupy the total area 
of apparently suitable moist habitat. Neither do the associated species 
extend beyond the sharply delimited boundaries of the groves. It is 
understandable that the cypress trees may not be able to grow down 
the beds of streams for any distance below the serpentine to which they 
are usually restricted, but it is surprising that the associated species are 
similarly limited. As already mentioned, the two northern species of 
Carex, C. mendocinensis and C. luzulina, do not grow in the water- 
courses below the boundaries of the cypress groves. The cypress groves 
seem to offer these species a special ecological niche upon which their 
persistence depends. Although the causal relationships are not fully 
apparent, it seems probable that the discontinuous occurrence of the 
cypress groves, their usual sharp delimitation, and the restriction of the 
associated species to the groves depend both on the influence of the ser- 
pentinized soil, which may act by reducing competition from the neigh- 
boring chaparral and forest species, and on the obvious need of all these 
species for a relatively abundant water supply. 

It is possible that cypress forests were formerly more extensive in the 
Santa Lucia Mountains. Typical cypress forest species occur in at least 
three areas where cypress trees do not grow. Carex obispoensis and Monar- 
della palmeri, the latter showing evidence of integration with M. villosa, 
have been found in one or two places on serpentine south of the Cuesta 
cypress grove as well as on Cerro Alto north of that grove. Similarly, 
near the Burnett Peak cypress groves Galium hardhamae and Monar- 
della palmeri grow in occasional chaparral openings where there are no 
cypresses. 

Vouchers for all these cypress forest species have been deposited at 
the Herbarium of the California Academy of Sciences. 

Mr. James Collord, geologist for the Madonna Construction Company 


1962 | JEPSON: BIGELOW 179 


of San Luis Obispo, described for me the geology of the Pine Mountain 
fault zone where most of the cypresses occur and helped me to explore 
the Burnett Peak and Chris Flood Creek groves. 


Creston Star Route 
Paso Robles, California 


CALIFORNIA BOTANICAL EXPLORERS—XII 
WILLIS LINN JEPSON 


John Milton Bigelow 


[The following account of a significant portion of Bigelow’s whereabouts and 
itinerary as a botanical traveler in central California is as pertinent and needful 
today as it was when Dr. Jepson originally wrote it, pin-pointing as it does the 
type localities of a number of well-known -and more or. less widespread species 
originally collected by Bigelow at the annoyingly elusive “Duffield’s Ranch” of 
the Torrey account in the Pacific Railroad Reports. The first five paragraphs have 
already been published (in the twice resuscitated journal “Erythea,” vol. 3, No. 13, 
pp. 102-104, Dec., 1938), this initial portion ending with the promise “to be 
continued.” Inasmuch as this number of “Erythea” had a very limited circulation 
and is not likely to be available to many readers, the entire article, from a type- 
script copy recently come to light among Dr. Jepson’s accumulated papers, is offered 
herewith —-R. BacicaLupi, Curator, Jepson Herbarium, Department of Botany, 
University of California, Berkeley. ] 


The name of Bigelow is associated with many of the early discovered 
plant species of California. John Milton Bigelow was born at Peru, Ben- 
nington County, Vermont, on June 23, 1804. When he was eleven years 
old, his family moved to Ohio where he went to the public schools and 
attended the Medical College of Ohio, at which institution he was gradu- 
ated March 8, 1832. In November of this same year, he was married to 
Maria R. Miers of Lancaster, Ohio, where he began the practice of medi- 
cine. In 1850 he was appointed surgeon to the Mexican Boundary Com- 
mission. While on duty with this survey, he collected and studied the 
native vegetation of the boundary. His specimens formed part of the 
material used by John Torrey in the preparation of the Report of the 
Mexican Boundary Survey (1859), which was under the charge of Major 
W.H. Emory. Bigelow’s name is frequently cited in its pages. In 1853 
Dr. Bigelow accepted the position of surgeon and botanist to the expedi- 
tion of the Pacific Railroad Survey under Lieutenant A. W. Whipple, 
which left Fort Smith, Arkansas, and proceeded along the Canadian 
River and across the Llano Estacado to Albuquerque. The Rio Grande 
was crossed on November 10, 1853 and the expedition, following down 
the Bill Williams Fork, crossed the Colorado River at the mouth of Wil- 
liams Fork on February 28, 1854, ferried it about sixty miles above and 
traversed the Mohave Desert to the Cajon Pass, thence to Los Angeles 
and San Pedro. Here the survey of Lieutenant Whipple ended; but it was 


180 MADRONO [Vol. 16 


now the most favorable botanizing season in California and Dr. Bigelow 
in his own interest came north to San Francisco and Marin County to 
collect more fully the native vegetation of California. His zeal took him 
across the Great Valley into the Sierra Nevada where he remained until 
nearly June 1, 1854. He was not the first, but he was amongst the first 
explorers to botanize at all extensively in the Sierra Nevada. Of Bige- 
low’s entire journey, John Torrey said that “his ample collections,” made 
_ in 1853 and 1854, “were brought home in perfect order,” and adds: “A 
number of new genera and more than sixty new species have been dis- 
covered by Dr. Bigelow, and he has added much valuable information 
upon many heretofore imperfectly known plants.” 

In the course of the years the student of California botany needs to 
consult and study repeatedly the pages of volume four of the Pacific 
Railroad Reports where, in about one hundred pages, John Torrey has 
given an account of the Bigelow plants supplemented by twenty-five very 
fine lithographic engravings. In the citation of localities for California, 
several extremely local place names occur which are not to be found on 
maps and which have been difficult to place. One of these is ‘“‘Duffield’s 
Ranch,” which is important as the collecting station for a number of 
Bigelow plants and especially as the type locality for Allium tribracteatum 
Torr. and other plants then new to science. 

To the systematic botanist whose initial or primary interest is in the 
living plant in its natural state, in its biology, ecological form and as- 
sociates and habitat conditions, the type locality is vastly a more fruitful 
quest than the type specimens of early collectors, which are frequently 
unsatisfactory, fragmentary, or so decadent structurally as to be essen- 
tially useless. At the best the type specimen can usually do little more 
than enable one to make sure that a given binomial belongs to a particular 
unit; whereas the type locality opens up a new and living world whose 
potentialities may not be exhausted for years. It was for this reason, as 
well as for some uncertainties attaching to the identity of Allium tri- 
bracteatum, that I wished forty years ago to find Duffield ranch which 
is placed no more definitely in the text than “Sierra Nevada.” The name 
could not be found on any available map or in any index, and constant 
inquiry amongst pioneers failed in like manner. Resort was then had to 
a stratagem. I took such dates as are given in Torrey’s paper, arranged 
them in chronological sequence with their localities and I thus made an 
itinerary for Bigelow. It was soon found that the dates were not always 
dependable, but where the date agreed with the locality three times or 
more, it was used on the list. In this manner I obtained the following 
chronology: From April 4 to April 12, he was in Oakland and the Oakland 
Hills; April 17 and 18 at Point Reyes; April 20 at Corte Madera; April 
25 to 27 in the Napa Valley; May 7 and 8, Stockton to Knights Ferry; 
May 9 at Sonora; May 10, 11, 12 at ‘““Duffield’s Ranch”; May 14, Car- 
son’s, Stanislaus River; May 14, Robinson Ferry, Stanislaus River, and 


1962 | JEPSON: BIGELOW 181 


Murphy’s Camp; May 15, Washington Mammoth Grove (Calaveras 
Grove); May 17, Mokelumne Hill; May 18, Ione Valley; May 19 and 
20, Grass Valley; May 21, Nevada City and Downieville; May 23, Middle 
Yuba River; May 25, Marysville and Marysville Buttes. 

This list made, it was a natural inference that Duffield ranch was 
somewhere between Sonora and the Stanislaus River or Murphy’s Camp 
in Calaveras County. Any locality in this region was too low in altitude 
for the species concerned, so that the inference seemed in some way 
fallacious. 

No systematic effort had been made to find Duffield ranch but for two 
decades the query had been kept in mind and used on any likely prospect 
or as a sort of poser to the geographically minded. Now that, in 1916, 
I had made a Bigelow calendar, a wider appeal was made to local his- 
torians and editors of newspapers in the foothill towns, but the effort 
again proved unavailing. 

In 1918, my friend George James Peirce, Professor of Plant Physiology 
at Stanford University, was in Berkeley as Professor of Botany in the 
University of California Summer School. At my invitation, he joined in 
a week-end excursion to Sonora. We arrived in this county-seat town of 
Tuolumne County in mid-afternoon on the Fourth of July. It seemed that 
every old-timer on Sonora’s High Street was, on this holiday, willing and 
anxious to tell us about Duffield ranch; it was out near the old Bob 
Finney place; it was on the Bald Mountain road; it was in this direction 
and that, in no case more than a few miles away. It developed at once that 
Duffield was a real character of the 1850’s and a dead-shot. Ancient 
legends regarding him are still spun on the High Street. Finally we found 
Robert Marshall, a shrewd clear-headed “‘down-east”’ Yankee, ninety-one 
years old, who came to Sonora in the early 1850’s, and who recalled with 
remarkable clearness the local events during the gold days. Said Marshall: 
“Yes, I knew Duffield. There is a Duffield Mine near Soulsbyville but no 
Duffield ranch.” Then he went on to give an elaborate, particularized and 
colorful history of Duffield, his activities and associates. In the course 
of describing the personality of Duffield, he remarked that there was only 
one man whom Duffield feared, namely Ben Soulsby. “‘Was Ben Soulsby 
still living?” “Yes; Ben Soulsby lives in a cabin near Soulsbyville.”’ We 
went to Soulsbyville, found the cabin a few miles from the village and 
found Ben Soulsby. When I told my errand he seated himself on a stone 
step in front of his cabin door. Seventy-eight years of age, he was as 
utterly distrustful of the stranger and as taciturn as Marshall in Sonora 
had been hospitable and loquacious. Little could be had from him save 
that there was no Duffield ranch, but that there was a Duffield mine. It 
was up the canon a half-mile; the hole still there, any one could go look 
at it. Hereupon I played the only valuable card left me and which, it was 
felt, would be decisive one way or the other. It was obvious that of early 
events Soulby’s memory was clear and sharp. So I took from under my 


182 MADRONO [Vol. 16 


arm a large volume which I had brought along, volume four of the Pacific 
Railroad Reports. I opened the book to a marked page, placed it in 
Soulsby’s hands and pointed to ‘“Duffield’s Ranch” without a syllable of 
comment. Soulsby looked at the printed words a long, long time without 
remark. We, his visitors, sat on the ground near him and waited patiently. 
At length he turned to the title page and noted the date, 1856. Then he 
went on communing with himself. Finally, he closed the book, ran his 
rough hand through the enormous shock of iron-gray hair that thatched 
his head and said very simply and briefly: ‘“Now I bethink myself. There 
was a Duffield ranch. It was up beyant Confidence. Duffield was up there 
in 1854. The Ward Pike place we call it now.” 

There was little difficulty finding the Ward Pike place on the Sonora 
Pass road at about 4800 feet altitude. It seems a plausible inference that 
Bigelow, coming as a traveler into Sonora, would inevitably meet so showy 
and ubiquitous a character of the High Street as Duffield, and that Duf- 
field took Bigelow to his mountain ranch. At Duffield ranch, Bigelow 
spent three days, May 10, 11, and 12, 1854, the year corresponding with 
the year named by Ben Soulsby. From the ranch as a base he must have 
made excursions upwards, since some of the species collected by him 
belong at higher altitudes than Duffield ranch. In his own account of the 
forest trees of California, it is unmistakable that Bigelow was “high up 
in the mountains east of Sonora (almost in the snowy regions)”’ (p. 26) 
and he further states that he was “fifteen or twenty miles southeast of 
Sonora, on the headwaters of the Stanislaus and Tuolumne rivers”’ 
(p. 24), but the only practicable direction eastward from Sonora he was 
likely to have traveled is northeast rather than ‘“‘southeast.”’ The Ward 
Pike place lies on the top of the ridge between the South Fork Stanislaus 
River and the North Fork Tuolumne River, truly northeastward from 
Sonora. 

Duffield ranch then, or the region immediately above it, is the type 
locality for the following new species or varieties collected by Bigelow: 
Philadelphus lewisu var. parvifolius Torr., Potentilla tridentata (Torr.) 
Greene (Horkelia tridentata Torr.), Sanicula tuberosa Torr., Phlox occt- 
dentalis Durand ex Torr. [today generally considered a phase of Phlox 
speciosa Pursh|, and Allium tribracteatum Torr. In addition, some nine- 
teen other species are listed as of “‘Duffield’s Ranch” by Torrey. One of 
these, Pinus sabiniana, is perhaps a slip on Torrey’s part, since it is not 
found on the Sonora Pass road at this altitude. It may be noted that Bige- 
low himself in his account of the forest trees does not list Duffield ranch 
as a locality for Pinus sabiniana. The other species conform altitudinally, 
or one or two may have been collected a little below the ranch. 

Soon after the last date mentioned in the itinerary as given above, 
Bigelow returned to the eastern United States and reported to his com- 
manding officer in the War Department at Washington for filing his 
report of the expedition which is dated [but certainly not published in| 


1962 ] JEPSON: BIGELOW 183 


August, 1854. This report (in Pacific Railroad Reports, Vol. 4) consists 
first of a botanical narrative of the expedition from Napoleon on the Mis- 
sissippi River, by boat up the Arkansas River to Fort Smith, Arkansas, 
thence overland to Los Angeles and San Pedro (pp. 1-16) and second 
of a description of the forest trees of the route (pp. 17-21) with a special 
section on “Descriptions of Valuable or Remarkable California Forest 
Trees” (pp. 21-26). One of the most interesting contributions made by 
Bigelow to the Report is a large map giving a botanical profile in color 
of the forest trees from Fort Smith to San Pedro. He also collaborated 
with George Engelmann in the preparation of the paper entitled ‘“‘Descrip- 
tion of the Cactaceae” (pp. 27-58, pls. 1-24) in which many of the more 
commonly known cactus species of our desert regions were first described 
and named. 

In his honor, Asa Gray named the genus Bigelovia of the Compositae, 
a group now covered by other names. There were also dedicated to him 
Clematis bigelovu Torr., Aster bigelovii Gray, and Linosyris bigelovi 
Gray of New Mexico, but our commonest cholla, Opuntia bigelovii, a wide- 
spread cactus of the arid districts of southern California and adjacent 
areas, named for him by George Engelmann, will perhaps serve most 
widely to keep in mind a botanical traveler who was so fortunate as to 
view a considerable part of California while its native vegetation was still 
in nearly pristine freshness. 

In 1860 Dr. Bigelow made his home in Detroit where he was placed 
in charge of the meteorological work of the Northern and Northwestern 
Lakes Survey. Some years later he was appointed surgeon to the Marine 
Hospital in Detroit and Professor of Medical Botany and Materia Medica 
in the Medical College. He lived in Detroit the remainder of his life and 
died there July 18, 1878. 


LITERATURE CITED AND REFERENCES 


ATKINSON, W. B. 1878. The Physicians and Surgeons of the United States. C. Robson, 
Philadelphia, Pa. 


BIGELOW, JoHN Mitton, M. D. 1856. (1) General Description of the Botanical 
Character of the Country .... along the route Traversed, and (2) Description 
of Forest Trees. In Pacific Railroad Reports, Vol. 4, pp. 1-26. Dept. of War, 
Washington, D. C. 


ENGELMANN, GEorGE, M. D., and J. M. BicELow. 1856. Description of the Cactaceae. 
In Pacific Railroad Reports, Vol. 4, pp. 27-28, pls. 1-24. Dept. of War, Wash- 
ington, D.C. 

SARGENT, CHARLES SPRAGUE. 1890. The Silva of North America, Vol. 1, p. 88. Hough- 
ton, Mifflin & Co., Boston and New York. 

Torrey, JOHN. 1857. Description of the General Botanical Collections [Bigelow’s 
in large part]. In Pacific Railroad Reports, Vol. 4, pp. 59-182, pls. 1-25. Dept. 
of War, Washington, D. C. 

TorrEy, JoHN. 1859. Botany of the Mexican Boundary. Dept. of the Interior, Wash- 
ington, D. C. 


184 MADRONO [Vol. 16 


A SUBARBORESCENT NEW ERIODICTYON 
(HYDROPHYLLACEAE) FROM 
SAN LUIS OBISPO COUNTY, CALIFORNIA 


Puitire V. WELLS 


In June of 1960, the writer encountered colonies of an extremely 
tall yerba santa, measuring up to thirteen feet in height, on the sand- 
stone ridges of Indian Knob, four miles north of Pismo, San Luis 
Obispo County, California (fig. 1). In the characters of the inflorescence, 
there is a close resemblance to Eriodictyon californicum (H.& A.) Torr., 
but the strongly linear and revolute leaves and densely pubescent cap- 
sules suggest a relationship to the isolated E. capitatum Eastw., which 
possesses an inflorescence unique for the genus. The combination of 
characters presented by the San Luis Obispo County Eriodictyon clearly 
sets it apart from the rest of the genus. This was verified by comparison 
with collections at the California Academy of Sciences and the University 
of California, Berkeley. 


Eriodictyon altissimum P. V. Wells, sp. nov. Frutex glutinosus, 2—4 
m. altus; foliis linearibus, 6.0-9.0 cm. longis, 2-4 mm. latis, margine 
revolutis, supra glabris et glutinosis, infra albo-tomentosis; inflorescentia 
cymosa-paniculata, ramulis elongatis, 4.0-9.0 cm. longis, glutinosis, 
floribus secundis in ramulis cymosis; calycis segmentis anguste lanceo- 
latis, 2-3 mm. longis, glabris sed ciliatis et glutinosis; corollis purpura- 
scentibus, infundibuliformibus, 11-15 mm. longis, exteriore villosis, in- 
teriore glabris, staminibus inclusis, filamentis basi villosis; stylis pur- 
pureis, basi villosis; capsulis pubescentibus. 

Tall, straggling shrub to over 4 m. high, averaging ca. 2 m., with a 
trunk-like main stem up to 12 cm. in basal diameter; growth habit of 
some individuals excurrent, with falsely whorled lateral branches; suck- 
ers freely produced from stout rootstocks, probably serving for propaga- 
tion when the tops are destroyed by fire; bark of main stems smooth, 
grayish; branches long and slender, ascending; the branchlets glutinous; 
leaves alternate, with a tendency to be opposite below, sessile, narrowly 
linear, 6.0 -9.0 cm. long, 2-4 mm. wide, entire, strongly revolute, glabrous 
and somewhat glutinous above, densely white-tomentose beneath, with 
heavy, sweet aroma; inflorescence an open panicle of cymes; flowers on 
minutely bracteate ramules 1-3 mm. long, secund on the lax, glutinous 
branches of the inflorescence, the branches ranging from 4-9 cm. in 
length; sepals lance-linear, 2-3 mm. long, ciliate on the margins, other- 
wise glabrous but glutinous; corolla infundibuliform, 11-15 mm. long, 
the limb lavender and the tube pale lavender to whitish below, sparsely 
villous without, glabrous within; stamens unequal, the filaments villous 
on the basal half; styles 5-7 mm. long, lavender, sparsely hairy below; 
ovary ca. 2 mm. long, densely short-pubescent and glutinous; capsule 


1962 | WELLS: ERIODICTYON 185 


Fic. 1. Eriodictyon altissimum, showing excurrent growth 
habit present in some individuals. Summit of San Luis Range 
near Indian Knob, San Luis Obispo County. 


glutinous-pubescent, containing numerous polyhedral seeds, the seeds 
ca. 0.4 mm. long, brown, longitudinally finely striate and reticulate with 
cross-striations. 

Holotype. Sandstone ridges of Indian Knob, elevation 880 feet, four 
miles north of Pismo, San Luis Obispo County, California, P. V. Wells 75, 
June 30, 1960, OBI. Isotypes at CAS, GH, UC, US. 

Eriodictyon altissimum is apparently confined to shallow, sandy soils 
derived from siliceous sandstone (San Pablo group: upper Miocene) in 
the eastern part of the San Luis Range at elevations of 650 to 880 feet. 
It was not found on Franciscan rocks (feldspathic sandstone, radiolarian 
chert, diabase or serpentine), nor on siliceous Monterey shale, all of 
which outcrop at comparable elevations in the San Luis Range. The vege- 
tation on the siliceous sandstone to which £. altissimum is confined is 


186 MADRONO [Vol. 16 


largely chaparral interspersed with low woodlands of Quercus agrifolia 
Neé, and with one small stand of Pinus muricata D. Don. The chaparral 
matrix is dominated by Arctostaphylos pilosula Jeps. & Wies., with asso- 
ciated chamise, toyon, Ceanothus impressus Trel., C. cuneatus var. ramu- 
losus Greene, and the subligneous Salvia mellifera Greene. (Pinus muri- 
cata and Ceanothus impressus are conspicuous in the Lompoc endemism 
area which harbors the linear-leaved Eriodictyon capitatum. ) 

Eriodictyon altissimum, like other members of the genus, has a weedy 
or pioneer ecology. It is aggressive on roadsides, with numerous young 
plants invading such disturbed sites. The large production of minute 
seeds averaging 0.2 mg. in weight provides the necessary mobility. It is 
apparently a rapidly growing, short-lived shrub, often overtopping by 
five feet or more the even-statured young manzanitas dating from the 
last chaparral fire. Thrifty specimens with luxuriant foliage rarely occur 
in the chaparral, being mainly confined to road sides. By far the greatest 
number of individuals observed had a senescent appearance, open and 
straggling with sparse foliage confined to the tips of branches (fig. 1). 

Since this Ertodictyon combines characters of E. californicum and E. 
capttatum (or less possibly EF. angustifolium Nutt.), one might suppose it 
to be of recent hybrid origin. However, none of these species occurs in 
the San Luis Range; in fact, there are no previous records of linear-leaved 
yerba santas from San Luis Obispo County (R. F. Hoover, unpublished 
checklist). If hybridization should prove to be involved, the writer sees 
no reason why a large, apparently stable population evolved in this 
manner should be given the nomenclatural treatment ordinarily accorded 
to hybrids of sporadic and ephemeral character. 

Department of Biology 


New Mexico Highlands University 
Las Vegas, New Mexico 


A NEW SPECIES OF QUERCUS FROM 
BAJA CALIFORNIA, MEXICO 


CORNELIUS H. MULLER 


In a report upon his 1885 collection of the plants of Cedros Island, Baja 
California, Mexico, Greene (1888) made the following entry: 

“66. Quercus. _____. A merely shrubby species of the White Oak 
series; leaves small, spinose-toothed and persistent; midway up the 
canons.” Greene’s collections (with Geo. W. Dunn) were made in several 
canyons located on the northeast side of the island. Presumably a speci- 
men of the oak is preserved in his herbarium. In 1922 G. Dallas Hanna 
collected the same oak at the “north end” of Cedros Island. His specimen 
is preserved in the herbarium of the California Academy of Sciences. 

The plant first came to my notice in the form of a small flowering col- 
lection made on the island by A. L. Haines and G. O. Hale in 1939 and 


1962 | MULLER: QUERCUS 187 


submitted to me through the kindness of Dr. Mildred Mathias from the 
herbarium of the University of California at Los Angeles. Subsequent 
inquiry concerning additional material brought to light two collections on 
the mainland in the vicinity of San Vicente some 200 air miles northward 
from Cedros Island, one near Rancho San Antonio del Mar and the second 
south of San Vicente. This latter collection was mentioned by Epling and 
Robinson (1940) under the name, Quercus dumosa Nutt. 

In February, 1960, I visited the above two mainland localities as well 
as a stand in Canon del Rio San Ysidro where there is also a hitherto 
unreported grove of Pinus muricata. | am indebted to Mr. Richard Broder 
for aid in the field on this occasion. In June, 1960, I spent four days in 
the mountains in the southern half of Cedros Island, principally about 
Cerro Cedros, which reaches an elevation of 3950 feet.’ In August, 1960, I 
encountered typical stands of the species at 3000 feet elevation on the 
western foot slopes of the Sierra San Pedro Martir. This locality is char- 
acterized by a dry chaparral dominated by Adenostema fasciculatum with 
A. sparsifolium and Arctostaphylos sp. About the same time Dr. John M. 
Tucker kindly called to my attention a collection made in 1956 by Mr. 
John Thomas Howell in San Carlos Canyon above Agua Caliente de San 
Carlos (the village southeast of Ensenada—not the race track). 

Haines and Hale had reported the oak to occur at 1750 feet elevation 
(according well with Greene’s ‘‘midway up the cafions”); I found the 
main body of the Cedros Island population occurring between elevations 
of 3300 and 3940 feet on north- and west-facing slopes. Here it is associ- 
ated with Juniperus californica, Arctostaphylos bicolor, Rhus laurina, and 
Eriogonum fasciculatum. Adjacent south-facing slopes are dominated by 
Pachycormus discolor var. veatchiana, Franseria spp., and only scattered 
Juniperus californica. Very distinct lines separate the mesic and desert 
slopes. The San Vicente localities, on the other hand, all lie below 300 
feet and-are usually characterized by Pinus muricata, although at the San 
Antonio del Mar locality Pinus is lacking and the community is repre- 
sented only by Ribes viburnifolium, another common associate of the oak. 

In spite of a long history, this overlooked species is still sketchily known 
as to its geographic range. It is safe to assume that far fewer than half its 
localities are known and that still further unexpected extensions of range 
will eventually be discovered. It may be found with or without Pinus 
muricata and the common associates of that species, but on the mainland 
pine groves constitute promising indicators of the oak. 


1 Tn this undertaking I incurred extensive obligations for aid without which suc- 
cess would have been impossible: to the U. S. Fish and Wildlife Service for trans- 
portation aboard the M. S. Black Douglas; to the crew of the Black Douglas for 
unusual courtesy; to Sr. Francisco Amaya, manager of the Atun-Mex fish cannery 
on Cedros Island, for most helpful cooperation and facilities; to Sr. Eduardo Her- 
nandez-Bello of the Mexican fisheries laboratory at Mazatlan for smoothing the 
way with officials; and especially to Mr. Campbell Grant, my good companion on 
this and other ventures. 


188 MADRONO [Vol. 16 


An examination of sterile and flowering materials early suggested that 
the Cedros oak was not a “‘white oak” at all but rather was a member of 
the series Chrysolepidae in the subgenus Protobalanus, the ‘‘intermediate 
oaks” in which, except for its spinose-toothed leaves, it strongly suggested 
Q. vaccinifolia Kell. An examination of fallen cups and acorns in the San 
Vicente localities and, more recently, of attached fruit on Cedros Island 
confirmed this opinion. The presence of tomentum on the inner surface 
of the acorn shell and the characteristically swollen and puberulent bases 
of the cup scales are distinctly characters of the series Chrysolepidae of 
Protobalanus. 

In spite of the superficial resemblance of this species to Q. vaccintfolia, 
it is amply distinct in several basic but scarcely obvious characters. Its 
position in the subgenus Protobalanus is actually fully as anomalous as 
that of the four previously known species of that group. Each species 
seems to be a relic and the end point of its own ancient line within the 
subgenus. 

Quercus cedrosensis sp. nov. Arbor parva vel frutex; ramuli 1 mm. 
diametro, sparse vel dense stellato-pubescentes; folia sempervirentia, 
coriacea, 6-20 (35) mm. longa, 4-14 (20) mm. lata, integra vel dentata, 
dentis spinosis, ovata vel lanceolata, acuminata vel obtusata, basi rotun- 
data vel cordata, supra sparse pubescentia vel glabrata, nitida, subtus 
glaucescentia; venis utrinque 4—8, haud prominentibus; petioli 1.5—2.5 
mm. longi; fructus biennis, brevipedunculatus; cupula 5-6 mm. alta, 
7-12 mm. lata; glans 15-22 mm. longa, 6-10 mm. lata, angusto-ovoidea 
vel fusiformis, ad basim tantum cincta. 

Small trees to 15 ft. tall with a trunk 2 dm. in diameter with flaky gray 
bark or, on windswept sites, the trunk decumbent forming a large shrub 2 
or 3 m. high and 6 or 8 m. broad, or the shrub quite prostrate and forming 
a mat as little as 2 dm. high, the stumps sprouting vigorously following 
fire and prostrate branches rooting freely or the underground parts 
rhizomatous; twigs about 1 mm. thick, the internodes very short, smooth, 
brown becoming dark gray, sparsely or densely pubescent with short 
stellate hairs which persist into the second year; buds about 1 mm. long, 
broadly ovoid or subrotund, light brown, sparsely pubescent; stipules 


EXPLANATION OF FIGURES 1-5 


Fics. 1-3, 5. Quercus cedrosensis: 1, The type collection from Cedros Island, 
Muller 10775 (x 1.4); 2, a common leaf form from near San Vicente on the main- 
land, Muller 10724 (xX 1.4); 3, a typical stump sprout in the Cedros Island popu- 
lation, Muller 16777 (x 1.4); 5, a typically toothed specimen from Cedros Island, 
Muller 10772 (x 10)-—note the extreme elongation of the spinescent teeth and 
the breadth of the sclerenchymatous sheaths of the veinlets, particularly on the 
lower surface, which almost occlude the chlorenchymatous alveolae. 


Fic. 4. Quercus vaccinifolia: A typically toothed specimen from Siskiyou County, 
California, Muller 9667 (x 10)—in this lower surface view note the very short tips 
of the teeth and the moderately sclerenchymatous veinlets bordering large chlo- 
renchymatous alveolae. 


MULLER: QUERCUS 


Fics. 1-3, 5. Quercus cedrosensis. 


Fic. 4. Q. vaccinifolia. 


190 MADRONO [Vol. 16 


2 to 3 mm. long, ligulate-spathulate, the apical end quite thin, persistent 
the second year; leaves evergreen, thick and chartaceous, persisting 2 or 
3 seasons, densely crowded on the short twigs, 6 to 20 (35) mm. long, 4 
to 14 (20) mm. broad, flat or sometimes distinctly concave beneath, entire 
or irregularly few-toothed or 6- to 8-toothed on each side, the teeth 
elongate and spinescent (consisting of 1 to 1.5 mm. of sclerenchymatous 
tissue extending beyond the chlorenchyma), the blade ovate or lanceolate, 
sometimes oblong or elliptic to subrotund, basally rounded or sometimes 
cordate, apically acute or sometimes broadly rounded, characteristically 
spinescent-tipped or the spine rarely lacking, upper surface glabrous or 
very sparsely stellate-pubescent at the base of the midrib, glossy green, 
lower surface glabrous or the midrib minutely strigose, glaucous and waxy, 
marked by white dots (juvenile leaves sparsely pubescent with stellate 
and simple hairs, especially on the upper surface about the midrib, the 
blade heavily anthocyanous on the upper surface, the lower surface 
green) ; veins 4 to 8 on each side, very inconspicuous or slightly raised on 
the lower surface, both veins and reticulum highly sclerenchymatous and 
white, almost eclipsing the chlorenchymatous alveolae (the diameter of 
each green area little greater than the width of the adjacent white veinlet) ; 
petioles about 1.5 to 2.5 mm. long, pruinose, glabrous or sparsely pubescent 
with small stellate hairs; staminate catkins about 10 to 15 mm. long, 
sparsely flowered, the rachis somewhat stellate-pubescent, the filaments 
inserted in a tuft of white pubescence on the receptacle and scarcely longer 
than the 4 to 8 glabrous red anthers, these barely exserted from the ciliate 
red perianth; pistillate catkins 4 to 8 mm. long, 1- to 3- flowered on a 
sparsely pubescent rachis; fruit biennial, maturing in July, simultaneously 
with or following the next flowering, solitary or paired, subsessile or on 
a peduncle to 10 mm. long; cups 7 to 12 mm. broad, 5 to 6 mm. high, cup- 
shaped, the scales very broad and thickened basally, appearing as though 
fused, green and densely silver-puberulent, the thin brown apices elon- 
gate, appressed and ciliate; acorns 15 to 22 mm. long, 6 to 10 mm. broad, 
very narrowly ovoid to fusiform, acute at apex, glabrous and brown except 
the silvery puberulent apical quarter, less than one-quarter enclosed at 
the base. 

Range. Baja California, Mexico, on Cedros Island and from the vicinity 
of San Vicente inland and northward on the peninsula. 

Specimens examined: MEXICO, Baja Catirornia: Isla Cedros: north end, 
August 9, 1922, G. D. Hanna s.n. (CAS, SBC-MU?) ; north slope of Cerros [Cedros] 
Peak, elev. 1750 ft., March 9, 1939, A. L. Haines & G. O. Hale 969 (UCLA, SBC-MU) ; 
elev. 1700 ft., Haines & Hale 970 (SBC-MU); in chaparral on north slope of Man- 
zanita Peak [north end of island], elev. 2700 ft., February 6, 1939, Haines & Hale 


935 (SBC-MU); “Cerros Mountains,” elev. 3900 ft., February 15, 1939, Haines & 
IIale sn. (SBC-MU); head of Cafion de Calabasas, southwest slopes of Cerro 


2 The abbreviations of herbaria are those of Lanjouw and Stafleu (1959) ; 
SBC-MU refers to my private collection of Quercus on deposit at the University 
of California, Santa Barbara. 


1962 | MULLER: QUERCUS 191 


Cedros, ca. 3600 ft. elev., June 16, 1960, C. H. Muller 10771-10777 (SBC-MU) [of 
which no. 10775 is the type (SBC herbarium no. 8766, the holotype, and widely 
distributed) ]; June 17, 1960, Muller 10799-10803 (SBC-MU); heavily wooded 
north-facing canyon wall near waterway on northwest slope of Cerro Cedros, 
ca. 2900 ft. elev., June 17, 1960, Muller 10791-10794 (SBC-MU) ; northwest slope 
of summit of Cerro Cedros ca. 3940 ft. elev., June 19, 1960, Muller 10817-10818 
(SBC-MU). Municipio de Ensenada: Cafion San Carlos above Agua Caliente, March 
18, 1956, J. T. Howell 31102 (DAV); south side of San Antonio Canyon about 
2 miles inland from San Antonio del Mar (Johnson’s Ranch), September 8, 1930, 
I. L. Wiggins & D. Demaree 4762 (DS, SBC-MU); Pine canyon 6 miles south of 
San Vicente, April 11, 1936, C. Epling & W. Stewart s.n. (UCLA) ; northwest-facing 
slope of Cerro Colorado in Canon de Rio San Ysidro, 6.6 miles southwest of San 
Vicente, in and above grove of Pinus muricata, ca. 300 ft. elev., February 14, 1960, 
Muller 10723-10727 (SBC-MU) ; near arroyos on north-facing slope in Cafion de los 
Pinitos, 8 miles south of San Vicente (the Epling and Stewart locality for which 
this is a mileage correction), ca. 250 ft. elev., February 15, 1960, Muller 10729-10732 
(SBC-MU); southeast side of Canon San Antonio, 2.7 miles north-northeast of 
Rancho San Antonio del Mar (Johnson’s Ranch) on road to Rancho Cerro Blanco 
(Rancho Guzman) and San Vicente (the Wiggins and Demaree locality for which 
this is a mileage correction), February 15, 1960, Muller 10733-10736 (SBC-MU) ; 
4 miles below Socorro, 7 miles above San José (Meling Ranch) on western slope of 
Sierra San Pedro Martir, elev. 3000 ft., August 16, 1960, Muller 10888-10890 
(SBC-MU) ; August 22, 12960, Muller 10937 (SBC-MU). 


Quercus cedrosensis is apparently rather remotely related to Q. vac- 
cinifolia, a species of high elevations in the Sierra Nevada of California 
and descending below 5000 feet only in northern California and Oregon. 
The Mexican species is distinguished from Q. vaccinifolia by its tree habit 
(although it matures as a shrub in windswept situations and elsewhere 
on the mainland), its generally maritime and frequently low elevation 
distribution, the heavily sclerenchymatous nature of its leaf reticulum, 
its spinescent teeth whenever teeth occur, and its elongate, acute acorns. 
Further, the brevity of its internodes produces a marked crowding of the 
leaves which is not approached by Q. vaccinifolia and the uniform occur- 
rence of anthocyanin in juvenile leaves is totally lacking in that species. 
Despite this rather lengthy list of significant differences, the two species 
are superficially quite similar and might, in the instance of a few speci- 
mens, prove difficult to distinguish without intimate acquaintance. 

The more characteristic populations of Q. cedrosensis are found on 
insular and inland sites of relatively high elevation (about 3000 feet) ; 
the less typical occur on the mainland coast at low elevation. In general, 
the broader leaf forms with more rounded apices and less frequently 
spinescent teeth are more common in the coastal San Vicente populations 
of Q. cedrosensis than in the insular and inland populations, suggesting 
that some low degree of introgression of the San Vicente populations is 
being reflected. It is impossible at this time to demonstrate the source of 
such an influence. The upper levels of the Sierra San Pedro Martir have 
been searched diligently for Q. vaccinifolia with the view of explaining 
some of the polymorphy of Q. cedrosensis about San Vicente. In addition 
to finding Q. cedrosensis at 3000 feet, three additional members of the 


192 MADRONO | Vol. 16 


Chrysolepidae were encountered as follows:Q. palmeri Engelm. at 2800 
to 4000 feet. O. chrysole pis Liebm. at 5500 to 7500 feet, and an anomalous 
entity suggesting QO. chrysolepis at 7000 to 9650 feet. With the possible 
exception of Q. chrysolepis, these San Pedro Martir species would not be 
expected to have contributed the aberrations of the San Vicente popula- 
tions, and the presence in the San Pedro Martir of typical O. cedrosensis 
makes even this extremely unlikely. 

University of California, 

Santa Barbara, 
University, California 
LITERATURE CITED 


GREENE, E. L. 1888. The botany of Cedros Island. Pittonia 1: 194-208. 
Eprinc, C., and W. Rosison. 1940. Pinus muricata and Cupressus forbesii in Baja 
California. Madrono 5: 248-250. 
Lanyouw, J., and F. A. StaFrevu. 1959. The herbaria of the world. Index Herbari- 
orum, part I, ed. 4. Regnum Veg. 15. 


PARASITISM IN PEDICULARIS ! 
ELIZABETH F. SPRAGUE 


The parasite-host relationship for many European species of Pedicu- 
laris has been well-documented by Wettstein (1891), Boeshore (1920), 
Hayek and Hegi (1918), and others. Such parasitism accounts for the 
difficulty in culture noted by such workers as Don (1838) and Tsoong 
(1955), although a few species have been cultivated and a few are sold 
for ornamental value. Tsoong (loc. cit.) states that P. fletcheriana 
Tsoong “may be easily raised from seed” and that at Perthshire, Eng- 
land, it established itself and freely reproduced. This indicates that at 
least some species under gven conditions may be saprophytic, mycor- 
rhizal, or completely autotrophic. Sperlich (1902) found some species of 
Pedicularis to be both parasitic and saprophytic, with haustoria of the 
same plant attached to both dead organic matter and living roots. Cer- 
tainly many of the meadow-dwelling species are quite opportunistic with 
regard to available host plants. In Europe, P. sylvatica L., and, in Amer- 
ica, P. canadensis L. and P. lanceolata Michx. are regularly sold and 
cultivated without apparent hosts. They probably thrive as saprophytes 


1 This paper is adapted from a portion of a doctoral dissertation prepared at the 
Rancho Santa Ana Botanic Garden and the Claremont University College, Claremont, 
California. I wish to acknowledge the assistance of Doctors Verne Grant, Sherwin 
Carlquist, and Philip A. Munz. The illustrations were prepared with the assistance 
of Messrs. William Klein and C. Dodson. The research was partially financed by 
two grants from the Claremont University College, Claremont, California, and a 
fellowship from the Southern Fellowships Fund, Chapel Hill, North Carolina. A 
grant from the University Center, Richmond, Virginia, assisted with typing and 
photographic expenses. 


1962 | SPRAGUE: PEDICULARIS 193 


because a soil rich in humus is required, although even then they may 
be difficult to maintain. Attempts to grow thirteen plants of P. canadensis 
at Rancho Santa Ana Botanic Garden, Claremont, California, were not 
successful either in the greenhouse or in deep humus under Quercus 
agrifolia Neé. Only four to six small leaves per plant were produced, 
and only five plants persisted to the second year. 


Attention was focused on the parasitic nature of seven California 
species when an attempt was made to cultivate them for experimental 
purposes. Plants of Pedicularis densiflora, P. semibarbata, P. groen- 
landica, P. attollens, and P. dudleyi were transplanted to clay pots and 
grown in the glasshouse at Rancho Santa Ana Botanic Garden, Clare- 
mont, California. None bloomed and even those which were accompanied 
by associated plants produced only a very few small leaves the second 
season. A single plant of P. dudleyi was grown successfully for the two- 
year period. Its native habitat, the redwood forest, was probably more 
successfully simulated under greenhouse conditions. 

No species of Pedicularis is recorded as requiring exact host specificity 
even as to genus, but some do “show obvious preference for definite 
species” (Hayek and Hegi, 1918, page 112, the quotation a translation 
from the German). The majority of European species are meadow- 
dwelling and hence the hosts listed are various grasses, sedges and wil- 
lows; however, Hayek and Hegi note that P. recutita L. is found usually 
on Deschampsia caespitosa (L.) Beauv. and Pedicularts verticillata L. on 
Sesleria caerulaea Scop. For the California species examined in this study, 
field observation and laboratory verification indicated the host relation- 
ships shown in Table 1. The alpine and meadow species of the Sierra 
Nevada, Pedicularis attollens, P. groenlandica and P. crenulata, show 
typical facultative association with various meadow plants. A fourth 
Sierran species, P. racemosa, grows in deep humus associated with A bies 
concolor and Pinus monticola; no haustorial connections were identified. 
Pedicularis dudleyi is associated with such redwood-forest understory 
plants as Vaccinium ovatum and Lithocarpus densiflora; roots were not 
observed. 

However facultative the parasite-host relationship may appear to be 
in most species, observations on Pedicularis densiflora and P. semibarbata 
show restrictions unlike those reported for other species. These two spe- 
cies parasitize principally roots of trees or woody shrubs. In addition, 
there is evidence that the local populations of P. densiflora are physiologi- 
cally distinct. In southern California, P. densiflora is associated pri- 
marily with Adenostoma fasciculatum, in the Santa Lucia Mountains, 
some populations are associated with Pinus coulteri and Arbutus men- 
ziesi, others with Adenostoma and Arctostaphylos; in the San Francisco 
area, Arbutus menziesti and Quercus kelloggi are the conspicuous hosts 
except on the top of Mount Diablo, where Pedicularis grows in a pure 
stand of Adenostoma. In the two areas on Mount Diablo and in the 


194 


MADRONO 


LVol. 16 


TABLE 1. Hosts at Vartous LOCALITIES OF OBSERVATION 
FOR FIVE SPECIES OF PEDICULARIS! 


PEDICULARIS 
SPECIES 


P. densiflora 
subsp. densiflora 
Benth. 


P. densiflora subsp. 
aurantiaca (Benth.) 
E. F. Sprague 


(Probable host—P. 


LOCALITY 


Del Mar mesa, 
San Diego County 


Cobal Canyon, San 
Gabriel Mountains, 

Los Angeles County 
Topanga Canyon, Santa 
Monica Mountains, 

Los Angeles County 
Refugio Canyon burn, 
San Marcos Pass, 

Santa Barbara County 
Klau Mine on serpentine, 
Adelaide, 

San Luis Obispo County 
Marquart Ranch, 
Cambria-Adelaide Road, 
Santa Lucia Range, 
San Luis Obispo County 
7X Ranch pass, on 
serpentine, Santa Rita 
Canyon, 

Santa Lucia Range, 

San Luis Obispo County 
La Honda, 

Santa Cruz Mountains, 
San Mateo County 
Phoenix Lake, 

Marin County 

Mount Diablo, Toyon 
Road, Rocky Point, 
Contra Costa County 


Mount Diablo, below 
Rocky Point, 

Contra Costa County 
Jackson County, Oregon” 
Lake Almanor, 

Piumas County 


Butte Meadows, 
Butte County 


Near Viola, Shasta County 
Near Mineral, 
Tehama County 


Confirmed host—C.) 


Host 


(C) Adenostoma fasciculatum 


H.& A. 


(P) Arctostaphylos glandulosa 


var. crassifolia Jepson 


(C) Adenostoma fasciculatum 


(C) Adenostoma fasciculatum 


(C) Adenostoma fasciculatum 
(P) Arctostaphylos glandulosa 


Eastw. 


(P) Adenostoma fasciculatuim 


(P) Arbutus menziesii Pursh. 


(?) Pinus coultert Don. 


(C) Adenostoma fasciculatum 


(?) Quercus dumosa Nutt. 
(?) Arctostaphylos sp. 


(2?) Rhus diversiloba T.& G. 
(?) Diplacus puniceus Nutt. 


(P) Arbutus menziesiz 
(P) Rhus diversiloba 


(P) Diplacus aurantiacus Jepson 


(P) Arbutus menziesii 


(P) Quercus kelloggii Newb. 
(C) Adenostoma fasciculatum 


(P) Arbutus menziesii 

(P) Quercus kelloggii 

(P) Pinus sabiniana Doug}. 
(P) Rhus diversiloba 

(P) Ceanothus sp. 


(P) Arbutus menziesiz 


(C) Pinus ponderosa Dougl. 


(P) Pinus jeffreyi 
Grev. & Balf. 

(P) Abies concolor 
Lindl. & Gord. 

(?) Pinus ponderosa 


(?) Pinus ponderosa 


1962 | 


PEDICULARIS 
SPECIES 


SPRAGUE: PEDICULARIS 195 


TABLE 1, continued. 


LOCALITY 


Host 


P.attollens Gray 


P. groenlandica 
IRGtz: 


P. crenulata 
Benth. 


P. dudleyi 
Elmer 


P. racemosa 
Dougl. 

P. semibarbata 
Gray 


Tioga Pass, Mono County 
Slate Creek, 

Middle Ridge, 

Mono County 

Slate Creek, 
Timberline Station, 
Mono County 

Echo Pass (Phillips 
and one mile north), 
El Dorado County 
Fallen Leaf Meadows, 
El Dorado County 


Tioga Pass and 

Tioga Lake, 

Mono County 

Slate Ridge, Middle Ridge, 
Mono County 

Slate Creek, Timberline 
Station, Mono County 
Sonora Pass, 
Tuolumne County 
Norden, 

Placer County 


Convict Creek, 
Mono County 


Portola State Park, 
Santa Cruz County 


Rainbow Tavern, 
Highway 40, Placer County 


Above Lake Arrowhead, 
San Bernardino Mountains 
San Bernardino County 
Mount San Gorgonio, at 
foot of trail to peak, 

San Bernardino County 
Wrightwood, San 

Gabriel Mountains, 

San Bernardino County 
Mount Pinos, Kern County 
Echo Lake, 

Eldorado County 
Norden, 

Placer County 


(P) Phleum alpinum L. 
(C) Carex heteroneura 
W. Boott. 


(C) Carex heteroneura 


(C) Trifolium monanthum Gray 
(P) Phleum alpinum 


(P) Carex sp. 


(C) Carex helleri Mkze. 

(C) Deschampsia caespitosa 
(P) Carex fissuricola Mkze. 
(P) Deschampsia caespitosa 


(C) Poa sp. 
(P) Poa sp. 
(P) Carex sp. 


(C) Deschanpsia sp. 

(C) Trifolium monanthum 

(P) Poa sp. 

(P) Vaccinium ovatum Pursh 

(P) Lithocarpus densiflora 
(H.& A.) Rehd. 

(P) Ceanothus thyrsiflorus Esch. 

(P) Abies concolor 

(?) Pinus monticola Don. 

(C) Pinus ponderosa 


(C) Pinus ponderosa 


(C) Pinus ponderosa 

(?) Poa scabrella (Thurb.) 
Benth. 

(C) Pinus ponderosa 

(C) Arctostaphylos patula 
Greene 

(?) Abies concolor 


1 All localities are from California except one, as stated, from Oregon. 
2 Hitchcock 64988, June, 1931 (RSA). 


196 MADRONO [Vol. 16 


Santa Lucia Mountains, where two distinct populations associated with 
different hosts were observed within potential breeding range, the plants 
in each area appeared distinctive, although the difference could not be 
defined. In addition, Pedicularis densiflora subsp. aurantiaca seemed to 
be very nearly host specific; at least, haustoria were never found on 
roots of species other than Pinus ponderosa. Pedicularis semibarbata, 
likewise, was almost exclusively on Pinus ponderosa; at Echo Lake, El 
Dorado County, in an old burn, the nearest yellow pine was 80 feet away, 
but the Pedicularis was attached to its far-reaching roots. 

Pedicularis densiflora and P. semibarbata have undergone an extreme 
divergence from a habitat of moist meadows and cool mountains such 
as that most species occupy to an almost arid habitat. The water re- 
quirements of such species must be critical. Under such conditions, woody 
shrubs and trees would provide both the most adequate supply through- 
out the growing season and an adjacent perennial root which could be 
annually tapped. 

In addition to these close vascular plant associations, there seems to 
be some evidence that under more favorable climatic conditions certain 
species may be largely saprophytic or mycorrhizal rather than parasitic. 
On plants of P. densiflora at Phoenix Lake, Marin County, where humus 
and climate provide more continuously available moisture than in many 
areas within their range, very few small haustoria and no actual connec- 
tions with adjacent plants were observed on six uprooted individuals. 
In a low drainage area near Viola, Shasta County, no haustoria were 
observed on P. densiflora subsp. aurantiaca. In both places, abundant 
mycelia in the rhizosphere suggested that mycorrhizae many function. 
Mycorrhizae may be important where species are largely saprophytic, 
also. The only material examined microscopically was taken from south- 
ern populations and showed no evidence of fungal elements. 


It was generally agreed among earlier investigators that members of 
the tribe to which Pedicularis belongs (Euphrasieae of Pennell, 1935; 
Rhinantheae of Wettstein, 1891) “take much from the earth” (Pitra, 
1861, page 66, the quotation a translation from the German) and little 
from their hosts. Kerner (1895) observed the close contact of the epi- 
dermal cells with the humus. Hayek and Hegi (1918) reported that some 
species take organic materials to the detriment ci the host. The impor- 
tance of the host to the parasite is evidently due to a “disproportion 
between the parasite’s water requirement and powers of the root to 
satisfy them” (Skene, 1924). Kostytschew (1922) showed that the cut 
shoots of Euphrasia absorbed water two times as fast, and those of 
Melampyrum ten times as fast, as the uncut shoots absorbed through 
their own roots. A similar distinction was found in the water absorption 
capacities of cut versus uncut shoots in Pedicularis densiflora, P. semt- 
barbata, and P. groenlandica. According to Tubeuf (1923, page 564, 
referring to unpublished work of Senn and Hagler), Euphrasia stricta 


1962 | SPRAGUE: PEDICULARIS 197 


Fics. 1-5. 1,2, haustoria of Pedicularis densiflora on Adenostoma fasciculatum, 
Del Mar, San Diego County; 3-5, haustoria of Pedicularis semibarbata on Pinus 
ponderosa: 3, Wrightwood, San Bernardino County; 4, 5, Mount Pinos, Kern County 
(note wrinkling of the large contractile root). (Fig. 1, X 54; 2, K %; 3, iy; 
Oe Ges, 2) 


Host. and Pedicularis sylvatica exhibit osmotic pressures significantly 
higher than those of their hosts. Although the loss of water to the parasite 
must put a considerable strain upon the absorbing system of the host, 
there are few references in the literature to any deleterious effects upon 
the host. 

The nature of the haustorium was discussed and illustrated by Wett- 
stein (1891), who considered it a reduced lateral root produced in the 


198 MADRONO [Vol. 16 


Fics. 6-12. 6,7, haustoria of Pedicularis attollens on Poa sp.; 8, median section 
through haustorium of Pedicularis semibarbata on Pinus ponderosa; 9, section 
through haustorial primordium of Pedicularis attollens; 10, longitudinal section of 
old haustorial connections of Pedicularis attollens on Poa sp.; 11, section of haus- 
torium of Pedicularis semibarbata on Pinus ponderosa; 12, section of haustorium 
of Pedicularis densiflora on Adenostoma fasciculatum. (Fig. 6, X %; 7, X1%;3 
R, x 17: 9) 255 10, 22. 11, Se 15: 12, Kx 15) 


spring. Assimilation from the host continues through the summer, then 
the haustorial connection weakens. With resorption of the organic union 
with the host (fig. 10), the haustorium serves as a storage organ. Accord- 
ing to Maybrook (1917), Leclerc du Sablon studied the haustoria in 


1962 | SPRAGUE: PEDICULARIS 199 


1886 and concluded that they are exogenous in origin in both Melam- 
pyrum and Pedicularis, arising from peripheral parenchyma which is 
stimulated by contact with the host to multiply and by ordinary elonga- 
tion penetrate the host, destroying the forepart and perforating the tissues 
by chemical absorption. Maybrook’s work on P. vulgaris Tournef. led 
him to agree with these findings. The position of haustoria on rootlets 
of P. groenlandica and P. attollens (figs. 6, 7) would indicate that con- 
tact with the host stimulates production of haustoria. The present obser- 
vations, however, agree with those of Wettstein (op. cit.), namely, that 
the haustoria are modified branch roots, endogenous in origin, and usually 
annual, as indeed they would have to be considering the contractile nature 
of the principal fleshy root system. However, observations on the very 
large haustoria of P. densiflora indicated they were the result of two or 
more years’ growth. 

Haustoria on all the species have a similar appearance; they are pale, 
fleshy lumps, usually occurring on the smaller branch roots. They vary 
in size with the species, with lesser variation between individuals of a 
given species. Large connections over 1 cm. in diameter of P. densiflora 
on Adenostoma (figs. 1, 2) are not frequent; they were found close to 
the ‘“‘trunks” of the host and in the drier localities (Topanga Canyon 
and Del Mar mesa, table 1). The largest haustoria of Pedicularis densi- 
jlora subsp. aurantiaca on Pinus were 7 mm. in diameter. The haustoria 
of Pedicularis semibarbata (figs. 3-5, 11) were mostly small in com- 
parison, but three of 4, 5, and 6 mm. were measured. Most haustoria 
observed on other species were very small, 1 to 3 mm. or less (figs. 6, 7), 
but even these small ones enclosed a large portion of the rootlets to which 
they were attached. 

In transection the mature haustorium exhibits an outer zone of thick- 
walled tissue, lacunar collenchyma, then a wide cylinder of thick-walled 
parenchyma filled with starch and other material. The vascular cylinder 
consists of a comparatively wide outer phloem band and an inner core 
of protoxylem and metaxylem tracheids and vessels, together with con- 
siderable xylem parenchyma (figs. 8-12). A mass of short vessel elements 
arranged randomly intermingle with those of the host (figs. 11, 12), so 
that a most intimate and effective contact is made. 

Sperlich (1902), in his examination of Pedicularis and related genera, 
frequently found tracheids wanting when the haustoria were saprophytic. 
He also identified storage products in addition to starch, such as albumi- 
noid crystals, amylodextrin, phosphoric acid and nitrates, varying with 
the species and seasons. Examination of the present species under con- 
sideration showed starch and also large quantities of other products 
which had the appearance of albuminous material. 

How early the haustorial attachment must be made probably depends 
on the vigor of the seedling as well as on the amount of soil moisture 
available. We do not know whether annual species are more or less de- 


200 MADRONO LVol. 16 


pendent upon host plants than perennials are. Neither Prain (1891), who 
listed fourteen annual species in India, nor others have commented on 
this. The fibrous roots of P. groenlandica and P. attollens appear to have 
a greater number of haustoria than other California species observed; 
this may reflect a specific need for more food and water (although the 
former often grows in running water) from the host plant, or it may be 
that the smaller haustoria are less effective than the larger ones (e.g., those 
of P. densiflora). Also, this greater number may be an artifact of prepara- 
tion; one can wash out the mass of P. groenlandica rootlets in a piece 
of sod more easily than one can dig extensive areas in the sun-baked 
hardpan of a chaparral-covered slope to obtain the entire root system 
of P. densiflora. 

Seeds of P. densiflora, P. semibarbata, P. groenlandica, and P. attol- 
lens were germinated in loam. The seedlings were transplanted to humus, 
or in the case of P. densiflora and P. groenlandica, to pots containing 
appropriate host plants. None of these became established nor did the 
roots develop macroscopic haustoria. Lack of success in establishing seed- 
lings and in transplanting these species was probably due in part to their 
parasitic nature and to the lack of adequate haustorial connections or 
of an appropriate host. 


Sweet Briar College 
Sweet Briar, Virginia 


LITERATURE CITED 


BoesHore, I. 1920. The morphological continuity of Scrophulariaceae and Oroban- 
chaceae. Contr. Bot. Lab. Univ. Penn. 5:139-177. 

Don, G. 1838. A general history of the dichlamydeous plants. Vol. 4. Londan. 

Hayek, A., von, and G. HEctI. 1918. Dicotyledones [Scrophulariaceae to Compositae | 
V. Teil) in Hegi, G. Illustrierte Flora von Mittel-Europa. 6(1):1-544. Munchen. 
Munchen 

KERNER, A. J. 1895. The natural history of plants. Vol. 1. Ed. F. W. Oliver, 
Blackie & Son, Ltd. London. 

KostyTscHEW, S. 1922. Uber die Ernahrung der griinen Halbschmarotzer. Ber. 
Deutsch. Bot. Ges. 40:273-279. 

Maysrook, A. C. 1917. On the haustoria of Pedicularis vulgaris Tournef. Ann. 
Bot. 31:499-511. 

PENNELL, F. 1935. The Scrophulariaceae of eastern temperate North America. Proc. 
Acad. Phila. Monog. No. 1. 

Pitra, A. 1861. Uber die Anheftungsweise einiger phanerogamen Parasiten an ihre 
Nahrpflanzen. Bot. Zeit. 19:61-67. 

Prain, D. 1891. The species of Pedicularis of the Indian empire and its frontiers. 
Ann. Roy. Bot. Gard. Calcutta, 3:1-196. 

SKENE, M. 1924. The biology of flowering plants. Sidgwick and Jackson, Ltd. 
London. 

SPERLICH, A. 1902. Parasitism in Rhinanthaceae. Beih. Bot. Centr. 11(7) :437-485. 

Tsoonc, P. C. 1955. New Himalayan species of Pedicularis. Bull. Brit. Mus. Bot. 
2:1-34. 

TusBeur, K. F., von. 1923. Monographie der Mistel. Munich. 

WETTSTEIN, R., von. 1891 and 1893. Scrophulariaceae in Engler and Prantl, Die 
naturlichen Pflanzenfamilien IV (3b) :39-107. 


1962 | CUMMINS & BAXTER: UREDINALIS 201 


NOMENCLATURE, LIFE HISTORIES, AND RECORDS OF 
NORTH AMERICAN UREDINALES! 


GEORGE B. CUMMINS AND JOHN W. BAXTER? 


NOMENCLATURAL NOTES 

1. PUCCINIA AGRIMONIAE (Arth.) Arth. Manual of Rusts in U.S. and 
Canada, p. 295. 1934. The type specimen consists of leaves of A grimonia 
pubescens Wallr. bearing uredia of Pucciniastrum agrimoniae ( Diet.) 
Tranz. and unattached teliospores that are identical with those of Puccinia 
lateripes Berk. & Rav., Grev. 3:52. 1874. The contaminant teliospores 
probably are from infected Ruellia strepens L. which Bartholomew col- 
lected at the same place and time (Sumner, Missouri, 7 Oct. 1907, F. 
Columb. No. 2667). Therefore, Puccinia agrimoniae falls into synonymy 
under Puccinia lateripes. 

2. PUCCINIA BOUVARDIAE Griff., Bull. Torrey Club 20:297. 1902. Puc- 
cinia anisacanthi Diet. & Holw., Bot. Gaz. 31:329. 1901. Reidentification 
of the host plant of the type of P. bouvardiae as Anisacanthus thurberi 
(Torr.) Gray instead of Bouvardia triphylla Salisb. made clear the rela- 
tionship of these two rusts which are recorded from southern Arizona 
and Mexico. Puccinia bouvardiae falls into synonymy under P. ani- 
sacanthi, 

3. Puccinia eumacrospora Cumm. nom. nov. Puccinia macrospora 
Arth. Mycologia 1:244. 1909; not Puccinia macrospora (Lk.) Spreng. 
Syst. 4:569. 1827. 

4. PUCCINIA XANTHIIFOLIAE Ell. & Ev., Jour. Myc. 6:120. 1891. P. 
helianthi Schw., Schr. Nat. Ges. Leipzig 1:73. 1822. That these two 
entities are the same was demonstrated by Baxter (1958), who success- 
fully inoculated seedlings of Helianthus annuus L., grown in the green- 
house, with urediospores from Jva xanthifolia Nutt. collected near 
Guernsey, Wyoming, in 1957. In 1960, urediospores from Helianthus 
annuus collected by Baxter near Greeley, Colorado, infected Jva xanthi- 


folia at Milwaukee, Wisconsin. Puccinia xanthiifoliae falls into synonymy 
under P. helianthi. 


LirE History STUDIES 
PUCCINIA ESCLAVENSIS Diet. & Holw. Aeciospores of Aecidium mira- 
bilis Diet. & Holw. on Mirabilis longiflora L., produced uredia and telia 
of Puccinia esclavensis on Panicum bulbosum H. B. K. ina field inocula- 
tion by Baxter near Portal, Arizona, August, 1960. In May, 1961, over- 
: 1 Journal Paper No. 1707, of the Purdue University Agricultural Experiment 
tation. 


~ The first author acknowledges Grants-in-Aid from the Society of the Sigma Xi 
and the Purdue Research Foundation and the privilege of working at the Jackson 
Hole Biological Research Station, Dr. L. Floyd Clarke, Director; the second author 
acknowledges a Grant-in-Aid from the National Science Foundation and the use 
of facilities of the Southwestern Research Station, Dr. Mont L. Cazier, Director. 


202 MADRONO [Vol. 16 


wintered teliospores were used in greenhouse inoculations of Mirabilis 
jalapa L., producing spermagonia and aecia. 


NEw REcorRDS 


1. AECIDIUM BOUVARDIAE Diet. & Holw. On Bouvardia glaberrima 
Engelm. near Southwestern Research Station, Portal, Cochise County, 
Arizona, 16 August, 1960, Baxter (PUR); Garden Canyon, Huachuca 
Mountains, Cochise County, Arizona, 5 September, 1959, 10 September, 
1960, Goodding 239-59, 266-60 (PUR). These are the first records of 
the fungus from the United States. The species is heteroecious. In 1961, 
Cummins noted intimate association with rusted Leptochloa dubia (H. 
B. K.) Nees in the Santa Rita and Chiricahua Mountains, Arizona, and 
used aeciospores successfully to infect L. dubia in a field inoculation con- 
ducted at the Southwestern Research Station near Portal. The fungus is 
a species of Puccinia, as yet unidentified. 

2, AECIDIUM CHAMAECRISTAE Arth. On Cassia fasciculata Michx., 
Ames, Iowa, 5 June, 1960, Baxter (PUR). The species was known before 
only from Kansas and Nebraska. 

3. BUBAKIA MEXICANA Arth. On Croton sp., Garner State Park, Uvalde 
County, Texas, 26 June, 1961, Miller (PUR). This is the first record of 
this rust from the United States. 

4. MELAMPsoRA ARCTICA Rostr. On Salix anglorum Cham., Breccia 
Peak, above Togwotee Pass, Wyoming, 29 August, 1960, Cummins 60-98 
(PUR). This species has not been found previously in Wyoming and 
only rarely in the United States. The site is in alpine tundra. 

5. PHAKOPSORA CROTALARIAE (Diet.) Arth. On Crotalaria vitellina 
Ker., Acapulco, Mexico, Octeber 1894—March 1895, Palmer 217 (PUR). 
This material, the first North American record, was found on a phanero- 
gamic specimen in the Chicago Natural History Museum. 

6. PUCCINIA ACROPHILA Pk. On Synthyris pinnatifida S. Wats. var. pin- 
natifida, near timberline, north side of Teton Pass, near Wilson, Wyoming, 
5 September, 1960, Cummins 60-126 (PUR). This rarely collected 
species has not been recorded on this plant in Wyoming. 

7. PUCCINIA CORONATA Cda. On Agropyron trachycaulum Malte and 
Bromus anomalus Rupr., Slide Lake, Gros Ventre River near Jackson, 
Wyoming, 30 August, 1960, Cummins 60-99, 60-103 (PUR); on Cala- 
magrostis rubescens Buckl., Indian Paint Brush Canyon Trail, Grand 
Teton National Park, Wyoming, 17 August, 1960, Cummins 60-20 
(PUR). Old aecia (spermagonia lacking) occurred on Elaeagnus cana- 
densis (L.) A. Nels. in close association at all sites. 

8. PUCCINIA DESCHAMPSIAE Arth. On Deschampsia caespitosa (L.) 
Beauv., Signal Mountain, Grand Teton National Park, Wyoming, 6 
September, 1960, Cummins 60-127 (PUR); near Wind River Lake, 
Togwotee Pass, Wyoming, 25 August, 1960, Cummins 60-84 (PUR). 
Previous records are from Colorado, Alberta, and Alaska. 


1962 | CUMMINS & BAXTER: UREDINALIS 203 


9, PUCCINIA DRABAE Rud. On Draba incerta Payson, D. sphaerocarpa 
Macbr. & Payson, Breccia Peak, above Togwotee Pass, Wyoming, 29 
August, 26 August, 1960, Cummins 60-97, 60-86 (PUR). The site is in 
alpine tundra. D. sphaerocarpa is a new host for this rarely collected 
fungus. 

10. PucciniaA mMonoica Arth. On Poa secunda Presl., Breccia Peak, 
above Togwotee Pass, Wyoming, 23 August, 1960, Cummins 60-80 
(PUR). This is the first record of the species on Poa and the Festuceae. 
Old aecia on Smelowskia calycina (Stephan) Mey. occurred in the area 
and probably belong in the life cycle. 

11. PucCINIA MONTANENSIS Ell. On Agropyron spicatum (Pursh) 
Scribn. & Sm., Togwotee Pass road, 16 mi. east of Moran, Wyoming, 22 
August, 1960, Cummins 60-62 (PUR). Old aecia were common at the 
site on Berberis repens Lindl. and probably belong in the life cycle. The 
only demonstrated aecial host is B. fendlert Gray, but the distribution of 
the fungus on grasses far exceeds the distribution of this barberry. B. 
repens probably serves in northern areas. 

12. PUCCINIA MUSENTI Ell. & Ev. On Lomatium montanum C. & R., 
Breccia Peak, above Togwotee Pass, Wyoming, 26 August, 1960, Cum- 
mins 60-90 (PUR). This relatively rare fungus has not been reported on 
species of Lomatium. 

13. PUCCINIA PAGANA Arth. On Lloydia serotina (L.) Reichb., Breccia 
Peak, above Togwotee Pass, Wyoming, 23 August, 1960, Roger S. Peter- 
son (Cummins 60-80), (PUR). The location is above timberline. The 
only previous record is the type, collected on Pike’s Peak, Colorado, in 
1904 as on Allium reticulatum Don (Clements, Cryptog. Form. Colo. 
No. 141 as Puccinia mutabilis). Arthur questioned the identity of the 
host plant when he described P. pagana. Cummins visited the type local- 
ity in July, 1961, and found P. pagana on Lloydia serotina but no rust 
fungus on the intermingled Allium. There is no doubt that the host of the 
type is also Lloydia serotina. 

14, PUCCINIA PATTERSONIANA Arth. On Agropyron spicatum (Pursh) 
Scribn. & Sm., Togwotee Pass road, 16 miles east of Moran, Wyoming, 
22 August, 1960, Cummins 60-61 (PUR). This fungus has not prev- 
iously been recorded for Wyoming. 

15, PUCCINIA WULFENI4E Diet. & Holw. On Veronica wormsk joldii R. 
& S., summit of Togwotee Pass, Wyoming, 20 August, 1960, Cummins 
60-55 (PUR). This relatively rare fungus has not previously been re- 
corded on a species of Veronica. 


Department of Botany and Plant Pathology, 
Purdue University, Lafayette, Indiana. 
Department of Botany, 
University of Wisconsin, Milwaukee. 


LITERATURE CITED 
Baxter, J. W. 1958. Notes on Rocky Mountain rust fungi. Trans. Wis. Acad. Sci. 
47:131-135. 


204 MADRONO [Vol. 16 


REVIEW 


Southern California Gardens, An Illustrated History. By Victorta PapILLa. Uni- 
versity of California Press, Berkeley and Los Angeles. 1961. 379 pages. 170 illustra- 
tions, 15 in color. $10.00. 

The passing parade of plants, people and events presented to the reader of this 
history of ornamental horticulture in southern California is an impressive one indeed. 
The story of “Southern California Gardens” begins with the Franciscan missionaries 
who came northward from Baja California to form a chain of missions from 
San Diego to Sonoma from 1769 to 1823. They brought with them a number of 
plants mostly for agricultural purposes, but some were ornamentals and among these 
were several trees which are today so much a part of our landscape that we take 
them for granted. Olive, pepper, fig, and citrus trees are but a few. The story is 
carried on from this early beginning down to the present time. 

Through the early history, but particularly beginning with the last quarter of 
the 19th century, are told the stories of those horticulturists and gardeners responsible 
for bringing plants from other parts of the country and from Europe, those who 
had the vision to realize that new plants could be grown here, and who persisted 
and were successful in their efforts. It is surprising, in fact, how many plants were 
brought here before the beginning of our present century. For those who have 
lived in southern California, reading Miss Padilla’s personal sketches of these horti- 
culturists is reading the stories of one’s friends. Her characterizations are warmly and 
personally told. 

Miss Padilla is a gardener herself and has a personal knowledge of and acquaint- 
ance with the people and the plants that she has written about. She has been a 
prominent member for many years of the Southern California Horticultural Institute, 
the organization which sponsored the publication of this history. 

The format of the book is excellent, and the photographs, including several in 
color, are of good quality and well reproduced. A list of the plant introductions of 
Dr. Franceschi, Hugh Evans, E. O. Orpet, and the former Evans and Reeves nursery 
firm, and the horticultural features of the several parks of the City of Los Angeles 
add to the interest of the book. A bibliography gives some of the sources of material 
consulted by the author. The nomenclature of the plants is accurate. There is, 
however, one criticism which this reviewer offers regarding the way in which the 
names of the horticultural or cultivated varieties (cultivars) are written. According 
to the International Code of Nomenclature for Cultivated Plants, names of cultivated 
varieties (cultivars) are to be enclosed in single quotes and not double quotes. 
Throughout this book double quotes are used, this in direct contradiction to the 
International Code. There did not seem to be any explanation on the part o1 the 
publisher for not following this very simple international rule. 

Interesting and authoritative, and written in a charming style, this book is highly 
recommended for all those interested in southern California horticulture and gardens, 
and there is, in fact, enough of general interest to recommend it for all those inter- 
ested in California horticulture and gardens. EL1zABETH McCiinTock, Department 
of Botany, California Academy of Sciences, San Francisco, California. 


INFORMATION FOR CONTRIBUTORS 


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heading, “‘Notes and News.” 

Institutional abbreviations in specimen citations should follow Lanjouw 
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Utrecht. Second Edition, 1954). 

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

Membership in the California Botanical Society is normally considered 
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VOLUME 16, NUMBER 7 JULY, 1962 
Contents 

PAGE 
AN ANATOMICAL STUDY OF THE SECONDARY TISSUES 


IN Roots AND STEMS OF UMBELLULARIA CALIFOR- 
NICA NuTT. AND LAURUS NOBILIS L., Baki Kasapligtl 205 


Rurus Davis ALDERSON (1858-1932), Reid Moran 224 


THE OCCURRENCE OF NEW ArCTIC-ALPINE SPECIES IN 
THE BEARTOOTH MouNTAINS, WYOMING-MoNTANA, 
Philip L. Johnson 229 


THE UNIQUE MORPHOLOGY OF THE SPINES OF AN 
ARMED RAGWEED, AMBROSIA BRYANTII 
(ComposiTAE), Willard W. Payne SS) 


NOTES AND NEWS 236 


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 
MI prep E. Maturtas, 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—JOoHN H. THomMAS 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Roxana S. Ferris, Dudley Herbarium, Stanford University, Stanford, 
California. First Vice-President: Edward C. Stone, School of Forestry, University 
of California, Berkeley. Second Vice-President: Thomas C. Fuller, Department of 
Agriculture, State of California, Sacramento. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding Sec- 
retary: Margaret Bergseng, Department of Botany, University of California, Berke- 
ley. Treasurer: John H. Thomas, Dudley Herbarium, Stanford University, Stanford, 
California. 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS 205 


AN ANATOMICAL STUDY OF THE SECONDARY TISSUES 
IN ROOTS AND STEMS OF UMBELLULARIA CALIFORNICA 
NUTT. AND LAURUS NOBILIS L. 


BAKI KASAPLIGIL 


This paper, dealing with the secondary tissues of roots and stems of 
Umbellularia and Laurus is a continuation of the author’s comparative 
studies (1951, 1954) of these laureceous genera seeking evidence regard- 
ing their phylogenetic relationships. 

Normal secondary growth is essentially the same in the roots and stems 
of both gymnosperms and woody dicotyledons, but because routine collec- 
tions of woody plants do not usually include roots, details of root struc- 
ture are less well known than are those of stems. Roots are considered 
“conservative” organs, but Beakbane (1941) found the anatomy of 
apple roots to be affected by the environment. Fegel’s work (1941) 
demonstrated the relatively primitive structural features of roots, and 
Noelle (1910) applied the anatomical characters of roots to the classifica- 
tion of the Coniferae. 


MATERIALS AND METHODS 


Native material of Laurus nobilis L. was collected at Antalya, Turkey, 
while native material of Umbellularia californica Nutt. was collected in 
California. Cultivated material of both species was collected in the 
Botanical Garden of the University of California at Berkeley. 

The material was studied partly in freehand sections and partly after 
maceration. Useful microtechnical methods were found in the publica- 
tions of Gassner (1931), Ball (1941), and Foster (1949, Appendix). 


HISTOLOGY OF SECONDARY XYLEM ELEMENTS 


Roots 


PERIDERM. There are no essential differences in the periderm of Umbel- 
lularia and Laurus. The outermost cell layer of the pericycle functions as 
a phellogen and produces 8-10 regular rows of phellem toward the 
periphery. The primary cortex breaks up and disappears. Phellem cells 
die as their cell walls gradually acquire suberized thickenings. Mean- 
while, the phellogen also forms a phelloderm tissue toward the inside, 
which is composed of parenchymatous cells containing starch grains and 
oil globules. Phelloderm cells closely resemble cortical parenchyma cells 
and they join the phloem parenchyma farther toward the inside. It is 
almost impossible to make a sharp distinction between phelloderm and 
phloem. Idioblastic secretory cells are abundant and diffusely distributed 
in phelloderm, while lenticels appear in the periderm during later stages 
of secondary growth. 


Maprono, Vol. 16, No. 7, pp. 205-236, July 13, 1962. 


206 MADRONO [Vol. 16 


SECONDARY PHLOEM. The primary phloem is obliterated during the 
secondary growth of Umbellularia roots, but sometimes remains dis- 
tinguishable as a faint line in transverse section. The secondary phloem 
is composed of cells which are more or less uniform in cross section. A 
large portion of this tissue consists of parenchyma cells containing starch 
grains. Sieve tubes and companion cells form small groups which can 
be identified by the absence of starch grains. Phloem rays are indis- 
tinguishable and no fibers were observed. Parenchyma cells containing 
tannic substances are abundant; they are distributed at random in the 
secondary phloem and phelloderm. 

The secondary phloem of Laurus exhibits certain differences. It con- 
tains fibers, of reduced cross sectional area and angular form, with thick 
walls and reduced lumina. Phloem rays expand in conical shape and can 
be distinguished easily. The tannin-containing cells in the secondary 
phloem and phelloderm are arranged in regular tangential rows. 

SECONDARY XYLEM. a. Transverse sections. The root wood of both 
genera is diffuse porous (figs. 1 and 2). Huber (1935) and Gilbert (1940) 
consider diffuse porosity more primitive than ring porosity. Vessel elements 
in Umbellularia have large diameters in spring wood and small diameters 
in summer wood. Occasionally 10-12 vessel elements of varying diameters 
are grouped together in summer wood. Vessel elements with large or 
small diameters are distributed in spring and summer woods of Laurus 
more or less in the same ratio. As an average there are about 64 vessel 
elements per square millimeter of Umbellularia root compared to about 
100 for Laurus. No tyloses are produced; apparently the vessels in the 
roots of both genera are entirely functional. Alten (1908) pointed out 
the abundance of tylosis formation in root woods of many trees. How- 
ever, the studies of Klein (1923) Liese (1925) and Fegel (1941) show 
the absence of tyloses in the root woods of forest trees. 

The outer and inner boundaries of the growth layers of summer wood 
are distinct in both genera. The width of growth layers is variable, but 
the proportion of spring and summer woods within the growth layers 
seems to be constant. Summer wood occupies approximately one tenth 
of the growth layer in Umbellularia and about one third of the growth 
layer in Laurus. 

Xylem rays are heterocellular, uni-, or multiseriate in both genera. Ray 
parenchyma cells contain an abundance of starch grains and tannic sub- 
stances. Idioblastic secretory cells occur commonly within the xylem rays. 
Xylem rays are less abundant in Umbellularia than in Laurus. In Umbel- 
lularia, the cells of ray parenchyma are larger in spring wood than in 
summer wood; in Laurus they are of nearly uniform size. 

Xylem parenchyma is apotracheal-diffuse in both genera, i.e., the 
parenchyma cells are distributed throughout the root wood independent 
of vessel elements. This is a very different situation from the paratracheal 
and metatracheal distribution patterns of the xylem parenchyma in stem 


1962 | 


Sp.W. 


SM. W. 


Fics. 1-2. Transverse sections of root wood: 1, Umbellularia californica; 2, Laurus 
nobilis. Legends: ves. e.==vessel element, sm.w.=summer wood, sp.w 


st.g.=starch grains, 


KASAPLIGIL: UMBELLULARIA AND LAURUS 


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i S <i 


y mm 


JX 


(es) 
a 


207 


Ves.e. 


xyl.par. 


ves.e. 


xyl.r. 
xyl par. 


id.sec.c. 


SM.W. 


Sp. W. 


spring wood, 


xyl.par.—xylem parenchyma, xyl.r.—xylem ray, id. sec.c.=idioblastic secretory cell 


208 MADRONO [Vol. 16 


woods which will be described later. Apotracheal-diffuse type is con- 
sidered as an unspecialized feature by Metcalfe and Chalk (1950). The 
xylem parenchyma cells form radially extending rows mixed with wood 
fibers. 

In Umbellularia the pith consists of thin-walled parenchyma cells; in 
Laurus it consists of sclerenchymatous cells. In both cases the cells con- 
tain much starch. 

b. Radial sections. In both genera, superimposed series of vessel 
members are very distinct if they fall on the plane of the sections (fig. 4). 
Xylem rays are composed of rectangular parenchyma cells containing 
starch grains. Libriform wood fibers are empty and dead, while the 
wood parenchyma cells are filled with starch grains. Idioblastic secretory 
cells appear rectangular or isodiametric in form, and they occur fre- 
quently in xylem rays as well as outside of the rays. 

c. Tangential sections. In Umbellularia roots the xylem rays are 1—4 
cells wide and 5-17 cells high (fig. 5), with a single cell at their pointed 
upper and lower margins. Xylem parenchyma occupies the spaces between 
the rays with a few libriform wood fibers. Generally, the xylem paren- 
chyma appears as long, narrow cells with tapering ends, but often the 
parenchyma cells also form superimposed vertical series. These series 
overlap one another so that there is no storied condition. Idioblastic 
secretory cells within the xylem rays are either scattered individually or 
form small roundish groups of 2—3 cells or even vertical series of 3-5 
cells (fig. 5). 

The xylem rays of Laurus roots are 1—3 cells wide and 1-13 cells high 
in transectional outlines. Idioblastic secretory cells may be scattered indi- 
vidually or may form small groups within the rays, but they generally 
occupy upper and lower margins of the rays. The rays taper gradually 
toward the upper and lower margins, which are generally straight instead 
of pointed (fig. 6). Usually these margins are in contact with xylem 
parenchyma. The “vertical xylem parenchyma” forms superimposed series 
of 3—8 cells, and these series run parallel to xylem rays. Libriform wood 
fibers occur in spaces between xylem rays and vertical series of wood 
parenchyma and are more abundant than in Umbellularia. 


Stems 


The secondary xylem of Umbellularia and Laurus is described and 
illustrated in various atlases and books for timber identification (Brown 
and Panshin, 1940, and Record, 1934, for Umbellularia; Greguss, 1945, 
and Huber, 1954, for Laurus; Stern, 1954, for these and many other 
Lauraceae). However, the secondary structure of stems will be described 
here briefly to provide a basis for the comparison with the root structure 
of Umbellularia and Laurus. 

UMBELLULARIA CALIFORNICA. A phellogen tissue is formed by the outer- 
most cell layer of cortex parenchyma. The epidermal tissue is broken in 


KASAPLIGIL: UMBELLULARIA AND LAURUS 209 


1962] 


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simple perforation 


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Fics. 3-4. Radial sections of root wood: 3, Umbellularia californica; 


plate, r.par.=ray parenchyma, 


nobiiis. Legends: sc.p.pl 


VES-E 


starch grain, 


wood fiber, id.sec.c.=idioblastic secretory cell, st.g 


ment, xyl.r.=xylem ray. 


210 MADRONO [Vol. 16 


places as the result of secondary thickening of the stem. Phellem and 
phelloderm layers produced as the result of the activity of this secondary 
meristematic tissue are shown in fig. 7. The cortex parenchyma below 
the periderm consists of 8—10 cell layers in which idioblastic secretory cells 
are distributed without a regular pattern. The innermost cell layers of 
the cortex—adjacent to the primary phloem—form one to several layers 
of hippocrepiform sclereids (cf. Bailey and Swamy 1948). The sheath 
of hippocrepiform sclereids in stems is composed of one or more cell layers 
which form a regular cylinder interrupted by phloem fibers. The inner 
tangential walls and the radial walls of these sclereids are thick and ligni- 
fied heavily, while their outer tangential walls are unthickened. Thus the 
hippocrepiform sclereids appear U-shaped in transectional view resemb- 
ling endodermal cells at the tertiary stage of thickening. The thick walls 
of these sclereids are provided with simple pits. These pits are generally 
opposite the pits of the adjacent cells, forming simple pit pairs. However, 
blind pits are also observed along the radial walls. Hippocrepiform 
sclereids of secondary stems 6—7 years old are living cells with large 
lumina. The cytoplasm is peripheral, while the central part of the cell is 
occupied by a vacuole. Some of these sclereids contain granular tannic 
substances and appear dark. 

The derivative cells of the vascular cambium form the secondary 
phloem externally and in this way the primary phloem is pushed out- 
ward. The primary phloem in older portions of the stem is crushed by 
the internal pressure and in the later stages of development the primary 
phloem may be entirely obliterated. However, the phloem fibers with 
thick and resistant cell walls remain in groups along the outer boundary 
of the primary phloem (fig. 7). 

Stem wood is hard and exhibits distinct growth layers. Heart and sap 
woods are distinguishable in old and thick stem portions. Heart wood is 
grayish or dark brown while the sapwood is whitish or light brown. 
Porosity is of the diffuse type as in the root wood. Xylem rays are very 
fine in transverse section and hardly distinguishable to the naked eye. 
Typically the rays are heterocellular and the xylem rays together with 
phloem rays form continuous vascular rays. The rays are not as dense 
as in the stem wood of Laurus and there are about ten rays per millimeter 
in transverse section. 

Wood fibers form regular rows extending radially. The xylem paren- 
chyma exhibits paratracheal-vasicentric arrangement. One to three cell 
layers of xylem parenchyma encircle the vessel elements as seen in the 
lower left corner of figure 7. The distribution of xylem parenchyma in the 
stem wood of Umbellularia exhibits a more advanced and specialized con- 
dition when compared to the apotracheal-diffuse type of arrangement in 
the root wood of the same species. The xylem parenchyma cells in the 
stem have thick lignified walls and contain starch grains. The primary 
xylem elements are readily identified in small groups adjacent to the pith. 


211 


KASAPLIGIL: UMBELLULARIA AND LAURUS 


1962 | 


seae> 


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SOS 


water, 


io) 


— 
oe 


? 


6 


Umbellularia californica; 


’ 


Fics. 5-6. Tangential sections of the root wood: 5 


Laurus nobilis. Legends: xyl.r.=xylem ray, id.sec.c 


.idioblastic secretory cell, r.par. 


simple pits, xyl.par.=xylem 


.==simple starch grains. 


si.p. 


si.st.g 


ray parenchyma, lib.w.fib.=libriform wood fiber, 
parenchyma, co.st.g.=compound starch grain 


) 


212 MADRONO [Vol. 16 


They possess thicker cell walls and small diameters and lumina as com- 
pared to the secondary xylem elements. The pith is composed of large 
isodiametric parenchyma cells. The outer 4—5 cell layers of the pith 
cylinder in the old portions of stems remain alive and contain starch 
grains while the inner cells of the pith die. The simple pits of the pith 
parenchyma are distinct, but the intercellular spaces are obscure. 

LAURUS NOBILIS. Epidermis, periderm and cortex tissues of the Laurus 
stem are essentially similar to those in the Umbellularia stem. 

The primary and secondary phloem groups are intersected by phloem 
rays. Figure 8 represents a transverse section of a young stem in which 
the primary phloem tissues are not obliterated. It was not possible to 
draw a clear demarcation line between primary and secondary phloem. 
However, the fiber groups shown in figure 8 help determine the approxi- 
mate position of the primary phloem groups. The phloem fibers appear 
as polyhedral, thick walled cells grouped compactly in transectional view. 
The position of the secondary phloem is determined approximately in 
figure 8 according to its position relative to the vascular cambium. The 
vascular cambium appears as if it contains 30 cell layers since the stem 
material was collected during cambial activity in July. Theoretically only 
one cell layer forms the cambial initials while the rest of the cells repre- 
sent undifferentiated derivatives of the cambium in both inner and outer 
directions. However, the vascular tissues produced by the earlier activity 
of the cambium are already differentiated into secondary structure. 

The secondary xylem is diffuse porous. The vessel elements are scat- 
tered individually or in twos in the spring wood, while three or four of 
them are arranged in small radial rows in the summer wood. The growth 
layers are distinct due to the fact that the wood fibers along the border 
lines of the growth layers are flattened and have very small diameters. 
Xylem rays are heterocellular, uni-, or biseriate. There are 12 xylem rays 
per millimeter in transverse section. The xylem parenchyma of the stem 
exhibits metatracheal arrangement which represents a more specialized 
condition compared to the apotracheal diffuse arrangement of the xylem 
parenchyma of the secondary roots of the same species. 


MORPHOLOGY OF THE SECONDARY XYLEM ELEMENTS 


Anatomical features exhibited by various sections of root and stem 
woods of Umbellularia and Laurus have been described above. In addi- 
tion, macerated material of root and stem woods was studied in the hope 
of finding other characters that might be considered of phylogenetic 
importance. 

The phylogenetic value of wood anatomy in systematic studies was 
shown clearly by Record (1934), Chalk (1937), Heimsch and Wetmore 
(1939), Tippo (1946) and others. Metcalf and Chalk (1950) emphasize 
the fact that wood structures exhibit more conservative characters than 
do external features of plants. 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS 213 


(X cu 
ss Z 2 
ZS, S55 2-6 phm 
KLE ae ace 
geo ge SS = ae aa ate ae eee Bhlen 
2Aq-eeaaH = =2>—-o ge senss— 
aogesee ae Saec Cll a= )  phdm 
Se eee ee ee ee 
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camb. 2. 
sec xyl 


Fics. 7-8. Portions of transverse stem sections showing secondary growth: 7, 
Umbellularia californica; 8, Laurus nobilis. Legends: cu.=cuticle, phm.=phellem, 
phlgn.—phellogen, phdm.=phelloderm, id.sec.c.=idioblastic secretory cell, cor.par.= 
cortical parenchyma, phl. fib.=phoem fibers, hip.scl.=hippocrepiform sclereids, 
prim.phl.=primary phloem, sec.phl.—secondary phloem, camb.z.—cambial zone, 
sec.xyl.=secondary xylem, xyl.r.=xylem ray, ep.=epidermis. 


214 MADRONO [Vol. 16 


The dimensions of wood elements are significant for both diagnostic 
and phylogenetic interpretations. In general, long and narrow wood ele- 
ments are considered more primitive than short and wide ones. However, 
the dimensions of wood elements exhibit considerable variation in various 
organs of the same plant or even in different regions of the same organ, 
and Sanio (1872) demonstrated the length increase of the wood elements 
from central toward the peripheral regions of Pinus sylvestris roots. Gerry 
(1915) showed that the root tracheids are longer than stem tracheids in 
P. palustris and P. strobus. Anderson (1951) found that the length of the 
tracheids in conifers tends to increase with the increase of distance from 
the pith and that the tracheids of spring wood are shorter than those of 
summer wood within the same “annual ring.”” Dimensional variations of 
the xylem elements in the root and stem woods of Umbellularia are obvi- 
ous in tables given below. An interesting subject for further investigation 
would be the relationship of these variations to different stages of growth. 

The morphological features of wood elements may also be used to 
interpret the degree of specialization. The conclusions regarding the primi- 
tive and advanced characters of the secondary xylem elements reached by 
wood anatomists (Bailey and Tupper, 1918; Bailey and Howard, 1941; 
Fegel, 1941; Frost, 1930a, b, 1931; Gilbert, 1940; Metcalfe and Chalk, 
1950; Tippo, 1946) are summarized below. 


PRIMITIVE CHARACTERS ADVANCED CHARACTERS 

1. Diffuse porous. 1. Ring porous. 

2. Scattered vessel elements. 2. Grouped vessel elements. 

3. Small perforation plates with 3. Wide perforation plates with a 
many bars. few bars. 

4. Scalariform perforation plates. 4. Simple perforation plates. 

5. Polyhedral vessel elements. 5. Round vessel elements. 

6. Inclined end-walls. 6. Transverse end-walls. 

7. Fiber tracheids with bordered 7. Libriform wood fibers with small 
pits. simple pits. 


8. Scalariform and bordered pits. 8. Simple pits. 


The anatomical features set forth above have been used as a basis to 
judge and compare the primitive and advanced characteristics of the 
xylem elements in the secondary structures of roots and stems in Umbel- 
lularia and Laurus in the present paper. According to the suggestion of 
Chalk and Chattaway (1934), the vessel elements were measured from 
tip to tip to obtain the length dimension. To calculate average dimen- 
sions, at least fifty measurements have been made for each element. The 
terminology proposed by the Committee on Nomenclature of the Inter- 
national Association of Wood Anatomists (1957) has been followed in 
describing the wood elements. 

SECONDARY XYLEM ELEMENTS OF UMBELLULARIA Roots. Wood 
parenchyma, fiber tracheids, and septate fiber tracheids are abundant. 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS a5 


The average length of vessel elements is 250 microns and their average 
width is 40 microns. Therefore the vessel elements in question fall into 
the group “small and short” in Metcalfe and Chalk’s (1950) classifica- 
tion. In general they are shorter and narrower than the vessel elements 
found in the stem wood of the same species. 

The ligulate tips of the vessel elements are generally long and broad 
(fig. 9, c). Vessel elements approaching a cylindrical shape are rare (fig. 
9, f-h). In transectional view they are polyhedral in form (fig. 1). 

The perforation plates of the vessel elements are generally simple, 
oblique, and distant from the ligulate tips. Scalariform perforation plates 
are very rare. A scalariform perforation plate with a single bar is shown 
in fig. 9, a. Although the vessel elements are usually provided with two 
simple perforation plates (fig. 9, c, e-h), vessel elements with three 
simple perforation plates have also been observed (fig. 9, b). In the 
latter instance, there are two simple perforation plates in one end of the 
vessel element and one perforation plate in the other end of the element. 
Some vessel elements have two perforation plates side by side in the 
middle of a vessel element without perforations at the cell ends (fig. 9, 
d). These latter two cases are characterized by profuse pitting at the 
ligulate tips of the vessel elements (fig. 9, a, d). 

The simple pits on the lateral walls of the vessel elements are small 
and generally at equal dimensions. Opposite and alternate pitting may 
be found on the walls of the same vessel element. The ligulate tips of the 
vessel elements are usually pitted, but no pitting is found in cases where 
the tips are short. 

The fiber tracheids and the septate fiber tracheids possess pits. The 
septations of these elements are primary walls and do not exhibit secon- 
dary thickenings (fig. 9, m). An interesting type of septate fiber tracheid 
is shown in figure 9, j, in which the lateral walls are about three times 
thicker than the terminal portions of the lateral walls. These tracheids 
recall “gelatinous fibers” (cf. Esau, 1953, p. 205), but their thick walls 
did not shrink during the process of dehydration. 

The wood fibers have very thick secondary walls with vestigial simple 
pits. Some of the wood fibers have wide lumina (fig. 9, 1), but some of 
them have an extremely reduced lumen (fig. 9, k) appearing like a line 
along the longitudinal axis of the cell. Septate wood fibers (fig. 9,1) are 
very similar to septate tracheids, but are distinguished from the latter 
by the presence of vestigial simple pits. Septate wood fibers may possess 
one or two partition walls of a primary nature. According to Metcalf and 
Chalk (1950), septate tracheids and septate fibers commonly occur in 
tropical woods and serve as a useful feature for determining phylogenetic 
relationships. As a matter of fact, most Lauraceae are distributed in 
tropical regions and contain either one of the septate elements or both 
types in their woods. The wood fibers of Umbellularia roots are shorter 
but wider than the fibers of the stem wood. 


[Vol. 16 


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Fics. 11-12. Tracheary elements of Laurus nobilis: 11, 


(Explanation in text). 


218 MADP.ONO [Vol. 16 


The parenchyma cells of the root wood possess lignified cell walls, but 
their cell walls are thinner than the cell walls of other xylem elements. 
Xylem parenchyma cells are elongated cells, while the ray parenchyma 
cells are smaller and tabular in shape. 

SECONDARY XYLEM ELEMENTS OF UMBELLULARIA STEMS. Vessel ele- 
ments of stem wood average 336 microns in length and 53 microns in 
width. This dimensional variation is not as pronounced as the size varia- 
tion of the vessel elements in the root wood. The transverse outlines of the 
vessel elements do not exhibit sharp corners. Generally, they approach 
drum shape in spite of much variation in their form (fig. 10, i). They are 
provided with simple perforation plates, and scalariform perforations 
have never been observed. The ligulate tips of the vessel elements are 
generally small, those with transverse perforation plates having no 
ligulate tip (fig. 10,1). These characters indicate that the vessel elements 
of the stem wood exhibit a higher degree of specialization than the vessel 
elements in the root wood of Umbellularia. 

The ligulate tips of the vessel elements may be long and wide at both 
ends of the cells, tapering gradually (fig. 10, a, b), or they may extend in 
slender needle-like form at both ends (fig. 10, c). These two are primitive 
types resembling tracheids. Some vessel elements have a single ligulate 
tip in one end of the cell that is mucronate, i.e., the end wall of the vessel 
element abruptly becomes a short tail (fig. 10, e). Some ligulate tips are 
curved inwardly. This is generally observed in short and wide vessel 
elements (fig. 10, h). 

Although the vessel elements usually have two simple perforation 
plates, some of them are provided with three perforation plates (fig. 10, b) 
and still others have only one (fig. 10, g). Two perforation plates may be 
located side by side in one end of a vessel element and the other end of 
the cell in such vessel elements tapers gradually (fig. 10, d, f). 

Intervascular pits on the lateral walls of the vessel elements are small 
and simple. Reticulate and broad simple pits are restricted to the surfaces 
of the lateral walls in contact with the xylem parenchyma cells (figs. 10, 
a, d, e, g). Scalariform pitting has not been observed. 

Fiber tracheids have approximately the same diameter as those in root 
wood. The fiber tracheids are either uniform in width, and terminating 
in small ligulate tips at both ends (fig. 10, n), or they may be constricted, 
terminating in sloping end walls (fig. 10, p). 

Septate fiber tracheids (fig. 10, m) are fusiform cells with gradually 
tapering ends. Their lateral walls are thickened equally in all directions. 
Fiber tracheids with unequal secondary wall thickening have not been 
found in the stem wood. 

Libriform wood fibers have very thick secondary walls and highly 
reduced lumina which can be distinguished sometimes as a fine line ex- 
tending lengthwise (fig. 10, k). Vestigial simple pits of the wood fibers 
are also extremely reduced, and they appear as small oblique lines along 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS 219 


both sides of the lumen trace. Wood fibers with wider lumina are not 
rare, and their diameters are smaller than the wood fibers of the root wood. 

Septate wood fibers are very similar to septate fiber tracheids in regard 
to their shapes and sizes, but they can be distinguished from the latter by 
the presence of simple pitting (fig. 10, 1). They possess one or two parti- 
tion walls; their lateral walls with secondary thickenings are thinner 
than the lateral walls of the septate fibers found in the root wood of 
Umbellularia. 

Xylem parenchyma cells have thick secondary walls with simple 
pitting, forming superimposed vertical series (fig. 10, 0). They resemble 
closely septate wood fibers even in macerated state, but are distinguished 
from the latter by the presence of secondary thickenings in their trans- 
verse walls (cf. fig. 10, a, 1). 

Ray parenchyma cells are rectangular cells with straight or slightly 
pointed end walls. Generally they possess simple pitting, but they also 
exhibit reticulate pitting on the areas in contact with vessel elements 
(fig. 10, r). 

TABLE 1. COMPARISON OF THE DIMENSIONS OF SECONDARY XYLEM ELEMENTS IN THE 
RooTs AND STEMS OF UMBELLULARIA CALIFORNICA AND LAURUS NOBILIS. 


Woop ELEMENTS AVERAGE LENGTH (microns) AVERAGE WIDTH (microns) 


Root Woop Stem Woop’ Root Woop’ STEM Woop 
Ue Lie er ee. Uc. Ln. Wice La. 


Vessel elements 250 263 $305 353 40 84 53 29 
Fiber tracheids 412 451 417 348 2 34 24 25 
Septate fiber tracheids See oS 504 459 26 30 ony) 23 
Libriform wood fibers 443 521 511 456 19 18 1) 12 
Septate wood fibers Se AS ee 19 Ba 16 oe 
Xylem parenchyma 167 160 170 93 ot Si 23 19 
Ray parenchyma 63 60 74 44 36 29 21 28 


* U.C. = Umbellularia californica Nutt. 

* Tn. = Laurus nobilis L. 

SECONDARY XYLEM ELEMENTS oF LAuRUS Roots. The average length 
of the vessel elements in the root wood is 263 microns and the average 
width is 84 microns (table 1). Obviously they are shorter and wider com- 
pared to the vessel elements of the stem wood. They fall into the group 
of ‘‘small and short vessel elements” in the classification of Metcalfe and 
Chalk (1950) since their average width is smaller than 100 microns and 
their average length shorter than 350 microns. The ligulate tips of vessel 
elements are short and pointed (fig. 11, a, b, d). The transectional out- 
line of the vessel elements is polyhedral. 

The perforation plates of the vessel elements are either scalariform or 
simple and exhibit three types of combinations in individual vessel ele- 
ments: a) both perforation plates may be scalariform (fig. 11, a); b) 
one of the perforation plates may be scalariform and the other one simple 
(fig. 11, b); c) both perforation plates may be simple (fig. 11, e, f). The 


220 MADRONO [Vol. 16 


number of bars in the perforation plates is generally 7—8, but rarely 
there may be only a single bar. Although in general there is one perfora- 
tion plate at each end of the vessel elements, there are exceptions to this 
rule. Some vessel elements may have three perforation plates. For example, 
figure 11, d, illustrates a vessel element with two simple and one scalari- 
form perforation plates. Still some vessel elements may have but one 
simple perforation plate (fig. 11, g). A solitary scalariform perforation 
plate has never been observed. The vessel elements with a single perfora- 
tion plate exhibit abundant pitting on their lateral walls, and the im- 
perforate end of the vessel element tapers gradually so that such vessel 
elements acquire a funnel shape. Both of the perforation plates may be 
located side by side in one end of some vessel elements, (fig. 11, c); the 
imperforate end of such a vessel element tapers gradually and possesses 
many simple pits. In general the simple perforation plates are either 
straight or slightly sloping (fig. 11, e-g). These three illustrations also 
show that the vessel elements terminating with simple perforation plates 
have either very short ligulate tips or none. The cylindrical vessel elements 
with simple perforation plates shown in figures 11, e and f, undoubtedly 
represent the most advanced types, but they occur together with the 
primitive types possessing scalariform perforation plates. 

The vessel elements exhibit three types of pitting: a) scalariform 
pitting (fig. 11, d-g); b) reticulate pitting (fig. 11, e, g); c) simple pitting 
(fig. 11, a-c). According to Bailey (1954) scalariform pits represent a 
primitive type, but interestingly enough the scalariform pits are asso- 
ciated quite often with simple perforation plates in the same vessel ele- 
ments, forming vertically arranged regular rows on the lateral cell walls. 
The scalariform pitting does not occur in the vessel elements of Umbel- 
lularia, hence they are more specialized than the vessel elements of Laurus. 
Reticulate simple pits are restricted to the wall areas of the vessel ele- 
ments in contact with the ray parenchyma cells in the root wood of Laurus. 
Simple pits, however, are located on the lateral walls as well as in the 
ligulate tips of the vessel elements. Although some vessel elements possess 
exclusively simple pits, still others exhibit all three types of pitting at 
the same time (fig. 11, e). 

The fiber tracheids are abundant in the root wood, and they are longer 
and wider than those in the stem wood (cf. table 1). The size variation 
of the fiber tracheids in the root wood is not as pronounced as the wide 
range of variation shown by the fiber tracheids of the stem wood. 

The septate fiber tracheids are also longer and wider than those of the 
stem wood. They may be provided with one or several septa. Lumina 
are wide and they taper gradually toward the ends (fig. 11, k). 

Libriform wood fibers possess secondary walls with varying degree of 
thickening. Consequently they exhibit wide as well as narrow lumina. 
Those with wide lumina resemble fiber tracheids, but they are distin- 
guished by the presence of vestigial simple pits (fig. 11, }). The propor- 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS 221 


tion of libriform wood fibers in the secondary xylem of Laurus roots is 
relatively smaller than the proportion of wood fibers in the secondary 
xylem of stems. Septate wood fibers have not been observed, and the 
septate elements resembling fibers are actually tracheids. 

Xylem parenchyma cells exhibit a greater size variation than those in 
the stem wood. They are provided with oval simple pits. Cell shape is 
elongated and terminates in straight or abruptly tapering end walls (fig. 
11,h). 

Ray parenchyma cells are isodiametric and show slight variation in 
size (fig. 11, 1). They are provided with simple as well as with reticulate 
pitting. 

SECONDARY XYLEM ELEMENTS OF LAURUS STEMS. The vessel elements 
exhibit considerable size variation (cf. fig. 12, g, j}). As an average they 
are longer and narrower than the vessel elements of the roots (cf. table 1). 
The ligulate tips of the vessel elements are more conspicuous, more slen- 
der and longer than those of the root wood (fig. 12, c, d, g, j, k). This 
situation is similar to that described for Umbellularia. The vessel elements 
which approach the cylindrical form are rare (fig. 12, 1). The transec- 
tional view of the vessel elements is angular in form. 

Tertiary thickenings in the form of spiral bands on the secondary walls 
have not been observed although they were reported by Greguss (1945). 
The vessel elements are generally plugged by tyloses which do not occur 
in the vessel elements of the root wood of Laurus. The formation of tyloses 
in the vessel elements of Laurus stem distinguishes these elements from 
the vessel elements of the root and stem of Umbellularia. The presence 
of tyloses in other members of Lauraceae is reported in the literature. 
For example, Carpenter and Leney (1952) demonstrated the formation 
of tyloses in vessel elements of Sassafras albidum (Nutt.) Nees. 

The vessel elements are provided with scalariform or simple perfora- 
tion plates like those in the secondary xylem of the roots. However, a 
notable difference is the occurrence of reticulate perforation plates (fig. 
12, e, {). The number of bars in scalariform perforation plates is reduced, 
and perforations with one or two bars are fairly common. The vessel ele- 
ments always possess two perforations which are terminal in position. 
The occurrence of one or three perforations has not been observed al- 
though this situation is common in the vessel elements of the secondary 
xylem of the root. Either both of the perforation plates may be scalari- 
form (fig. 12, a, j}) or one of them scalariform and the other one simple 
(fig. 12, b-d) or both of them may be simple (fig. 12, g, k, 1). The vessel 
elements with reticulate perforation plates also exhibit similar variation 
(fig. 12, e, f). In general, the perforation plates are inclined; occasionally 
they become horizontal in cylindrical vessel elements. 

Scalariform, reticulate, and oval simple pits occur in varying combina- 
tions. Scalariform and simple pits may occur together on the longitudinal 
walls of vessel elements (fig. 12, d, k). Reticulate pits are commonly 


222 MADRONO [Vol. 16 


associated with reticulate perforation plates (fig. 12, f). Simple pits are 
arranged oppositely as well as alternately in vertical rows. 

The fiber tracheids and septate fiber tracheids are rare in the secondary 
xylem but common in the primary xylem. The latter may be provided 
with one or two septa. The fiber tracheids in the stem are smaller than 
those in the roots. 

Libriform wood fibers are the dominant wood elements of the stem 
structure. They are characterized by their gradually tapering forms, 
thick secondary walls, narrow lumina, and very small diameters (fig. 
a) 

Septate libriform wood fibers do not occur in stem wood. 

Xylem and ray parenchyma cells of the stem wood are similar to those 
in the root wood. However, there are fewer parenchyma cells in the stem 
wood than in the root wood. 


CONCLUSION AND SUMMARY 

1. The secondary root and stem structures of Umbellularia californica 
and Laurus nobilis are compared anatomically. The pith of Umbellularia 
roots is composed of parenchyma cells, while the pith of Laurus roots 
consists of sclerenchyma cells. Idioblastic secretory cells and cells con- 
taining tannic substances are abundant throughout the secondary root 
and stem tissues of both genera. 

2. Diffuse porosity is a common feature of the secondary xylem in 
the roots and stems of both species. 

3. The wood parenchyma of Umbellularia roots exhibits an “‘apo- 
tracheal-diffuse” arrangement, while the wood parenchyma of Umbellu- 
daria stems exhibits a more advanced “paratracheal-vasicentric” arrange- 
ment. Wood parenchyma of Laurus roots is ““apotracheal-diffuse,” while 
the stem wood is ‘“‘metatracheal,” a more specialized condition. 

4. The vessel elements in the secondary xylem of Umbellularia roots 
are provided with elongated ligulate tips, inclined perforation plates, 
and in transectional view, they are angular in form. These are consid- 
ered primitive as compared to the vessel elements in the stem wood of 
the same species, the latter having short ligulate tips, transverse simple 
perforations, and a circular form in transectional view. 

5. The arrangement of wood parenchyma and the comparative mor- 
phology of the tracheary elements reveal the fact that primitive characters 
are retained in roots, thus providing a useful anatomical tool for phylo- 
genetic studies. 

6. The vessel elements of the root and stem woods of Umbellularia 
are devoid of tyloses. Likewise, the tracheary elements in the root wood 
of Laurus lack tyloses, but the vessel elements of the stem woods of 
Laurus are often plugged by the development of tyloses. 

7. The vessel elements in the root and stem woods of Laurus possess 
scalariform, reticulate, and simple perforation plates, and their longi- 
tudinal walls are provided with scalariform and simple pitting. In these 


1962 | KASAPLIGIL: UMBELLULARIA AND LAURUS 228 


respects they resemble the tracheids. The vessel elements of Umbellu- 
laria are more specialized than those of Laurus. 

8. Fiber tracheids, septate fiber tracheids, and libriform wood fibers 
occur throughout the secondary tissues of both species in varying sizes 
and proportions. On the other hand the septate libriform wood fibers and 
‘“oelatinous tracheids” are restricted to the secondary xylem of Umbellu- 
laria, but are entirely absent in the secondary xylem of Laurus. 

9. These differences suggest that the phylogenetic affinity between 
these two genera is somewhat distant within the family, although both 
genera perfectly fit the natural group of Lauraceae in many respects. 


Department of Biology, 
Mills College, Oakland, California 


LITERATURE CITED 


ALTEN, H. V. 1908. Beitrage zur vergleichenden Anatomie der Wurzeln, nebst Bemer- 
kungen ueber Wurzelthyllen, Heterorhizie, Lenticellen. Inaug. Diss. Gottingen. 
ANDERSON, E. A. 1951. Tracheid length variation in conifers as related to distance 

from pith. Jour. Forest. 49:38-42. 

BaiLey, I. W. 1954. Contributions to plant anatomy. Chron. Bot., Waltham, Mass. 

, and R. A. Howarp. 1941. The comparative morphology of the Icacinaceae: 

II. Vessels. Jour. Arnold Arb. 22:171-187. 

, and B. G. L. Swamy. 1948. Amborella trichocarpa Baill., a new morphol- 
ogical type of vesselless dictotyledon. Jour. Arnold Arb. 29:245-254. 

———, and W. W. Tupper. 1918. Size variation in tracheary cells: I. A. comparison 
between the secondary xylems of vascular cryptogams, gymnosperms and angio- 
sperms. Proc. Am. Acad. Sci. 54:149-204. 

Batu, E. 1941. Microtechnique for the shoot apex. Am. Jour. Bot. 28:233-243. 

BEAKBANE, A. B. 1941. Anatomical studies of stems and roots of hardy fruit trees. 
III. The anatomical structure of some clonal and seedling apple rootstocks stem- 
and root-grafted with a scion variety. Jour. Pomology and Hort. Sci. 18:344-367. 

Brown, H. P. and A. J. Pansuin. 1940. Commercial timbers of the United States. 
New York and London. 

CARPENTER, C. H. and L. LEeNney. 1952. 91 paper making fibers. N.Y. State Univ., 
Coll. Forestry, Tech. Publ. 74. 

Cuak, L. 1937. The phylogenetic value of certain anatomical features of dicotyle- 
donous woods. Ann. Bot. n.s. 1:409-428. 

and M. M. Cuattraway. 1934. Measuring the length of vessel members. Trop. 
Woods 40:19-26. 

COMMITTEE ON NOMENCLATURE, INTERNATIONAL ASSOCIATION OF Woop ANATOMISTS. 
1957. Glossary of terms used in describing woods. Trop. Woods 107:1-36. 

Esav, K. 1953. Plant anatomy. John Wiley & Sons, Inc., New York. 

Frecer, A. C. 1941. Comparative anatomy and varying physical properties of trunk, 
branch, and root wood in certain northwestern trees. N.Y. State Univ., Coll. 
Forestry, Tech. Publ. 55. 

Foster, A. S. 1949. Practical plant anatomy. D. Van Nostrand Co., New York. 

Frost, F. H. 1930 a. Specialization in secondary xylem of dicotyledons. I. Origin of 
vessels. Bot. Gaz. 89: 67-94. 

. 1930 b. Specialization in secondary xylem of dicotyledons II. The evolution 
of the end wall of the vessel segment. Bot. Gaz. 90:198—212. 

——. 1931. Specialization in secondary xylem of dicotyledons. III. Specialization 
of the lateral wall of the vessel segment. Bot. Gaz. 91:88—96. 

GAssNER, G. 1931. Mikroskopische Untersuchung pflanzlicher Nahrungs-und 
Genussmittel. Jena. 

Gerry, E. 1915. Fiber measurements studies. Science 61:179. 


224 MADRONO [Vol. 16 


GILBERT, S. G. 1940. Evolutinary significance of ring porosity in woody angio- 
sperms. Bot. Gaz. 102:105-120. 

Grecuss, P. 1945. Bestimmung der mitteleuropaeischen Laubholzer und Straeucher 
auf xylotomischer Grundlage. Ungar. Naturwiss. Mus. Budapest. 

Hermscu, C. Jr. and R. H. Wetmore. 1939. The significance of wood anatomy in 
the taxonomy of the Juglandaceae. Am. Jour. Bot. 26: 651-660. 

Huser, B. 1935. Die physiologische Bedeutung der Ring—und Zerstreutporigkeit. 
Deutsch. Bot. Gesell. Ber. 53:711—719. 

. 1954. Mikrophotographischer Atlas Mediterraner Hoelzer. Fritz Haller Verlag, 
Berlin. 

KASAPLIGIL, B. 1951. Morphological and ontogenetic studies of Umbellularia califor- 
nica Nutt. and Laurus nobilis L. Univ. Calif. Publ. Bot. 25:115-240. 

. 1954. The growth of the root apices in Umbellularia californica Nutt. and 
Laurus nobilis L. 8th Congr. Int. Bot. Rap. & Comm., Sect. 7-8:263-265. 

KLEIN, G. 1923. Zur Aetiologie der Thyllen. Zeitschr. f. Bot. 15:417-439. 

LirsE, J. 1925. Beitraege zur Anatomie und Physiologie des Wurzelholzes der Wald- 
baeume. Deutsch. Bot. Gesell. Ber. 42 :91-97. 

METcaALrFE, C. R. and L. CHALK. 1950. Anatomy of the dicotyledons. Clarendon Press, 
Oxford. 

NoELLE, W. 1910. Studien zur Vergleichenden Anatomie und Morphologie der Koni- 
ferenwurzeln mit Rucksicht auf die Systematik. Bot. Zeitung 68:169-266. 

Recorp, S. J. 1934. Identifications of the timbers of temperate North America. John 
Wiley & Sons, New York. 

Santo, K. 1872. Ueber die Groesse der Holzzellen bei der gemeinen Kiefer (Pinus syl- 
vestris L.) Jahrb. f. wiss. Bot. 8:401-420. 

STERN, W. L. 1954. Comparative anatomy of xylem and phylogeny of Lauraceae. 
Trop. Woods 100:1-72. 

Trppro, O. 1946. The role of wood anatomy in phylogeny. Am. Midl. Nat. 36:362-372. 


RUFUS DAVIS ALDERSON 
(1858-1932) 


REID MORAN 


The name of R. D. Alderson has been known to botanists both from 
his large collections in San Diego County, California, and from the writ- 
ings of E. L. Greene, who based several species on these collections and 
named for Alderson a phacelia, a helianthemum, and a rose. Yet to 
present-day botanists, Alderson is no more than a name. 

Rufus Davis Alderson was born in Alderson, [now West] Virginia, 
November 2, 1858, the younger son of Rufus Davis Alderson and Hester 
Ann Ammen Alderson. After teaching for three years in West Virginia, 
he attended the National Normal School, in Lebanon, Ohio, receiving a 
bachelor of science degree in August 1882. His subjects included botany, 
zoology, natural philosophy, physiology, herbarium, and astronomy. 
After two more years of teaching, he was from 1884 to 1887 the proprietor 
and editor of the Alderson Statesman. The word “PRINTERY?” on his 
door struck the fancy of a fellow editor, who, about 1885, wrote in the 
Pomeroy, Ohio, Democrat: 


1962 | MORAN: ALDERSON 229 


Fic. 1. Rufus D. Alderson. “In California about 1895.” 


“Tt has taken us full thirty-five years along printer’s lane to reach a 
printery. .. . We have seen binderys, bakerys, hennerys, piggeries, hog- 
geries, doggeries and groggeries, but never till auspicious fate led us into 
the Greenbrier Valley and up to the throne of the STATESMAN did we see 
a printery. In the Sancterry—we mean sanctum—was the editor. A pleas- 
ant, courteous young gentleman, who has served the State and a pleased 
constituency as a member of the Legislature, and who is greatly interested 
in all that pertains to Alderson and surrounding country... . The sanc- 
tum of the statesman who runs the STATESMAN serves the quadruple pur- 
pose of consultation room, library, editorial office and sleeping room... . 
Here into the bachelor boudoir come the exchanges and their editors, 
and here come the politicians, deacons, doctors, leaders, drivers and pro- 
prietors of enterprise to consult and move ahead.”’ 

In November 1887, apparently after a break with his childhood sweet- 


226 MADRONO [Vol. 16 


heart, Emma B. Perry, Alderson came to San Diego. For a time he was 
foreman in a print shop, and it is thought that he did some newspaper 
work. On September 6, 1888, he married Minnie E. Matchin; they were 
divorced two years later. Between 1889 and 1897, he taught school in 
San Diego County, his posts including Potrero, Warners, Descanso, E] 
Nido, Glencoe, Santa Ysabel, Spring Hill, and Bloomdale. His salary 
was $60 to $70 per month, and his contracts varied from half a month 
to eight months, one reading “length of term subject to amount of money 
available.” From 1891 to 1895, most of his teaching was at Santa Ysabel, 
about 35 miles northeast of San Diego. During this time, he rented a 
small ranch at nearby Witch Creek, whose name appears on so many of 
his herbarium labels. In 1892 he returned to West Virginia to marry 
Emma Perry on August 8, leaving with her immediately for California 
again. In 1897 he was listed as living in Del Mar. That year, in rather 
frail health, he returned to West Virginia to live. Advised by his doctor 
to work outdoors, he took up cattle breeding and dairy farming and 
during the next 28 years built up a high-producing herd of Jersey cattle. 
He died at Lewisburg, West Virginia, May 11, 1932. 


Alderson’s daughter, Mrs. Ira D. Humphreys, remembers him as self- 
disciplined and a perfectionist, with an inquisitive mind, broad interests, 
and an exceptional memory, a man who worked long hours, never took a 
vacation, and while indoors always had a book in his hand. He read aloud 
to his family from history and literature. 


At the age of fifteen, Alderson was already identifying native plants, 
as shown by notes and dates in his worn copy of Wood’s “American 
Botanist and Florist,” edition of 1874. It is not known how much he col- 
lected before leaving West Virginia, but after his return apparently his 
botany was mostly of a more practical sort; and seemingly few specimens 
of his from West Virginia have found their way into public herbaria. 
Millspaugh (1913:11) listed “R. W. Alderson” among those collectors 
a few of whose specimens were in the herbarium of Professor Sheldon, 
now in the herbarium of West Virginia University. Apparently the “D” 
of Alderson’s signature was misread for “‘W”’: according to a letter from 
Weldon Boone, this ““D” was sometimes open and could be so misread. 


In San Diego County, Alderson collected plants at least from 1891 
to 1896 but apparently most actively in 1893 and 1894. He also collected 
some insects and mollusks. Some details of his botany come from eight 
letters, now in the archives of the University of Notre Dame, written be- 
tween April 16, 1893, and March 27, 1895, mostly at Witch Creek, from 
Alderson to Professor E. L. Greene at the University of California. 
Greene’s letters have not been found. Alderson first wrote to Greene at 
the suggestion of H. W. Henshaw of the U.S. Biological Survey, who was 
visiting naturalist Frank Stephens, Alderson’s neighbor at Witch Creek.? 


1 Concerning the life of Frank Stephens, see Stephens 1918 and Huey 1938. 


1962 | MORAN: ALDERSON 22a 


Alderson introduced himself to Greene as a student and teacher of botany 
and inquired about Greene’s monograph on oaks, very highly recom- 
mended by Henshaw, and about other publications to help him identify 
plants. He later thanked Greene for a copy of Greene’s “‘Manual of the 
Botany of the Region of San Francisco Bay,” saying that for use in the 
schools it was much better than Rattan’s ‘““A Popular California Flora,” 
then in use. 


In October 1893, Alderson sent a large shipment of specimens to 
Greene for identification, saying that he had collected between 500 and 
600 species that season. Many of these were from about Witch Creek; 
but he had also made two collecting trips over the Cuyamaca Mountains 
to Campo and thence to San Diego, one early in May, the other in late 
fall. In April 1894, he wrote that he was collecting again and wanted to 
make a clean sweep, taking everything. This year, with Greene in Europe, 
he had some plants determined by Samuel B. Parish of San Bernardino, 
a keen student of the flora of southern California.” Though it was a 
dry year and collecting relatively poor, the next February he shipped 
Greene 328 specimens with labels and eight more for which he did not 
know the names. 


In April 1894, Alderson inquired of Greene as to possible purchasers of 
plant specimens to help defray cash expenditures in collecting. And in 
January 1895, he wrote that he preferred to sell to Greene rather than 
to exchange, for he had been getting many plants from other parts of the 
state by exchange and had not the room to store them. 


There is no record that Alderson was associated with the San Diego 
Society of Natural History, but it appears that he was acquainted at 
least with T. S. Brandegee® and therefore probably with the other botan- 
ists of San Diego. In February 1895, Alderson wrote Greene that he hoped 
to visit the Colorado Desert with a party of naturalists and a florist. 
Labels for that year show that he was at Palmetto Springs on the Colorado 
Desert on the last day of March, that Brandegee was there on the first 
of April, and that Frank Stephens was at Vallecitos, about 10 miles to 
the northwest, on the first and third of April. We may probably assume 
that they all got together. 


When Alderson went East, at least a good part of his herbarium went 
with him. Though much of it has since been lost, a remnant of some 275 
specimens has recently been given by his daughter to the San Diego 
Museum of Natural History, where there were already about 100 of his 
specimens received with the herbarium of Mary Snyder. Many of the 
specimens sent to Greene apparently are still in the herbarium of the 
University of California at Berkeley, though the types, at least, went 
with Greene to the University of Notre Dame. There is a large representa- 


2 Concerning the life of S. B. Parish, see Jepson 1932. 
® Concerning the life of T. S. Brandegee, see Setchell 1926. 


228 MADRONO [Vol. 16 


tion in the Dudley Herbarium, at Stanford University, from the Parish 
herbarium; but to judge from the specimens cited by Wiggins (1929), the 
set is far from complete. Other specimens are at Harvard University, the 
Missouri Botanical Garden, the University of Michigan, the United 
States National Herbarium, and probably various other institutions. 

Alderson sent his specimens to Greene under numbers, though Greene 
did not cite these numbers and apparently they were not always kept 
on the labels. Many of Alderson’s specimens in the Dudley Herbarium are 
numbered, not chronologically, perhaps in the same series; others are un- 
numbered. When he added printed labels to his own herbarium some- 
time after the collecting season of 1894—\the last season represented in 
what I have seen of this herbarium—Alderson renumbered his specimens 
to beyond 12,400. The new numbers are written on the labels where they 
might be taken for field numbers, not only on his own specimens but also 
on many collected by S. B. Parish and by Frank Stephens. Besides the 
new numbers, specimens in his own herbarium sometimes also bear num- 
bers corresponding to those on his specimens in the Dudley Herbarium, 
mostly penciled on the sheets, probably before the labels were added. 
Specimens from the herbarium of Mary Snyder and probably others 
sent out in exchange, bear his new herbarium numbers. Specimens he 
collected in 1895 and 1896 have numbers in the 900’s to 1200’s, ap- 
parently in continuation of the original series; but again, others are 
unnumbered. 

My thanks are due to Dr. Robert McIntosh of the Greene-Nieuwland 
Herbarium, University of Notre Dame, for copies of the Alderson-Greene 
correspondence; to the Serra Museum of San Diego for several old 
records; to Miss Annetta Carter and Mrs. R. S. Ferris for checking 
herbarium labels; to Professor Joseph Ewan for various suggestions; 
and especially to Mrs. Ira D. Humphreys of Ronceverte, West Virginia, 
for many details of Alderson’s life and for the fine portrait. 


Natural History Museum 
San Diego, California 


REFERENCES CITED 


Huey, Laurence M. 1938. Frank Stephens, pioneer. Condor 40:101-110. 

Jepson, WILLIs Linn. 1932. Samuel Bonsall Parish. Univ. Calif. Publ. Bot. 16:427- 
444, pl. 32. 

MitispaucH, C. F. 1913. The living flora of West Virginia. West Virginia Geol. Surv. 
5(A) 21-389, 454-487. 

SETCHELL, WILLIAM ALBERT. 1926. Townshend Stith Brandegee and Mary Katharine 
(Layne) (Curran) Brandegee. Univ. Calif. Publ. Bot. 13:155-178, pls. 13, 14. 

STEPHENS, FRANK. 1918. Frank Stephens—an autobiography. Condor 20:164—-166. 

Wiccrns, IRA Loren. 1929. Flora of San Diego County, California: a phytogeographic 
and taxonomic study. i-Ixxxi, 1-888. Unpublished thesis, Stanford University. 


1962] JOHNSON: ARCTIC-ALPINE PLANTS 229 


THE OCCURRENCE OF NEW ARCTIC-ALPINE SPECIES 
IN THE BEARTOOTH MOUNTAINS, 
WYOMING-MONTANA 


Puitip L. JOHNSON * 


During three summers of field work, 1958-1960, in the Beartooth 
Mountains, Wyoming-Montana, an extensive study was made of the alpine 
plant communities in relation to cryopedogenic (soil frost) processes and 
patterns (Johnson and Billings, 1962). Extensive alpine tundra is found 
in these mountains between elevations of 10,000 and 12,000 feet. The 
range is traversed by United States Highway 312 northeast of Yellow- 
stone National Park. This study was concentrated on the southern end 
of the mountain range because of the better developed vegetation, the 
diversity of patterned ground features, and accessibility. From field 
observation the Beartooth tundra in Wyoming appears to have received 
much less glaciation than farther north in Montana which may account 
for the floristic diversity. Of particular interest are the numerous alpine 
bog habitats which are decidedly less frequent in the Rocky Mountain 
alpine zone than in Arctic regions. Many of these bog habitats are 
underlain with permafrost within three feet of the surface. 

As a group the thirteen species discussed herein are of particular 
ecological interest because they are known primarily from the North 
American Arctic flora. It seems apparent that bog habitats and solifluc- 
tion slopes have served as a refugia for Arctic plants since the last 
glacial period. The present hypothesis is that a bog environment is 
capable of dissipating the present excessive summer heat load by the 
high latent heat of vaporization associated with evaporation from a wet 
site. This hypothesis is analogous with the investigations of Dahl (1951) 
in Scandinavia where the lower altitudinal limit of many alpine species 
is correlated with maximum summer temperatures. It is probable that 
this correlation affects the plant through a critical maximum temperature 
as suggested by field studies of photosynthetic and respiration processes 
(W. D. Billings, personal communication). 

Seven species, Phippsia algida, Carex misandra, Kobresia macrocar pa, 
Eriophorum callitrix, Koenigia islandica, Rumex acetosa, and Draba 
glabella, are new records in the Beartooth Mountains and in Wyoming. 
The remaining six species, Festuca baffinensis, Kobresia bellardu, Carex 
capitata, C. nelsoni, Juncus albescens, and J. castaneus, are new records 
in the Beartooth Mountains and rare species in Wyoming. Only Carex 
capitata and Rumex acetosa have been reported from Montana. 


PHIPPSIA ALGIDA (Phipps) R. Br. is a densely caespitose plant having 
boat-shaped leaf tips and resembling a small Poa, but it is a member of 


1 Present address: Rocky Mountain Forest and Range Experiment Station, 
Laramie, Wyoming. 


230 MADRONO [Vol. 16 


the tribe Agrostideae. The plants are relatively common only in wet 
sand or gravel at the base of late melting snowbanks. These sites, drenched 
with snowmelt water most of the growing season, are subjected to con- 
siderable needle-ice activity throughout the summer and fall. Phippsia 
forms an open community between alpine bog vegetation and late snow- 
beds devoid of plants. Koenigia islandica L. and Epilobium alpinum L. 
are frequent associates. The species, according to Porsild (1952), is 
strongly nitrophilous. It has a circumpolar, widespread, high Arctic dis- 
tribution, and has also been collected at several sites in Clear Creek 
County, Colorado (Harrington, 1954). Collections were made on both 
sides of Beartooth Pass; one in the head of Wyoming Creek (18 July 
1960, Johnson 168), and another east of Frozen Lake (10 August 1960, 
Johnson 219). 


CAREX MISANDRA R. Br. is common to infrequent on slightly raised 
mineral soil within wet sedge bogs. It was found associated with Kobresia 
macrocarpa Clokey and Poa longipila Nash. The plants are densely 
caespitose with 3-5 spikes on long slender, often drooping peduncles. 
The species is known to have a circumpolar, high Arctic distribution, 
however, Harrington (1954) reports collections from north central Colo- 
rado and from northern Utah (Lewis, 1958). Collections were made on 
a solifluction terrace in the head of Wyoming Creek (5 August 1960, 
Johnson 204, and 10 August 1960, Johnson 233). 


KOBRESIA MACROCARPA Clokey | K. bellardiu var. macrocarpa (Clokey) 
Harrington| is an alpine sedge relative previously reported only from 
central Colorado (Harrington, 1954). It differs markedly from K. bel- 
fardit in having a larger inflorescence, stouter culms and wider leaves. 
No intergradation was observed. Both species are apparently rare in 
Wyoming, although they are close associates in the Beartooth tundra on 
well developed soils in alpine turf. The collections are: 30 August 1960, 
Johnson 184 on a gentle slope east of Twin Lakes; 5 August 1960, John- 
son 205A, and 10 August 1960, Johnson 233A from a sedge meadow on a 
solifluction terrace in the head of Wyoming Creek; and 20 August 1960, 
Johnson 257 from a steep northwest slope north of Gardner Lake. 


ERIOPHORUM CALLITRIX Cham., a cotton grass, is confined to three 
bog sites east of Beartooth Pass. Two of these sites are known to be 
underlain with permafrost. The species is codominant with Carex scopu- 
lorum Holm on water-saturated peats formed behind solifluction ter- 
races. The plants form individual tufts with solitary spikelets subtended 
by black to lead-colored spathes and scales. The species is widely dis- 
tributed in the North American Arctic and subarctic regions, but no 
previous reports are known from the Rocky Mountains according to 
Porsild (1952). The collections are: 21 June 1958, Johnson 58B and 
27 July 1958, Johnson 5&8 from near the head of Twin Lake cirque; 
24-30 July 1959, W. M. Johnson, near the head of the North Fork of 


1962] JOHNSON: ARCTIC-ALPINE PLANTS 231 


Popo Agie Creek in the Wind River Mountains, Fremont County, Wyo- 
ming (identified by A. E. Porsild). 


KOENIGIA ISLANDICA L. is one of the few annuals in the Arctic-alpine 
flora. The plants, which seldom exceed 3 cm. in height, are locally very 
numerous in saturated sands, moss mats and organic soil which receive 
snowmelt water throughout the summer. Koenigia may extend into wet 
sedge meadows around the base of sedge hummocks, but it is mostly 
confined to the margins of alpine bogs, lake shores, and stream drainages 
adjacent to late snowbeds. The one or two pairs of sessile cauline leaves 
and the terminal, apetalous flower of plants exposed to direct sunlight 
develop more anthocyanin pigment than plants growing in partial shade. 
The distribution of this plant is considered circumpolar, low Arctic. 
It has recently been collected at several alpine stations in Colorado 
including Mount Evans and Rocky Mountain National Park. The species 
was observed at several sites on both sides of Beartooth Pass (4 August 
1959, Johnson 116). 


RUMEX ACETOSA L., green sorrel or sour dock, is naturalized from 
Europe throughout much of temperate, eastern North America. It 
appears, however, to be native in the American Arctic. A single rec- 
ord is known from Glacier National Park, Montana (6 July 1922, 
J. W. Severy 36) along Gunsight Pass Trail. The only plants of the 
species encountered recently were growing in moist alpine turf on a 
steep northwest slope north of Gardner Lake (26 July 1958, John- 
son 63B). It is assumed that this collection represents a southern range 
extension of native Arctic populations rather than an introduced weed, 
since the plant is not otherwise known to occur in Wyoming. 


DRABA GLABELLA Pursh is the probable identification of a collection 
which has been tentatively confirmed by Dr. Reed C. Rollins of the 
Gray Herbarium (Rollins, 1961). If true, this site represents a long 
southern extension of its known Arctic distribution. Porsild (1952) 
reports the species as, ‘“‘strongly nitrophilous, favouring animal dung.” 
The widely scattered plants were growing in sheep trails on a very steep, 
exposed northwest slope north of the highway at Gardner Lake (8 July 
1959, Johnson 112). 

It should be pointed out that this species is in addition to eleven alpine 
Draba species reported by Rollins (1953) from Clay Butte, a glacial 
monadnock of sedimentary rocks seven miles west of Beartooth Pass. 
Five of these species and D. glabella were collected from granitic parent 
material in the course of this study (Johnson and Billings, 1961). 

Several additional collections from the Beartooth Mountains are from 
similar habitats. Thanks to the annotations of Dr. A. E. Porsild, Festuca 
baffinensis Polunin, is now distinguished from F. brachyphylla Schult. 
by a culm which is puberulent on the upper half and a usually shorter, 
dark purple panicle. A previous collection from Park County, (Porter 


232 MADRONO [Vol. 16 


& Rollins 5875) has been so annotated. The present collection (5 August 
1960, Johnson 203) is from a frost boil in the head of Wyoming Creek. 


KOBRESIA BELLARDII (All.) Degland [by some, K. myosuroides (Vill. ) 
Fiori and Paol.] was first collected in Wyoming near the head of the 
North Fork of Popo Agie Creek, Fremont County in the Wind River 
Mountains (24-30 July 1959, W. M. Johnson). It is now recorded from 
Park County (30 July 1960, Johnson 185) from a gentle alpine slope 
east of Twin Lakes. It was associated with, but distinct from, K. macro- 
carpa. In the Colorado alpine tundra, K. bellardii is dominant, forming 
nearly pure stands on undisturbed snow-free ridges; the genus is evi- 
dently rare in Wyoming. 


CAREX CAPITATA L. is known from one previous collection in Wyoming 
(1893, Frank Tweedy 3, Big Horn Range, Sheridan County). It was 
again encountered at 11,000 feet elevation on a ridge top one mile north- 
west of Beartooth Pass (20 August 1958, Johnson 55). The species 
is known from Eurasia, southern South America, and Arctic North 
America, extending southward as far as Colorado, Utah, and Nevada 
(Lewis, 1958). 


CAREX NELSONTI Mack. is restricted to Colorado, Utah, and Wyoming. 
Within Wyoming three previous collections represent the species in 
Carbon and Albany Counties, all within 30 miles of the Colorado state 
line. A very substantial northern range extension is represented by 
plants found in a wet sedge meadow in the head of Wyoming Creek 
(10 August 1960, Johnson 232). 


JUNCUS ALBESCENS (Lange) Fern. (J. triglumis L.) and J. CASTANEUS 
J. E. Smith are known in Wyoming by one previous collection, both 
from the Medicine Bow Mountains, Albany County. Both species are 
low Arctic-Alpine species known in Colorado. It is not surprising, then, 
to find them in wet stream gravel with Juncus biglumis L. in northern 
Wyoming. The collections are: J. albescens, Johnson 221A, 31 August 
1959; 187B, 29 July 1960; 205B, August 1960; 235, 10 August 1960; 
J. castaneus, 121B, 31 August 1959; and 186, 29 July 1960. All speci- 
mens were collected from wet gravel on solifluction terraces in the head 
of Wyoming Creek; the two species are usually found together. 

Specimens of these species are deposited in the Rocky Mountain 
Herbarium, University of Wyoming, Laramie, Wyoming. All Johnson 
collections cited without initials are those of the author. Help with the 
identifications was received from C. L. Porter, A. E. Porsild, and Reed C. 
Rollins and is gratefully acknowledged. Appreciation is also expressed 
to the National Science Foundation for financial support under a N.S.F. 
grant (G-5574, W. D. Billings, Environmental Biology). 


Department of Botany, 
University of Wyoming, Laramie. 


1962 | PAYNE: AMBROSIA 233 


LITERATURE CITED 


Dant, E. 1951. On the relation between summer temperature and the distribution of 
alpine vascular plants in the lowlands of Fennoscandia. Oikos 3:22-52. 

HarrincTon, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver. 
666 pp. 

Jounson, P. L. and W. D. Brtincs. 1962. Alpine vegetation of the Beartooth 
Plateau in relation to cryopedogenic processes and patterns. Ecol. Monog. (In 
Press). 

Lewis, M. E. 1958. Carex—its distribution and importance in Utah. Brigham Young 
Univ. Biol. Ser. 1(2): 1-43. 

Porsttp, A. E. 1952. Illustrated flora of the Canadian Arctic Archipelago. Nat. 
Museum Canada Bull. 146: 1-209. 

Rotting, R. C. 1953. Draba on Clay Butte, Wyoming. Rhododora 55: 229-235. 

. 1961. Personal communication, March. 


THE UNIQUE MORPHOLOGY OF THE SPINES OF AN ARMED 
RAGWEED, AMBROSIA BRYANTIT (COMPOSITAE)! 


WILLARD W. PAYNE 


The true ragweeds, wind-pollinated composites of the genus Ambrosia, 
include our most serious hay fever plants. They are generally herbs or 
subshrubs. Their indument commonly consists of delicate hairs and 
glands, although some species become more or less hispid. Spines are not 
characteristically borne on the vegetative body of ragweeds, in spite of 
the fact that most ragweed species are found in open or disturbed habitats 
where spiny plants are common. The species to be discussed, A. bryantii, 
is interesting not only for possessing spines, but for the nature of the 
spines themselves, which, to my knowledge, are unique among similar 
structures in vascular plants. 

Armature of plants is accomplished in a number of ways, and the fact 
that many unrelated species possess spines is frequently used in teaching 
to illustrate convergent evolution. With the exception of the case of A. 
bryanti, presented below, spines which serve to protect the plant (thorns, 
prickles and other spine-like structures being included here under the 
term “‘spines”’) are formed from organs and tissues which are not directly 
associated with the flowers or fruits. They may be modified leaves (Ber- 
beris thunbergu DC.), leaf margins (Cirsium spp.), stipules (Robinia 
pseudoacacia L.), lateral branches (Gleditsia triacanthos L.), terminal 
shoots (Rkamnus cathartica L.), or epidermal emergences (Rosa spp.). 
Only one near relative of Ambrosia is spiny, i.e., Xanthium spinosum L. 
In this species the spines appear to be modifications of the two prophylls 


1 Publication Number 22 on atmospheric pollution by aeroallergens, under re- 
search grant Number E-1379 from the National Institute of Allergy and Infectious 
Diseases, Public Health Service. Thanks are due Mr. D. M. Porter who supplied the 
specimens for the drawings, and Dr. W. H. Wagner, Jr. who helped in preparing the 
manuscript. 


234 MADRONO [Vol. 16 


of the lateral branches, each prophyll being entirely changed to a three- 
branched spine. 


In addition to such vegetative spines, which are more or less perma- 
nently associated with the plant, many species produce spiny fruits. 
Fruit spines may be of value to the organism as a means of dissemina- 
tion of the seeds, or as protective structures which tend to prevent animals 
from eating the developing embryos, or often they serve both functions. 
Ordinarily, however, such spiny fruits are not retained by the parent 
plant as protective organs for the plant per se, but are shed when the 
seeds have developed. 


In all respects the species under discussion here conforms to the genus 
Ambrosia, although I fail to find that the proper name combination has 
been made. Curran (1888) placed it in the genus Franseria before the 
true nature of the group was known. It was placed in the monotypic genus 
Acanthambrosia by Rydberg (1922) on the basis of having more than 
one achene per fruit. As Shinners (1949) pointed out, however, characters 
of this nature are not sufficient to distinguish genera in the Ambrosieae. 
Accordingly, the new combination is made below: 


Ambrosia bryantii (Curran) Payne, comb. nov. Franseria bryanti 
Curran. Proc. Calif. Acad. ser. 2, 1:232. 1888. Acanthambrosia bryanti 
(Curran) Rydb.,-N.Am: Plo33222. 1922. 


In the genus Ambrosia, considerable modification of the floral struc- 
tures has occurred. Pollen and fruit production are carried out by differ- 
ent heads on different parts of the plant. Staminate heads are borne in 
spikes at the tips of the branches. Each head consists of a cluster of cen- 
tripetally developing, sterile flowers partially enclosed by a cup-shaped 
involucre, the phyllaries of which are fused laterally. Pistillate heads are 
found in the axils of leaves and bracts located below the staminate spikes. 
The pistillate flowers are borne singly or in clusters of from two to five. 
The involucre of the pistillate head has become concrescent, the phyllaries 
being united to form a hard, resistant, flask-shaped structure within which 
the achenes are borne. The tips of the phyllaries which form the involu- 
cral case are usually represented by more or less prominent spines. The 
spines may be blunt or sharp, straight or hooked at the tips, but in all 
species except A. bryantii they are short, usually shorter than the 
body of the fruit (fig. 1,C). The pappus is entirely lacking on both the 
male and the female flowers. Thus the fruit consists of one or more 
achenes enclosed by the spiny, indehiscent covering formed by the in- 
volucre of the pistillate head. In most species all of the fruits are shed at 
the end of the growing season, or as rapidly as they mature. 


Ambrosia bryantii is found on the desert plains of central Baja Cali- 
fornia, Mexico, where it is common and often quite abundant. It forms a 
small, perennial shrub which bears clusters of long, chalky spines along 
its stems. These spines are usually more abundant toward the stem apices 


1962 | PAYNE: AMBROSIA 


Bho 
WN 
OU 


Fic. 1. A, habit sketch of Ambrosia bryantii showing spiny aspect of a branch; 
B, single fruit of A. bryantii showing the long spines at the apex of the fruit; C, fruit 
of giant ragweed, A. trifida L., a common, annual species of the eastern United States. 
A and B drawn mainly from D. M. Porter 451, from 29 miles south of El Crucero, 
Baja California, Mexico. 


where one commonly finds inflorescences in all stages of development 
(fig. 1,A). When examined closely, these spines are seen to be borne on 
the fruits (fig. 1,B). The spines appear in every way to be homologous 
with the processes of the pistillate involucres of other species of Ambrosia. 
In A. bryantu, however, they are greatly exaggerated, forming very sharp 
spines 1.5 to 3.5 cm. long, with a basal diameter of 2 to 3 mm. 


236 MADRONO [Vol. 16 


The unusual and significant fact is that some of the fruits remain per- 
manently attached to the plant through several growing seasons. Exam- 
ination of many specimens has shown that a certain number of the fruits 
which develop during the perennial growth of the plant are thus retained 
and serve the function of armature. 

In summary, the spines of A. bryant represent what appears to be 
a unique morphological type of protective device, at least in the North 
American flora. They are actually borne on the fruits of the plant. The 
whole plant tends to become spiny because some of the fruits remain per- 
manently attached to the stems. This unusual armature of A. bryantii 
adds another striking illustration to the many examples of convergence 
in the evolution of vascular plants. 

Department of Botany 


The University of Michigan 
Ann Arbor, Michigan 


LITERATURE CITED 


Curran, M. K. 1888. Botanical Notes I: Plants from Baja California. Proc. Calif. 
Acad. ser. 2, 1:227-269. 

RypBerc, P.A. 1922. Carduales: Ambrosiaceae, Carduaceae, in N. Am. Fl. 33(1): 
1-46. 

SHINNERS, L.H. 1949. Notes on Texas Compositae. III. Field and Lab. 17:170-176. 


NOTES AND NEWS 


The following publications are of interest. 


Drawings of British Plants, by Stella Ross-Craig. Since mention was last made in 
Maprono of this beautifully executed series, the following numbers have appeared: 
Part XIII. Umbelliferae (2), Araliaceae, Cornaceae, 30 plates, 1959 [this completes 
Volume IV which comprises Parts X—XIII, 40 shillings. Cloth bound]. Part XIV. 
Adoxacaeae—Dipsacaceae, 39 plates, 1960. Part XV. Compositae (1), 28 plates, 
1960. Part XVI. Compositae (2), 33 plates, 1961. Part XVII. Compositae (3), 36 
plates, 1962. G. Bell and Sons, Ltd., London. Parts XIV and XVII are quoted at 
10/6; the others at 9/6. The publishers state that photographs or blocks may be 
obtained for reproduction purposes. 


Arizona Flora, by Thomas H. Kearney, Robert H. Peebles, and collaborators. 
Second edition, 1085 pp. University of California Press, Berkeley and Los Angeles. 
1960. A fifty-page supplement by John Thomas Howell and Elizabeth McClintock 
of the California Academy of Sciences, and collaborators, provides material accumu- 
lated since the publication of the first edition in 1951. Three species (Typha angusti- 
folia, Potamogeton richardsonii, and Elatine californica), reported in the January, 
1961, issue of MApRONO as new to the flora of Arizona, could have been included 
in the Supplement had the editors of MApRONo realized the imminent appearance 
of the second edition. 

Ecosystems of the East Slope of the Front Range in Colorado, by John W. Marr. 
University of Colorado Studies, Series in Biology, No. 8, pp. 1-134. University of 
Colorado Press, Boulder, November, 1961. 

The Systematics of Oenothera, Subgenus Chylismia, by Peter H. Raven. Univer- 
sity of California Publications in Botany 34 (1): 1-122. University of California 
Press, Berkeley and Los Angeles, 1962. 


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. 


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Hosting 


MADRONO 


VOLUME 16, NUMBER 8 OCTOBER, 1962 


Contents 


PAGE 
Factors INFLUENCING SURVIVAL AND GROWTH OF A 
SEEDLING POPULATION OF ARBUTUS MENZIESII IN 
CALIFORNIA, John Pelton 237 


A NEw SPECIES OF DOWNINGIA, 
John H. Weiler 256 


THREE NEW SPECIES RELATED TO MALACOTHRIX 
CLEVELANDII, William S. Davis and Peter H. Raven 258 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 266 


Review: Anne Ophelia Todd, The Little Hill, a chronicle 
of the flora on a half acre at the Green Camp, 
Ringwood, New Jersey (H. L. Mason) 268 


NoTEs AND NEws: WYOMING PINYON REVISITED, Roger 
S. Peterson; A CONTROVERSIAL TREATMENT OF THE 
PoLEMONIACEA4E, Edgar T. Wherry; CNEORIDIUM DU- 
MosuM (NUTTALL) HOOKER F. COLLECTED MARCH 
26, 1960, AT AN ELEVATION OF ABOUT 1450 METERS ON 
CERRO QUEMAZON, 15 MILES SOUTH OF BAHIA DE Los 
ANGELES, BAJA CALIFORNIA, MEXICO, APPARENTLY 
FOR A SOUTHEASTWARD RANGE EXTENSION OF SOME 
140 Mites, Retzd Moran; PUBLICATIONS OF Marcus 
E. JoNES AVAILABLE, Robert Ornduff. 269 


INDEX 273 


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 
EpGAR 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 
Miuprep E. Matuias, University of California, Los Angeles 24 
Marion OWNBEY, State College of Washington, Pullman 
REED C. RoLiins, Gray Herbarium, Harvard University 
Tra L. Wiccrns, Stanford University, Stanford, California 


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


Business Manager and Treasurer—JoHN H. THomas 
Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Roxana S. Ferris, Dudley Herbarium, Stanford University, Stanford, 
California. First Vice-President: Edward C. Stone, School of Forestry, University 
of California, Berkeley. Second Vice-President: Thomas C. Fuller, Department of 
Agriculture, State of California, Sacramento. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley. Corresponding Sec- 
retary: Margaret Bergseng, Department of Botany, University of California, Berke- 
ley. Treasurer: John H. Thomas, Dudley Herbarium, Stanford University, Stanford, 
California. 


1962 | PELTON: ARBUTUS SEEDLINGS 237 


FACTORS INFLUENCING SURVIVAL AND GROWTH OF A 
SEEDLING POPULATION OF ARBUTUS MENZIESII 
IN CALIFORNIA 


JoHN PELTON 


Differential survival in the seedling stage of potentially competing 
species is widely recognized to be of major significance in determining 
vegetational patterns and succession. Furthermore, an understanding of 
seedling ecology is essential to evaluate relative adaptation of different 
life cycle stages of a species, and to understand the mechanics of natural 
selection in plants. A brief review of these problems has been previously 
published by the writer (Pelton, 1953). 

Arbutus menziesii Pursh, Pacific Madrone, is an ericaceous tree rang- 
ing from southern California to southern British Columbia. Tarrant 
(1958) has summarized much of what is known of the autecology of this 
species. The present study of a naturally-occurring seedling population 
of Arbutus was done from February to August of 1958 in the Santa Cruz 
mountains of central coastal California. 

Grateful acknowledgment is extended to Dr. Victor C. Twitty, Head 
of the Department of Biological Sciences, and to Dr. Ira L. Wiggins, for- 
mer Director of the Natural History Museum, both of Stanford Univer- 
sity, for kindly providing facilities; to Mrs. Roxana S. Ferris, Dr. John 
H. Thomas, and Dr. Wallace Ernst for aid in plant identifications; to 
Dr. H. N. Hansen and Dr. J. R. Parmeter of the Department of Plant 
Pathology of the University of California, and to Dr. W. W. Wagener of 
the Division of Forest Disease Research of the Southwestern Forest and 
Range Experiment Station for examinations of pathological material; 
and to my wife, Jeanette S. Pelton, for help throughout the study. 


GENERAL METHODS 


The individual fates of several hundred emerging Arbutus seedlings 
were followed over a six month period, and mortality was correlated with 
observed or measured microenvironmental factors. Smaller numbers of 
seedlings of a few other species, especially Sequoia sempervirens (Lamb. ) 
Endl. and Heteromeles arbutifolia (Ait.) M. Roem., also occurred in the 
plots and were included in the study. 

The study area consisted of several hectares of second-growth Arbutus- 
Quercus-Sequota forest on the northeast-facing slope of the Santa Cruz 
mountains, at about the 184 meter contour on Martin Creek. The plots 
were situated on both sides of the Old La Honda Road which bisects the 
area, 0.8 km. beyond the intersection with Portola Road, in San Mateo 
County. 

Fifteen plots 30 cm. on an edge were established arbitrarily where seed- 
lings were emerging in large numbers and representing the range of micro- 


Maproxo, Vol. 16, No. 8, pp. 237-276. October 25, 1962. 


238 MADRONO [Vol. 16 


habitats in the study area. All seedlings were individually staked and 
charted to insure re-identification. Twice a week for most of the study 
period each seedling was examined and probable causes of mortality eval- 
uated. Measurements of selected seedlings were made once a week, and 
the root systems of representative individuals adjacent to the plots exca- 
vated monthly. 

Environmental measurements were made within the seedling stratum 
of air and soil temperatures, precipitation, vapor pressure deficit, evap- 
oration, light intensity, soil moisture, pH, wilting percentage, and other 
conditions. In addition, rodent and bird exclosures were utilized for cer- 
tain plots, and the vegetational structure and composition analyzed. 
Details of all these procedures are described below. 


RESULTS 


VEGETATIONAL ANALYSIS. The results of the vegetational analysis are 
summarized in Tables 1 and 2. The study area was not homogeneous, but 
ranged from a closed dense Arbutus-Sequoia forest above the road 
(“shade plots,” fig. 1) to a semi-open and lower Arbutus-Quercus forest 
below the road (“sun plots”). Plots which were intermediate in environ- 
ment were grouped in a third category (‘‘all other plots”). Both evergreen 
(QO. agrifolia Nee) and deciduous (Q. kelloggii Newb.) Quercus species 
were of significance in the latter two forest types. These forests were on a 
northeast-facing slope of a narrow ravine, and graded into chaparral on 
adjacent ridges and south-facing slopes. 

The entire area had been both lumbered and burned, the latter having 
been indicated by fire scars and multiple trunks. The closed forest in 
which the shade plots were located had almost no saplings of any species. 
The semi-open sun plot forest showed considerable numbers of Arbutus 
and Quercus seedlings and saplings. Less common trees not listed in 
Table 1 included Aesculus californica (Spach) Nutt., Umbellularia cali- 
fornica (Hook. and Arn.) Nutt., and Pseudotsuga menziesu Franco. The 
rarity of the latter tree and the almost complete absence of Sequoia seed- 
lings suggest that a conifer overstory will not dominate the study area in 
the foreseeable future. 

Tall shrubs (averaging over 1.5 m. at maturity) were locally abundant 
on the sun and miscellaneous plots, but rare on the shade plots. Hetero- 
meles arbutifolia and Corylus californica (A. DC.) Rose were most fre- 
quent, with occasional individuals or clumps of Ceanothus sorediatus 
Hook. and Arn., Rhkamnus californica Esch., Holodiscus discolor v. fran- 
ciscana (Rydb.) Jepson, and Sambucus mexicana Presl ex DC. Low 
shrubs included mostly Svmphoricarpos mollis Nutt., Lonicera hispidula 
v. vacillans A. Gray, and Rhus diversiloba T. & G., with occasional Rosa 
gymnocarpa Nutt. and Rubus vitifolius subsp. ursinus (Cham. & Sch.) 
Abrams, all least abundant again in the shade plots. Herbaceous species 
included Trientalis latifolia Hook., responsible for most of the density 
figures for “herbs” in Table 2, Drvopteris arguta (Kaulf.) Watt., which 


1962 | PELTON: ARBUTUS SEEDLINGS 239 


Fic. 1. View of forest in an area where Arbutus dominates over one of the shade 
plots (not visible). 


240 MADRONO [Vol. 16 
TABLE 1. SUMMARY OF THE TREE STRATUM WITHIN THE STUDY AREA 
Density per hectare 
Saplings 
less than Trees? over Basal 
2.5 cm. DBH! 2.5 cm. DBH area of Fre- 
trees quency? 
Over 1 over of 
year & 10 cm. trees 
under Over Over DBH over Aver- 
PLOT 30 cm. 30cm. 2.5-10cm. 10c¢.m. (sq. m. 10 cm. age % 
LOCATION TREE SPECIES tall tall DBH DBH per ha.) DBH cover 
A. menziesii ) @) @) 561 49.0 60 49 
SHADE Q. agrifolia @) 239 0 0 0 0 0 
S. sempervirens ) 0 Se 478 90.7 60 42 
Totals @) 239 79 1039 139.7 94° 
A. menziesii 1580 400 99 200 34.2 25 38 
SUN Q. kelloggii 99 498 197 200 6.9 50 15 
Q. agrifolia 988 597 99 200 62 50 24 
Totals 2667 1495 395 600 47.3 tS 
ALL A. menziesii 0 67 0) 1065 Be) 83 59 
OTHER Q. kelloggii 67 200 0 133 30 17 2 
PLots Q. agrifolia 534 400 0 67 ile | iby) 12 
Totals 601 667 0 1265 58.0 85° 


* Diameter at breast height. 4 Of sprout, not seedling, origin. 
2 Or multiple trunks. 5 Includes minor species not listed. 
3 Percentage of 5 X 5 meter plots occupied. 


was absent from the sun plots, and Bromus laevipes Shear., Satureja doug- 
lasu (Benth.) Brig., and Pityrogramma triangularis (Kaulf.) Maxon, all 
three absent from the shade plots. The moss stratum, composed of several 
mosses and an Anthoceros sp., was present only on the sun plots where 
litter was scant. Mineral soil exposure averaged 8% on the sun plots and 
0% on the shade plots. Nearly 30 additional vascular plants not listed 
above also occurred in the study area. Voucher specimens for most of 
these, including several seedling stages of Arbutus and certain other spe- 
cies, are in the herbaria of the writer or Butler University. 

The average maximum height of the Arbutus canopy was 15 m. in the 
dense Arbutus-Sequoia phase, with occasional Sequoia emerging to a maxi- 
mum of 32 m. The maximum height of the Arbutus-Quercus canopy aver- 
aged only 10 m. The 75% cover provided by the latter forest was at its 
maximum during the major part of this study; the deciduous Q. kelloggi 
had fully leafed out by the end of March, and the forest canopy did not 
begin to open significantly until Arbutus began abscising part of its foliage 
in August. 

In general terms, that portion of the study area referred to as Arbutus- 
Quercus is comparable to the “Broad Sclerophyll Forest” of Cooper 
(1922), the “Black Oak-Madrone Forest” of Mason (1947), the “Mixed 
Evergreen Forest” of Munz and Keck (1959), Whittaker (1960), and 
Thomas (1961a), and the ‘“‘Woodland” of Jensen (1939). The “shade”’ 
plots, on the other hand, are transitional with the ““Redwood” (Sequoia) 


1962 | PELTON: ARBUTUS SEEDLINGS 241 


TABLE 2. SUMMARY OF SHRUB AND HERB STRATA WITHIN THE STUDY AREA 


AVERAGE 

SHRUB AND AVERAGE DENSITY PER CENT 

PLOT LOCATION HERB STRATA PER SQ. METER COVER 
Tall shrubs 02 | 

SHADE Low shrubs 14 «+ 27 
Herbs 6 | 
Moss ea 0 
Tall shrubs ale 

SUN Low shrubs 45 } 36 
Herbs 6 J 
Moss ae 16 
Tall shrubs ae) 

ALL OTHER PLOTS Low shrubs 7) |e 64 
Herbs 34. | 
Moss se 0 


forest of most of these authors. The entire study area is mapped as poten- 
tial conifer cropland by the Forest Service (1950). 

A number of vegetational and environmental studies have been made 
in the Santa Cruz mountains near the study area, including the classic 
researches of Cooper (1917, 1922) as well as the more recent ones of 
Springer (1935), Moeur (1948), and Oberlander (1953). Floristic work 
in the area has been recently reviewed by Thomas (1961b). 

Puysicat Factors. Measurements of certain environmental conditions 
near the sun and shade plots are in figures 2 and 3. Where practicable, 
measurements were made within the air or soil strata in which the seed- 
lings actually grew. Air and soil temperatures were recorded with Six’s 
type maximum-minimum thermometers, the former at 2 cm. above the 
soil or litter surface and the latter at 3 cm. below this surface. A small ven- 
tilated reflective metal shelter was employed for the air thermometers. 
Maximum air temperatures on sun plots (Station 1, lower graph of fig. 2) 
reached 42°C. on July 15, but although actual surface soil temperatures 
were much higher, no stem girdle of seedlings resulted owing to hypo- 
cotyls having long since passed the “‘succulent stage” (Baker 1950:255). 

Minimum air temperatures were likewise reached in the semi-open 
Station 1, but at no time did they fall below 2°C. Frost was therefore not 
a factor in mortality even though seedlings were emerging during the 
normally coldest part of the year. The nearest official weather station 
having long term temperature records (Redwood City) suggests that Jan- 
uary and February of 1958 were appreciably warmer than normal al- 
though there was only slight deviation during the remainder of the study 
(United States Department of Commerce 1959). Under dense Sequoia 
canopy (Station 2) the maximum temperatures were greatly depressed 
but the minimum temperatures were only slightly raised, resulting in a 
much smaller diurnal variation. Soil temperatures at 3 cm. (upper graph 


242 MADRONO [Vol. 16 


STATION | MAXIMUM \ yea 


STATION 2 dee.) 


80 


ae) 
aA 
Harte i eo 
77 y eo 
TION 2 MINIMUM 


y 
2 
& 


STA 


60 


50 


STATION | 


MINIMUM 
40 


oF °C 
40 
100 
35 
~ STATION | MAXIMUM 
30 as 
STATION 2 MAXIMUM 
80 
25 R ha 
/ / N 
Pea? prada Aen ye 
7 Boe Cant ane or 
Naa <0 Near), v 
20 . N 
y v 
607 15 


STATION 2 MINIMUM =~ 
y 0 ed 
50 


9 26 4 Ul i8 25 1 8 15 22 29 6 i) ANG 17 
FES. MARCH APRIL MAY NE 


20 27 3 24s Bul P 22 29 


Fic. 2. Soil temperatures (above) at 3 cm., and air temperatures (below) at 2 cm. 
Station 1 semi-open, Station 2 in deep shade. 


fig. 2) showed a pattern similar to that of air temperatures except for a 
tempering of the extremes. It is significant, however, that temperatures 
of 33°C. were reached at this level twice at Station 1. Such high tempera- 
tures at this depth help explain the soil drought which rapidly obtains in 
the upper soil level of semi-open areas following rains (fig. 3). 

Vapor pressure deficit measurements were taken on each visit to the 
area among the sun plots between 12 noon and 1 p.m., as close to the 
ground as a small sling psychrometer could be used (about 30 cm.). There 
was a detectable upward trend as the season progressed (fig. 3), but the 
summer advection fogs from the ocean characteristic of the Sequoza belt 
(Cooper 1917) raised the humidity significantly above that which would 


1962 | PELTON: ARBUTUS SEEDLINGS 243 


Ww 
Sf 
eo te 
Ww 
at 6 
S) 
w 
4 
eo 
< 
<7 50 
15 
_ le 
a 
Se 
Hea z~ 
ry 6 Ox 
° r& 
a. <e 
<2 3 ro 
zz S 
ft) ga 
a~ 
cM. 
50 
Wt 
“40 
« PLOT 14 12-15 CM. mg 
K- 
” 30 8 
3 rh oe = 6 
oO eee ares me 
: kee po ge oa eee y 
ro) PLOT II 12-15 cm. A Sav 
107 1] [MV tb | 0 SSO geese. 2 
e f WT |® TUB STE TOS 
9 26 4 tt #@ 25 | 8 IS 22 29 6 13 20 27 3 10 7 24 | +8 I5 22 29 
FEB. MARCH APRIL MAY JUNE JULY 


Fic. 3. Vapor pressure deficit, evaporation, precipitation, and soil moisture. Plot 11 
in semi-open, plot 14 in shade. Shaded bars represent precipitation under dense can- 
opy, unshaded bars under an opening, only the latter record being available on Feb- 
ruary 26. 


otherwise prevail. Although the study area was near the inland edge of 
the fog belt, this factor is doubtless significant in prolonging survival in 
the face of intensifying summer drought. In addition, several light show- 
ers in May and June had a detectable effect on depressing the vapor pres- 
sure deficit. 

Evaporation data as shown in figure 3 represent the average of corrected 
readings of two standardized and periodically checked spherical porcelain 
atmometers placed with the base of the spheres 10 cm. above the soil 
surface among the sun plots. The figures represent the maximum evapora- 
tion stress to which the seedlings were subjected. Again, an upward trend 
is apparent in spite of depression during rainy and foggy periods. 

Precipitation data were obtained from two 7.5 cm. diameter rain gauges, 
one under an opening in the canopy among the sun plots, and the second 
under complete cover adjacent to the shade plots. Precipitation was con- 
centrated in winter and early spring, as is normal for Mediterranean-type 
climates. Excluding the February rain, at which time only one gauge was 
operating, an average of 30% more precipitation was received under the 
light than under the dense canopy. Since the gauges were but 50 m. apart 
and in similar positions with regard to topography, rainfall interception 
by the vegetational canopy was probably responsible for this difference. 
Official precipitation records from nearby weather stations indicate that 
the rainfall during the study period was 2.7 times normal, reflecting the 
unusually wet year for the region. 


244 MADRONO [Vol. 16 


Light intensity was measured photoelectrically by reflection for each 
seedling plot. This was done five times between 9 a.m. and 3 p.m. on a 
clear day in early May. The resulting foot candle figures were arbitrarily 
corrected for estimates of the cover of low vegetation below the level at 
which readings could be made. The final results facilitated the distinction 
of “‘sun” from “shade” plots, although not permitting expression in abso- 
lute terms. In the former plots, sun reached the seedlings a small to mod- 
erate part of each day, but only an occasional sun fleck penetrated to the 
shade plots. 

Soil moisture determinations were made adjacent to six sun and shade 
plots at two-week intervals. Two depths were sampled in duplicate with 
a soil tube, the top 3 cm. of mineral soil, and the depth range of 12 to 
15 cm. Data for one representative shade plot (Plot 14) and one sun plot 
(Plot 11) are given in figure 3. Greenhouse determinations of permanent 
wilting percentages (method of Daubenmire, 1959) for these soils gave 
the following results: 


DATE WILTING 
FIRST REACHED 


WILTING IN FIELD 
PLor DEPTH (cCM.) (PER CENT) (PER CENT) 
14 1-3 19.8 July 29 
14 12-15 1320 “ 8 eae es 
11 1-3 18.5 May 21 
11 12-15 11.8 June 17 


In marked contrast to Plot 11, Plot 14 reached the wilting percentage 
only by July 29 at the surface and at no time deeper than this. Soil drought 
arrived much earlier and reached a greater intensity on sun than shade 
plots. 

The contrast of mineral soil drought does not take into account the 
litter through which seedling roots often did not penetrate. The sun plots 
were all similar in having usually less than 1 cm. of litter, often exposing 
mineral soil or moss on small hummocks. Litter on shade plots averaged 
5 cm., but reached 8 cm. on Plot 14. Although the but slightly decom- 
posed (mor-type) litter was thicker under Sequoza, the large coriaceous 
leaves of Arbutus often forced roots to grow horizontally for long periods. 
Litter of sun plots was dry to the touch within a day following rains, but 
the lower layers of shade plot litter were moist in July on some plots. It is 
clear that the “sun” and ‘‘shade”’ plots differed in other important condi- 
tions than light alone. The shade plots can best be viewed as densely 
shaded sites with heavy litter and moist mineral soil, while sun plots are 
those of moderately open sites with thin litter, if any, and dry soil. Past 
and present soil disturbance by rodents was also much more important 
in the semi-open than in the shade, and the incidence of injurious inverte- 
brates and fungi also differed. 

The soils in the study area are classifiable as the “rough broken phase”’ 
of the Altamont clay loam, a residual group largely derived from inter- 
bedded sandstones and shales, and retentive of moisture but well drained. 
Rocks are uncommon, and a B horizon is not distinguishable. Electro- 


1962 | PELTON: ARBUTUS SEEDLINGS 245 


a 

au 

Re 

< 100 

z e——e SHADE PLOTS 

= x 

a« 90 oO 

iy oa o-—-o SUN PLOTS 
80 so. 

° 5. Qa ALL OTHER PLOTS 

oc 

Ww 

a 


SURVIVORS 


19 26 4 ul ig@ 25 | 8 15 22 29 6 13 20 27 3 10 17 24 1 8 15 22 29 
FEB. MARCH APRIL MAY JUNE JULY 


Fic. 4. Survivorship curves of Arbutus on log scale (top) and arithmetic sca‘e 
(below). 


metric pH determinations of the top 3 cm. of mineral soil ranged from 
5.7 to 6.6, averaging 5.96. At 15 cm. the range was 4.7 to 6.5 with a mean 
of 5.65. The surface soil pH of the thin-littered sun plots averaged slight- 
ly lower (5.9) than that of the thickly covered shade plots (6.2); this 
anomaly probably resulted from rodents mixing acid subsoil with topsoil 
on only the sun plots. 

Mortatity. Arbutus seedling mortality results are presented graphical- 
ly in figures 4, 5, and 6. The survivorship curves in figure 4 compare the 
seasonal advance in mortality on shade, sun, and all other plots. Since a 
straight line on a log scale indicates a constant mortality rate, the top 
curves show that such a condition prevailed only during certain periods 
in each habitat. The greater mortality in the shade at almost all times is 
evident in comparison with the sun plots. 

Most Arbutus seedlings had already germinated by the time the study 
was initiated on February 19, but all were in the early cotyledon stage. Of 
the ultimate total number, 92% had germinated on the sun plots, 77% 
on the shade plots, and 86% on all others. The remaining seedlings ap- 
peared sporadically up to April 8 on the sun plots and April 15 in the 
shade. Consequently, the curves of figure 4 are expressed as survivors per 
100 germinated rather than as survivors of an initial maximum popula- 
tion. A total of 829 Arbutus seedlings approximately equally divided be- 
tween the sun, shade, and other plots varied from 20 to 108 per plot 
30 cm. on a side, or 222 to 1200 per square meter. 


MADRONO [Vol. 16 


246 

o— INVERTEBRATES 

100 x g ? 

o—-© DROUGHT A h J 

90 I I ! 

ara FUNGI i I I 

80 : \ in| | 

%--k COMBINATIONS 1a KR. 1 \ I 

70 LSS i \ fea ] 
= eunne MISCELLANEOUS ON ne 1 te x | \ | 
F 60 Mame Re BNR ee ae 
= Came Uf Veal \E2 ! vy \ f 
& 50 x ec ee Cae 
= i: a ‘ \ 

- * BO. yy, ee 
w : Pe ie Nea I 
=< 30 As Nts Sif 2 [: \ ! 
e Behe) i: z \ \ | 
= We See aE ee d : 

8 20 1 Be ee \ I 
x \ t, ; \ ! 
w J oce "th, \ | 
710 re ae fees y 
- \ \; As \ S “4, tS lo a8 J “MN, | \ ! 
PO Nangiafigh any dnytisy | ec eee Se ae ausrnennte 
13 20 27 3 #10 17 24 | 8 158 22 29. 
MAY JUNE JULY 


Fic. 5. Seasonal variations in causes of mortality of Arbutus seedlings on all plots 
combined, expressed as a percentage of total mortality for a given week. 


At the termination of the study on August 2 no Arbutus seedlings were 
present on the shade plots, 2“ had survived in the sun, and 1% on all 
other plots. Seedlings of other species which happened to occur on the 
plots, totalling 75, were also included in the study. Of these, Sequoia was 
represented by 32 individuals, all on the shade plots, of which none sur- 
vived beyond April 23. Twenty-one seedlings of Heteromeles were present 
on sun and other plots (none in the shade), one of which was living at the 
end of the study. There were no survivors from three seedlings of Sanicula 
crassicaulis Poepp. ex DC., three of Satureja douglasiu, or of eight uniden- 
tified plants of several other species. Four seedlings of Pyracantha sp. and 
one of an Acacia sp., probably carried in by birds from nearby gardens, 
also failed to survive. On the other hand all three seedlings present of 
Lonicera hispidula survived the entire period. Other species having occa- 
sional seedlings in the locality but not on the plots included what were 
probably Rhus diversiloba, Osmorrhiza chilensis H. & A., Rosa gymno- 
carpa, Rhamnus californica, Dentaria californica Nutt., Bromus laevipes, 
Sym phoricarpos mollis, and Hedera helix L. (probably bird-disseminated 
from nearby gardens). 
Figure 6 isa summary of Arbutus mortality under different conditions. 
The damage by invertebrates was probably mostly by slugs (Order Pul- 
monata). A small species of this group was found feeding upon seedlings 
on several occasions in early morning, although most damage was noctur- 
nal. Frequently the cotyledons were completely eaten, leaving the decapi- 
tated hypocotyl, and sometimes the entire shoot disappeared. Two plots 
were temporarily covered on sides and top with 3 mm. mesh hardware 
cloth exclosures to test for bird or rodent feeding, but without effect on 
injury by this cause. By comparing the data in figure 5 with that in figure 
3, it can be seen that mortality of this type correlated positively with pre- 
cipitation. It was also of greater relative importance on the shade plots 
than the sun plots (fig. 6). In addition, covering of seedlings by falling 
leaves increased their chances of being eaten. All these observations would 
tend to implicate slugs or other soil-dwelling invertebrates. A mite (Order 


1962 | PELTON: ARBUTUS SEEDLINGS 247 


100 
90 
> 80 
570 
eG SHADE SUN ALL OTHER 
a PLOTS PLOTS PLOTS 


1 23 4 5 | 2 3 4 5 1 2 3 
CAUSE OF DEATH a 


Fic. 6. Causes of mortality of Arbutus seedlings in different habitats. Numerals on 
horizontal line refer to: 1 = invertebrates, 2 = drought, 3 = fungi, 4 = combina- 
tions, 5 = miscellaneous. 


Acarina) was found in abundance on hypocotyls of several seedlings early 
in the season, but damage seemed slight. Mites have been reported to be 
vectors of damping-off fungi, however, and may have contributed to mor- 
tality from this cause (Wilde, 1958). Insects were only rarely found feed- 
ing on seedlings. A total of 29.2% of the total mortality on all plots was 
classified under the heading of invertebrates, slugs probably accounting 
for nearly the whole amount. 

Drought, when acting alone, is operative very soon after germination, 
and again, towards the end of the study, in July, when only a handful of 
seedlings remained (fig. 5). The early mortality, which included the great 
majority of the drought-killed plants, was due to recently emerged seed- 
lings being rooted only in rapidly drying surface litter, even though this 
mortality occurred at the height of the wet season. The presence of a thick 
litter on the shade plots and its absence on the sun plots explains the 
anomalously larger mortality from drought in the shade (fig. 6). Actually, 
there was no mineral soil drought even in the surface soil on the shade 
plots until July (fig. 3). The more extreme sun plots, on the other hand, 
lacked growth water (moisture above the wilting percentage) at the sur- 
face by late May and at 15 cm. by late June. So few seedlings had sur- 
vived by this time, however, that the toll from drought alone was very 
small (6% ) on the sun plots in terms of the initial number of seedlings. 
Total mortality on all plots from drought acting alone was only 10.4%. 

Death of Arbutus seedlings from fungus attack was more important 
quantitatively, comprising 28.1% of total mortality on all plots. This 
factor was positively correlated with precipitation, by June becoming of 
little importance as a factor in seedling mortality (figs. 3 and 5). Surpris- 
ingly, fungi were of much greater relative importance on sun than on 
shade plots. In part, this is a natural consequence of the lesser mortality 
from invertebrates on the sun plots, thereby leaving a larger proportion 


248 MADRONO [Vol. 16 


to be killed by other agents. Also, several etiological agents were involved 
in the fungus attacks, one of which occurred only in the sun. Fungus 
damage was classified in the field into three categories: (1) Typical post- 
emergence damping-off, with collapse of the hypocotyl (and usually root 
decay also); (2) Root decay alone; (3) Leaf spot, usually with root decay 
as well. On the sun plots these conditions were in the proportion of 39%, 
25‘~7, and 36%, respectively. Leaf spot was absent from shade plots, giv- 
ing a ratio of 59%, 26%, and 0%. The other plots were intermediate. 


From external appearances it is probable that most of the damping-off 
and root decay of the Arbutus seedlings was produced by water molds 
(Pythium, Phytophthora, etc.). The unusually heavy rains in March and 
April (fig. 3) correlated well with maximum seedling damage, as is normal 
for these fungi. In addition, R/izoctonia seemed also to be present on a 
number of seedlings, and semi-parasitic mycorrhizal fungi may have 
played a role as well. The leaf spot of Arbutus, which was present only 
on the seedlings growing on the sun plots, developed at the height of the 
March rainy period, but owing to its crippling effect, mortality was mostly 
delayed until April. The spots generally started as small necrotic areas 
which gradually enlarged until coalescence of several spots killed most of 
the cotyledon, although epicotyledonary leaves, the epicotyl, or the hypo- 
cotyl were often also affected. The disease was mainly concentrated on 
plants of a single sun plot with much exposed mineral soil. If the etiologi- 
cal agent were a bacterium which required splashing from mineral soil to 
the cotyledons by raindrops, the absence of the disease from the thick- 
littered shade plots might be understood. 


A total of 22.7% of all mortality was attributed to “combinations” of 
factors. Of this amount 67% was a consequence of rather mild drought 
preceded by crippling from root decay fungi. If fungi had not limited the 
depth and volume of root penetration, the seedlings presumably could 
have survived at least until the summer drought intensified. This category 
predominated after the rainy season when the weakened root systems 
resulted in drought-death (fig. 5). The effect of the light showers in May 
and June in temporarily reducing mortality from this cause is clearly 
shown. 

On the shade plots 25% of the “combination” category was considered 
to have been a consequence largely of deep shade, which predisposed the 
seedlings to death from another cause such as drying litter. Again, root 
fungi were usually involved here also, and occasionally mechanical injury 
from rains, falling leaves, leaf chewing, and other factors. Even on the 
most densely shaded plots, low light intensity was probably never by itself 
a cause of death during the period of study, the proximal or immediate 
cause of mortality always being some other factor. One completely albino 
seedling even survived for 25 days. 

An average of 9.6%. mortality was attributed to “miscellaneous” causes. 
Much of this category resulted from causes which were too uncertain to 
classify. On the sun plots, however, 67% was the result of mechanical 


1962 | PELTON: ARBUTUS SEEDLINGS 249 


disturbance by rodent undermining. Also included were seedlings covered 
by fallen leaves or exfoliating bark of Arbutus for extended periods. The 
average angle of slope of the plots (18.4°), however, resulted in a gradual 
shifting of the litter downslope such that a seedling would usually be re- 
leased within a few days and in such cases covering was considered a pre- 
disposing or contributing (“crippling”) rather than a proximal (or imme- 
diate) environmental factor contributing to mortality. Temporary cover- 
ing not only predisposes seedlings to injury from drought and fungi but 
during the moist season to attacks by slugs as well. Conversely, drought 
predisposes seedlings to covering by litter, since wilted seedlings are easily 
bent by falling leaves or even by heavy rains, and once bent horizontally 
are readily covered by downslope shifting of litter. 

On October 7, 1961, the study area was again revisited. The winter 
and spring of this year received less than half normal precipitation, and 
yet the number and size of visible Arbutus seedlings of the 1961 season 
was comparable to that observed in 1958 on and near certain sun plots. 
All but two seedlings, adjacent to one sun plot, were dead but had prob- 
ably survived to about mid-summer. The two living seedlings had 4 to 6 
live but wilted and reddish leaves (the cotyledons had dried) and prob- 
ably could have been expected to survive if the normal fall rains material- 
ized. Neither living nor dead seedlings were visible on the shade plots nor 
on any sun plot where mineral soil was not exposed, emphasizing again 
the inhibiting influence of litter as well as of shade. 

With regard to seedlings of other species than Arbutus, a similar com- 
plex pattern of mortality causes occurred. Of the Sequoza seedlings, 32% 
were eaten by invertebrates and 6% succumbed to drought in the rainy 
season as a result of rooting in thick litter which soon dried out in the 
upper layers. Damping-off and root decay fungi accounted for 13%, but 
at least 23% died from drying of the litter combined with attacks by root 
fungi. Other combinations (16% ) or miscellaneous (10% ) accounted for 
the remainder of the Sequoza seedlings by the early date of April 23. 

Over half of the Heteromeles seedlings dried primarily by direct fungus 
attack, either typical damping-off, or much more commonly (38% of total 
mortality from any cause) from a leaf spot apparently caused by Fusi- 
cladium dendriticum (Wallr.) Fckl. v. eriobotryae Scalia. Combinations 
of factors killed 14%, the remaining seedlings dying from miscellaneous 
causes excepting one survivor which was nearly dead from leaf spot and 
probably did not survive the year. 


GrowtTH. Weekly measurements of shoot length, cotyledon and leaf 
size, and hypocotyl diameter of several marked seedlings on each plot 
were made until death of the plants. Plants adjacent to and of size com- 
parable to those being measured were excavated monthly (fig. 7). Sev- 
eral points are noteworthy. First, the very small amount of growth of 
all Arbutus seedlings over the span of half a year is evident. Even by 
August the cotyledons had not died, although by then they were wilted, 
and only three to five nodes were visible by this time. The greater root 


250 MADRONO [Vol. 16 


system was present on plants in the sun, except where fungus attacks re- 
versed this relation. Rotting of the root tip often stimulated branching. 
A slight bend in the hypocotyl usually resulted when a seedling was bent 
by rain or leaf litter for a few days, although one of the “E” seedlings 
probably grew between two impenetrable leaves before becoming ver- 
tical. Leaf injury by slugs (or insects) was evident on several plants. In 
the sun the hypocotyls changed from pale green or yellowish-white to 
pink or even red in about a month, but remained pale in the shade for 
much longer. By early May in the sun hypocotyls began shrinking with 
the collapse of the cortex following periderm production, as is normal in 
woody species, becoming brown or dark red and hard and wiry. On 
the other hand in the shade the hypocotyls were still quite succulent even 
by June, and none survived there long enough to appreciably harden. 

Within a few months a number of the roots developed what appeared 
to be short, stubby mycorrhizal nodules (fig. 7), similar to the “root 
tubercles” in Arbutus unedo L. (Rivett, 1924). They occurred at the tips 
of certain main roots or on short branches, and developed into small and 
frequently branched pear-shaped tubercles enveloped in a dense mantle 
of hyphae from which minute root hair-like setae projected. Root hairs 
appeared to be absent from all naturally-occurring Arbutus menziesiu 
seedlings, which agrees with observations by Rivett (1924) on other 
species of the genus. 

Arbutus menziesii provides an interesting juvenile morphology which 
may not have been previously recorded. The epicotyledonary leaves of 
seedlings for several nodes are opposite, doubly serrate, glandular hairy, 
and thin (fig. 7), while mature foliage is alternate, essentially entire, 
glabrous, and coriaceous. Leaves of sprouts and saplings intergrade be- 
tween these extremes. 


DISCUSSION 

In spite of good seed crops being produced regularly (Tarrant, 1958), 
Arbutus seedlings are reported to be uncommon in comparison with 
stump sprouts (Jepson, 1910). On nearby Jasper Ridge, Cooper (1922) 
recorded no Arbutus seedlings in a Quercus-Arbutus-Aesculus forest. Ar- 
butus seedlings and saplings of a number of age classes were present on 
the study area (Table 1), however, but only in the semi-open forest, 
along road cuts, or at the bases of large fallen trees. Healthy two- to sev- 
eral-year old seedlings and saplings were also present under moderate 
Arbutus-Quercus canopy. Nevertheless, the classification of this species 
by a majority of silviculturists as ‘‘tolerant” of shade (Baker, 1949) is 
probably based entirely on the behavior of mature trees and stump 
sprouts rather than seedlings, and the writer agrees with the considerable 
segment of dissenting opinion reported by Baker to class the tree as “‘in- 
termediate,” based upon its local behavior. Even in the absence of gen- 
eral fire or other disturbance, Arbutus probably maintains its position in 
dense forests not only by stump sprouting but perhaps also by a process 
similar to “gap phase” reproduction described elsewhere by Bray (1956). 


1962 | PELTON: ARBUTUS SEEDLINGS Zoi 


Fic. 7. Representative seedlings of Arbutus showing the usual range of behavior 
found in the field. A to F represent monthly intervals from March to August. The 
seedling on the left of the letter is in each case from near one of the shade plots while 
that on the right is from near a sun plot. 


Shade can best be considered a predisposing factor increasing suscepti- 
bility of Arbutus seedlings to other more immediate causes of mortality, 
as has been found to be the case with other species (Baker, 1950: 266— 
267). In the present study, shade was always associated at least with a 
thick drying litter. Much earlier mortality occurred on the shade plots, 
in spite of the fact the mineral soil never reached the wilting percentage 
even at the surface at these sites (fig. 4). This is, however, partly due to 
the greater incidence of slugs on the shade than on the sun plots (fig. 6). 
The direct effects of shade were seen in the prolonging of the succulent 
hypocotyl stage and the usually smaller root-shoot ratio. Since one albino 
seedling survived for 25 days, we may assume that enough food reserves 


252 MADRONO [Vol. 16 


were present in green seedlings so that shade alone could not eliminate 
them before this time. 


Seedbed requirements of Arbutus menziesu elsewhere are not known. 
Sudworth (1908) states that germination is best in moist soil when seed 
is well covered, while lack of seed covering and drier soils inhibit seed- 
lings. There is apparently nothing in the nature of Sequoia or Arbutus 
litter which prevents germination. Instead, the ease of drying of upper 
litter layers (associated with extremely high wilting percentages of litter), 
in conjunction with the dense shade usually provided in such habitats, 
renders them highly unfavorable. Normal germination varies from Feb- 
ruary to April, depending on climate (Tarrant, 1958). The early (mostly 
February) germination in the present study probably reflects the unusu- 
ally warm January and February of 1958, and also the southerly loca- 
tion of the area in relation to the main range of the species. 


Germination of Arbutus is epigeous, and the cotyledons remain photo- 
synthetic for several months. The mesomorphic traits of the strikingly 
different juvenile foliage, although perhaps in part of a vestigial nature, 
probably are adaptive in certain respects. The dense glandular hairs may 
inhibit the occasional foliage-eating insects, but apparently have no in- 
hibiting effect on slugs. The thin, non-coriaceous, serrate leaf of the 
juvenile plants may also be photosynthetically more efficient at low light 
intensities than xeromorphic mature foliage which is usually more ex- 
posed to desiccation. 


The occurrence of drought-death as classified in figures 5 and 6 is mis- 
leading, since this mostly occurred at the height of the rainy season and 
on the shade plots, when a few sunny days would dry out the upper litter 
layers. Somewhat later, drought eliminated most of the seedlings which 
had been crippled by root fungi, often before even the upper layers of 
the mineral soil had dropped below the wilting percentage. But by June 
17 the sun plot soils had dropped below the wilting percentage at the 
12-15 cm. level. Although a healthy root system could probably pene- 
trate below this depth within four months, the usually continued drought 
for four or five more months usually presents an insurmountable ob- 
stacle to seedlings of most perennials. That a few Arbutus seedlings ac- 
tually do survive nearly every year is shown by the several seedlings and 
sapling age classes present on the area, usually in the same microsites in 
which the healthier current-year seedlings occurred (Table 1). Two or 
three exceptionally vigorous seedlings of the present season which oc- 
curred off the plots in favored sites appeared as if they might survive 
the summer, but all the Avbutus on the plots doubtless would not have 
survived many more weeks even had they not been excavated for root 
studies. The fact that two live 1961 seedlings were found near one plot 
near the end of the dry season in October, in spite of the unusual drought 
of this year, suggests that the abnormally heavy precipitation of 1958 
did not significantly reduce death from drought over that of more normal 
years, perhaps because precipitation was concentrated in early spring. 


1962 | PELTON: ARBUTUS SEEDLINGS 299 


Damping-off and root decay fungi have not been frequently incrimi- 
nated in studies of tree establishment under natural conditions, although 
they are often destructive in coniferous tree nurseries. The prevalence 
of these agents in this study, both as predisposing and as proximal causes 
of mortality, was probably in part associated with the unusually wet 
spring of 1958. A problem arises, however, in interpreting the relation 
of root decay and mycorrhizal fungi. It is possible that some of the root 
decay of seedlings was caused by the same fungus producing the tubercles. 
The tubercles resemble those of Arbutus unedo, where they are consid- 
ered to be ecto-endotrophic mycorrhiza in which the relationship is one 
of balanced parasitism rather than symbiosis (Rivett, 1924). If this is 
true also in A. menziesii, it would be understandable that the fungus 
might gain the upper hand during an unusually wet season and become 
a crippling parasite. It is probable, however, that the typical damping-off 
symptoms, and perhaps much of the root decay itself, were produced by 
other agents even if mycorrhizal fungi were also involved. 


Shade and summer drought are doubtless normally more significant 
in seedling establishment of Arbutus than is suggested by the histogram 
(fig. 6). Shade does not there appear among the causes of mortality be- 
cause it was classified as a predisposing condition under the heading of 
“combinations” of factors. Furthermore, on the shade plots, the early 
and complete mortality resulting from other conditions than shade pre- 
vented shade alone from exerting its full potential influence. In a similar 
fashion, summer drought was a minor factor in this study, most of the 
drought-killed seedlings of figure 6 having perished at the height of the 
wet season owing to a thick drying litter. Summer drought would be ex- 
pected to act as a final “coup-de-grace” for most of the few seedlings 
which have survived prior dangers and which are often crippled by fungi 
or shade-induced exhaustion of food reserves. The results of this study 
emphasize the importance of predisposing or contributing (‘“crippling’’) 
factors even though these may not be the proximal or immediate cause 
of death. 

With regard to Sequoia sempervirens, the present limited results con- 
firm the conclusion of Fritz (1950) that litter and shade do not prevent 
germination of this tree, but rather strongly inhibit survival. The present 
unusual results, however, which implicate fungi and invertebrates as 
being of greater importance than drought in causing mortality, may be 
due to the shady site and unusually wet spring. The Fusicladium leaf 
fungus of Heteromeles was also probably favored by this wet weather. 


In evaluating survival in seedling populations the following important 
questions are asked. To what extent are adaptive genetic differences in- 
volved in differential survival within seedling populations (‘‘natural se- 
lection” in the present sense)? Or, is seedling survival dependent pri- 
marily upon the chance vagaries of seed dispersal to physically favorable 
microsites (the “‘safe sites” of Harper e¢ al., 1961) combined with the 
subsequent chance absence of injurious biotic agents? 


254 MADRONO [Vol. 16 


-The difficulty of demonstrating a correlation between genotypes and 
survival in nature explains the paucity of field data supporting natural 
selection in plants (Stebbins, 1950: 106). No genetic races of Arbutus 
menziesu have been yet recognized within its range (Tarrant, 1958). 
Nevertheless, genetic differences were probably present among the Ar- 
butus seedlings in the present study which were significant to survival. 
This was obvious in the case of the few albino seedlings. Date of germina- 
tion ranged over almost a four month period in Arbutus, and probably 
involved both genetic and environmental factors. Late germination was 
conducive to drought injury and in fact no late germinating seedlings 
survived long. Arbutus cotyledons varied greatly in size and shape, prob- 
ably again partly owing to genetic differences; cotyledon structure cer- 
tainly is involved in shade tolerance. 


Presumably, distasteful biochemical products such as may be present 
in the glandular hairs of Arbutus seedlings could influence destruction 
by slugs. Possibly this feature helps explain why some seedlings of Ar- 
butus remained untouched adjacent to others which were thoroughly 
chewed. Also, adjacent seedlings of this species were not always affected 
similarly by fungus parasites. In view of the problem of root penetration 
of layers of rapidly drying litter as well as of their maintenance of con- 
tact with moist subsoil during the prolonged summer drought, any genetic 
variation promoting the development of a diffuse rather than a tap root 
system by Arbutus seedlings would be expected to be selected against. 
Injury of the tap root by fungi was one of the most frequent causes of 
mortality. 


The fact remains that at least all the “shade” plots were located in 
“unsafe sites” where the lack of seedlings or saplings of any but the cur- 
rent year showed that survival depended on thinning of the canopy and 
litter by fire, lumbering, or the falling of one or more trees. Under such 
conditions, the genotypes within the population could influence only the 
duration of survival by a few weeks, and natural selection, in the sense 
of either differential survival to reproductive age or differential repro- 
duction by the resulting survivors, has not occurred. Chance “catas- 
trophic” events, such as covering of seedlings by falling leaves or ex- 
foliating bark and perhaps much of the biotic injury, also were important 
“non-selective” factors, little influenced by seedling genotypes. On the 
sun plots, selective pressures would be expected to operate severely on 
the 2% survivors, although if all seedlings eventually succumbed before 
maturity the effects of natural selection would there also be nullified. 
Selection pressure, however, would be somewhat relaxed on these few 
survivors if the effects of the intensifying summer drought were more 
than compensated for by the reduced competition for light and moisture 
occasioned by the lower seedling density resulting from early high mor- 
tality. But in the present case this was probably not the situation, even 
if all survived, since seedling size was small and densities were only lo- 
cally high enough to result in much mutual shading or root competition 


1962 | PELTON: ARBUTUS SEEDLINGS 295 


during the first year. Consequently, in spite of much or most mortality 
resulting from ‘non-selective’ factors, a// survivors must be fit, even 
though the genetically “fittest” may have been eliminated by chance 
events. The rigors of the environment are such that although the “fittest” 
may not survive, the unfit never do. 


SUMMARY 


Emerging seedlings were marked and their fates followed in comparison 
with microenvironmental records from February to August in an Arbutus- 
Quercus-Sequoia forest in the Santa Cruz Mountains of central Califor- 
nia. By August 2, 2% of the Arbutus seedlings had survived in a semi- 
open forest and 0% in deep shade. Invertebrates, primarily slugs, ac- 
counted for 29.2% of the mortality, and were of greatest importance in 
the shade and during the rainy season. Death from drought acting alone 
was minor (10.4%), and occurred mainly soon after germination during 
the wet season in deep shade where thick litter prevented seedling roots 
from reaching moist mineral soil. Attacks by fungi, especially post- 
emergence damping-off and root decay types, accounted for 28.1% of 
the mortality, and were more important for several reasons in the semi- 
open than in shade. Combinations of factors killed 22.7% of the seedlings, 
mostly from mild drought preceded by crippling root decay fungi, or 
weakening by shade. Unknown or miscellaneous causes, especially under- 
mining by rodents or covering by leaves or bark, took the remainder 
(9.6% ) of the Arbutus seedlings. A number of seedlings of several other 
species, especially Sequoia sempervirens and Heteromeles arbutifolia, were 
also included in the study and showed a similar high mortality resulting 
from complex causes. Although chance or non-selective factors were prob- 
ably responsible for most mortality, natural selection still operates 
severely on the few survivors. 


Butler University, 
Indianapolis, Indiana 


LITERATURE CITED 


BAKER, F.S. 1949. A revised tolerance table. Jour. Forestry 47:179-181. 

. 1950. Principles of silviculture. McGraw-Hill Book Co. New York. 

Bray, J. R. 1956. Gap phase replacement in a maple-basswood forest. Ecology 
37:598-600. 

BUREAU OF Sorts. 1917. Reconnaissance soil survey of the San Francisco Bay Region, 
California. U.S. Government Printing Office, Wash., D.C. 

Cooper, W. S. 1917. Redwecods, rainfall, and fog. Plant World 20:179-189. 

. 1922. The broad-sclerophyll vegetation of California. Carn. Inst. Wash. 
Publ. 319. Wash., D.C. 

DAUBENMIRE, R. F. 1959. Plants and environment. Wiley & Sons. New York. For- 
est Service Station Staff. 1950. Timber stand-vegetation cover map. Calif. Forest 
and Range Exper. Sta. Berkeley, Calif. 

Fritz, E. 1950. Spotwise direct seeding of redwocd. Jour. Forestry 48:334-338. 

Harper, J. L., J. N. CLlatwortuy, I. H. McNAuGcuHToNn, and G. R. Sacan. 1961. The 
evolution and ecology of closely related species living in the same area. Evolution 
15:209-227. 


256 MADRONO [Vol. 16 


JENSEN, H. A. 1939. Vegetation types and forest conditions of the Santa Cruz 
Mountains unit of California. Calif. Forest and Range Exper. Sta. Survey Re- 
lease No. 1. Berkeley. 

Jepson, W. L. 1910. The silva of California. Memoirs of Univ. of Calif., Berkeley. 

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

Moevr, J. C. 1948. An ecological and taxonomic survey of the spermatophytes of 
Jasper Ridge. Master’s degree thesis (unpublished). Stanford Univ. 

Muwnz, P.A., and D. D. Keck. 1959. A California flora. Univ. of Calif. Press. 
Berkeley. 

OBERLANDER, G. 1953. The taxonomy and ecology of the flora of the San Francisco 
Watershed Reserve. Ph.D. thesis. Stanford Univ. 

PELTON, J. F. 1953. Ecological life-cycle of seed plants. Ecology 34:619-628. 

Rivett, M. F. 1924. The root-tubercles in Arbutus unedo. Ann. Bot. 38:661-677. 

SPRINGER, MarTua E. 1935. A floristic and ecologic study of Jasper Ridge. M.S. thesis. 
Stanford Univ. 

STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press. New 
York. 

SupwortTH, G. B. 1908. Forest trees of the Pacific slope. U.S. Forest Service. (un- 
numbered). 

TARRANT, R. F. 1958. Silvical characteristics of Pacific madrone. Pacific N. W. For- 
est and Range Exper. Sta. Silvical Ser. No. 6. Portland, Oregon. 

Tuomas, J. H. 1961a. Flora of the Santa Cruz Mountains of California. Stanford 
Univ. Press. 

. 1961b. The history of botanical collecting in the Santa Cruz Mountains 
of central California. Contr. Dudley Herb. 5:147-168. 

UnItTEeD STATES DEPARTMENT OF COMMERCE. 1959. Climatological data. California. 
Annual Summary for 1958. Vol. 62:418-443. Govern. Printing Office. Wash., D.C. 

UnitTED STATES ForREST SERVICE. 1948. Woody plant seed manual. U.S. Dept. Agric. 
Misc. Publ. 654. Govern. Printing Office. Wash., D.C. 

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and Cali- 
fornia. Ecolog. Monog. 30:279-338. 

Wipe, S.A. 1958. Forest soils. Ronald Press. New York. 


A NEW SPECIES OF DOWNINGIA 
Joun H. WEILER 


A systematic study of the genus Downingia now in progress has re- 
vealed a new entity heretofore included by all workers in D. elegans 
(Dougl. ex Lindl.) Torr. Investigation of plants in the field and in cultiva- 
tion suggests that this new entity should be accorded specific recognition. 
The new species I have chosen to name for Dr. Rimo Bacigalupi, Curator 
of the Jepson Herbarium, who has stimulated my interest in Downingia 
and given me a great deal of time and valuable advice. 


Downingia Bacigalupii sp. nov. Planta annua caulibus simplicibus vel 
e basibus ramosis, 0.5—3.0 dm. altis; foliis bracteisque linearibus vel 
lanceolatis; lobis calycis adscendentibus vel rotatis, quam sinibus laterali- 
bus tubi corollae longioribus; corollis caesiis lineis conspicuis azureis 
reticulato-venosis, labio inferiore concavo, labium superiore longitudine 
aequante vel quam id paulo longiore, centraliter albo haud venoso maculis 


1962 | WEILER: DOWNINGIA 257 


A 


Fic. 1. Perianths of D. elegans (A) and D. Baciga/upii (B) showing differences 
in color patterns and conformation of the corollas, <3. 


duabus flavis plerumque ovalibus in medio areae albae obsito; corollae 
tubo brevissimo latissimoque; lobis labii superioris cuneato-lanceolatis, 
divergentibus, erectis vel recurvatis; tubo staminali quam corollae tubo 
valde longiore, prorse curvato, antheris inferioribus duabus apice seti- 
geris; ovario uniloculari, placentis parietalibus duabus; seminis ellipticis 
angustis striis plus minusve parallelibus obsitis. 

There are similarities between D. elegans and D. Bacigalupu which 
make it difficult to differentiate the two on herbarium sheets. Both have 
widely opened corolla-tubes with the sides of the tube forming approxi- 
mately a 90-degree angle, and both have a long exserted staminal column 
with a sharply curved anther-tube. It is the combination of these two 
characters easily seen on mounted specimens which causes people to con- 
fuse the two. 

When living plants are compared, the most striking differences between 
plants of the two species are in the colors of the corolla. Corollas of D. 
elegans are a smooth, bright blue, with the lower lip having a central 
bilobed white spot. This white area is sometimes veined with blue reticu- 
lations or may even be completely suffused with blue. In contrast, the 
corolla of the new species is a lavender-blue, usually with prominent, 
more deeply colored veins, especially noticeable on the lower corolla-lobes. 
The lower lip of the corolla has a central white area which is devoid of 
blue veins and contains two bright orange-yellow spots. Other differences 
neither as consistent nor as easy to recognize, are size and shape of corol- 
la-lobes. Lobes of the upper corolla-lip of D. elegans are narrow, usually 


258 MADRONO [Vol. 16 


parallel or crossed over each other but occasionally divergent, and sharp- 
ly reflexed. The lobes of the lower corolia lip are cuneate and tapered toa 
point. In D. Bacigalupu, the upper lobes are broader, widely divergent, 
and erect or arched backward, but not so prominently as in D. elegans. 
The lobes of the lower lip are broader, rounded, and abruptly pointed. 
Besides morphological differences, the gametic chromosome number of 
D. Bacigalupi, including plants of the type collection, is n=12, whereas 
that of D. elegans is n=10. 

Downingia Bacigalupu occurs from southwestern Idaho westward 
across southern Oregon, as far north as southern Wasco County east of 
the Cascade Mountains, and in northeastern California as far south as 
Lake Tahoe. This range overlaps that of D. elegans only in southern Was- 
co County, Oregon. D. Bacigalupu grows in vernal pools, roadside ditches, 
open areas of mountain meadows and in muddy margins of lakes at sites 
exposed to bright sunlight. 

Type. In heavy soil of a large howl-shaped depression littered with 
rocks, 2.7 miles southwest of the California-Oregon border along Ager- 
Beswick road, Siskiyou County, California, June 24, 1960, J. H. Weiler 
and A. P. Nelson 60205 (UC-1,199,666). 

Other collections. CALIFORNIA. Sierra County: 1 mile south of junction to 
Calpine, Bacigalupi 4276. Plumas County: 11.7 miles north of Sattley, Sierra Valley, 
Weiler 59190. Lassen County: 11.1 miles south of Eagle Lake, Weiler 59203. Shasta 
County: Dickson Flat 3.2 miles south of Shasta-Siskiyou County line, Weiler 60194. 
Modoc County: Pitt River Valley south of Alturas, Mason & Grant 13414. OREGON. 
Josephine County: 3.7 miles north of O’Brien, Weiler 61319. Jackson County: 1.5 
miles east of Klamath Falls Junction, Weiler 60177. Klamath County: Modoc Point, 
Klamath Lake, Constance 9682. Wasco County: 0.9 miles north of Schoolie Ranger 
Station road on the road to Mount Wilson, Weiler 61397. Harney County: 1.2 miles 
west of Riley, Weiler 61345. IDAHO. Owyhee County: 10 miles south of Riddle, 
Holmgren 7976. 


Department of Botany, 
University of California, Berkeley. 


THREE NEW SPECIES RELATED TO MALACOTHRIX 
CLEVELANDII? 


WILLIAM S. DAVIS AND PETER H. RAVEN 


Malacothrix clevelandii A. Gray, a cichoriaceous composite of the 
southwestern United States and northwestern Mexico, has been regarded 
as a homogeneous species by all previous authors, including the most 
recent monographer of the genus, E. W. Williams (Am. Midl. Nat. 58:494- 
512. 1957). Stebbins et al. (Univ. Calif. Publ. Bot. 26:401—430. 1953) 
reported the gametic chromosome number n=7 for a population of this 
species from coastal California, whereas the plants they examined from 


1 The authors are indebted to Professors Carl C. Epling, Harlan Lewis, and Henry 
J. Thompson for their constructive criticism of the manuscript. 


1962 | DAVIS & RAVEN: MALACOTHRIX 259 


Arizona were tetraploid (n=14) and differed morphologically from most 
California material. Subsequent chromosomal and morphological studies 
of M. clevelandu have shown that, far from being a non-variable species 
as its lack of synonyms or infraspecific entities might suggest, it is a com- 
plex of several closely related entities, for which we propose the taxonomic 
treatment below. 

Malacothrix clevelandiu and the related taxa that will be described here 
are recognized collectively by their erect, inconspicuous heads with the 
ligules barely exserted; by the presence of one or more persistent pappus 
setae; and especially by a ring of acute teeth at the summit of the achene. 
The last-mentioned has proved to be the most useful trait separating them 
from members of the closely related group of species of which M. foliosa 
A. Gray is a member (fig. 2). 

The taxa of the M. clevelandu group can be distinguished by means of 
the following key: 


Achenes less than 1.7 mm. in length, fusiform, with 5 of the 15 ribs more prominent 
than the rest; involucre less than 8 mm. high; persistent pappus seta 1 


Achenes brown or straw-colored; cauline leaves often toothed; plants usually 
unbranched below; mean pollen diameter 254; gametic chromosome number, 
n=/, ae: ee ee oe 2 i clevelandy 


Achenes dark purplish-brown, rarely paler; margins of cauline leaves entire; 
plants often well-branched from the base; mean pollen diameter 304; gametic 
chromosome number, n=14 aerate fw A ee ee a Les 


Achenes more than 1.7 mm. in length, subcylindrical, grey-brown to straw-colored; 
with 15 equally prominent ribs; involucre more than 8 mm. high; persistent 
pappus setae 1 or 2 


Persistent pappus setae usually 2, the upper portion of the achene smooth; mar- 
gins of the basal leaves irregularly and doubly dentate; mean pollen diameter 
25 : : 2. 2 : : ; . : : . «= «= UW. sonorae 


Persistent pappus seta usually 1, the upper portion of the achene with ribs; mar- 
gins of the basal leaves dentate; mean pollen diameter 30u . WM. stebbinsii 


Pollen diameter was found to be useful for distinguishing taxa of the 
M. clevelandui complex, and, in addition, provided a clue to the level of 
polyploidy in two of them. Po!len samples were taken from herbarium 
sheets of WM. clevelandiu, M. similis, M. sonorae, and M. stebbinsu, and 
the size of mature grains was measured with an ocular micrometer. The 
mean standard deviation and range of the sample from each species are 
shown in Table 1. The mean of the pollen sample from M. similis (n=14) 
was compared with the mean of the pollen sample from M. clevelandu 
(n=7) by use of Student’s ¢-test, and the results show a highly significant 
difference between the mean pollen sizes. On the other hand, the pollen 
of M. clevelandiit and M. sonorae was closely similar in size, as was that 
of M. similis and M. stebbinsu (Table 1). Since in addition the pollen 
from M. similis and M. stebbinsii was mostly tetra-aperturate while that 
from M. sonorae and M. clevelandii was predominantly tri-aperturate, we 
believe that M. sonorae is probably diploid, and that M. stebbinsii is 
probably tetraploid. 


260 MADRONO [Vol. 16 


TABLE 1. DIAMETERS OF POLLEN GRAINS OF SPECIES OF MALACOTHRIX. 


NUMBER OF SAMPLE STANDARD 
SPECIES PLANTS sI1zE1l RANGE(uw) MEAN(u) DEVIATION (1) 
M. clevelandii 29 394 21-31 25 1.8 
M. similis ike} 347 23-38 30 23 
M. sonorae 10 260 20-29 25 1.3 


M. stebbinsi 49 794 24-36 30 De 


1 Pollen from one plant from each locality, 20-50 grains per plant. A plant from 
the type collection was included in each case. Data obtained from herbarium speci- 
mens. 


DESCRIPTIONS OF THE SPECIES 


MALACOTHRIX CLEVELANDII A. Gray, Bot. Calif. 1:433, 1876 (fig. 1, 2a). 

Annual herb 5—60 cm. tall, with a single stem or, more rarely, numer- 
ous stems from the base; basal leaves linear to narrowly lanceolate, den- 
tate, pinnatifid or lobed, the rachis oblong or wider near the base; cauline 
leaves often toothed; heads cylindrical to narrowly campanulate, 10—160 
(median, 36), 4-8 mm. high, 2-5 mm. broad, 19—67—flowered; ligules 
yellow; pollen grains 21—3lu (mean=—=25,) in diameter; achenes trun- 
cate-fusiform, 1.4-1.8 mm. long, 0.22—0.38 mm. wide, slightly curved, 
brown to straw-colored, finely 15—ribbed, with 5 ribs more prominent than 
the rest, the achene pentagonal in transverse section, its apex flared, bor- 
dered by a ring of 14-17 white-scarious teeth, of which the basal portions 
extend above the achene lip, the teeth often irregularly cleft, outwardly 
curved, lance-deltoid, the persistent seta 1. Gametic chromosome num- 
ber, N==7" 

Type. San Diego, California, Cleveland (GH; isotype, K). 

Representative specimens.2 CALIFORNIA. Tehama County: 5 miles west of Pasken- 
ta, Baker 12581. Glenn County: 9 miles east of Alder Springs, Heller 11452. Colusa 
County: upper Sand Creek, Hoover 3212. Lake County: Scotts Valley, Tracy 1646. 
Contra Costa County: Mitchell Canyon, Mount Diablo, Bowerman 1415. Santa 
Clara County: Seeboy Ridge, Mount Hamilton Range, Sharsmith 3270. San Benito 
County: Pinnacles, Howell 12933. Monterey County: King City, K. Brandegee in 
1893. San Luis Obispo County: 8 miles east of Santa Margarita, Ferris & Rossbach 
9440. Santa Barbara County: Painted Cave Ranch, Eastwood 120. Kern County: 
Kern River, Pezrson 8835. Ventura County: Kinchers, Ojai Valley, Pettzbone & Hubby 
in 1896. Los Angeles County: east fork of Santa Anita Canyon, Howell 3778. San 
Bernardino County: Cajon Pass, S. B. Parish 4868. Riverside County: Santa Rosa 
Mountains, Munz 15087. San Diego County: Buckman Springs, Fosberg 8486. Tuo- 
lumne County: above Indian Creek, Williamson 80. Mariposa County, Sherlocks, 
Congdon in 1897. Calaveras County: Mokelumne Hill, Blazsdell. Amador County: 


2 The following herbaria have been consulted, and for this privilege the writers 
are grateful to the curators of the institutions concerned: University of Arizona, 
British Museum (Natural History), California Academy of Sciences, University of 
California (Berkeley), University of California (Los Angeles), Pomona College, 
Rancho Santa Ana Botanic Garden (where vouchers for our chromosome counts are 
deposited), Royal Botanic Gardens, Kew, and Stanford University. 


1962 | DAVIS & RAVEN: MALACOTHRIX 261 


M. clevelandii o 
M. similis r 
M. sonorae i 
M. stebbinsii x 
ow 
x 
a» 


oy e 


% ° 


@ 


Fic. 1. Distribution of Malacothrix clevelandi and allied species in the south- 
western United States and northwestern Mexico. 


Drytown, Hansen 401. BAJA CA irorniA, Mexico. Guadalupe Island, Palmer 51; 
13 miles southeast of Tecate, Munz 9520. 

As shown by the specimens cited above and by figure 1, this species 
occurs on the coastward slopes of the mountains of California and north- 
ernmost Baja California. Its occurrence on Guadalupe Island should be 
confirmed by additional material and by determination of chromosome 
number. In addition to the report of Stebbins e¢ al. of a chromosome 
number of 2n=14 from the Sharsmith collection cited above from Santa 
Clara County, we have obtained this number in a collection from the 
Santa Monica Mountains, Los Angeles County, California (Raven & 


262 MADRONO [Vol. 16 


Thompson 15034) and the gametic number of n=7 in a collection from 
the San Jacinto Mountains, Riverside County, California (Davis 99). 


Malacothrix similis sp. nov. (fig. 1, 2c). Herba annua; foliis ad radices 
linearo-lanceolatis, integris, dentatis, lobatis, vel pinnatifidis; capitulis 
anguste campanulatis, 6-10 mm. longis, 3-6 mm. latis, floribus 32-73; 
corollis flavis; achaeniis truncato-fusiformis, 1.4-1.7 mm. longis, sub- 
flexuosis, maximam partem purpureo-brunneis interdum stramineis, sub- 
tiliter 15-costatis, 5 costis prominulis, 5-angulatis in sectione transversa, 
parte superiore late dilata, ab annulo setarum circa 18 scabriosarum cir- 
cumdata; seta perstata 1. 

Annual herb 5—32 cm. tall, usually branched from the base, the stems 
1-11; basal leaves linear-lanceolate, entire, dentate, lobed, or pinnatifid, 
the rachis oblong; cauline leaves subentire; heads narrowly campanu- 
late, 5—50 (median, 14), 6-10 mm. high, 3-6 mm. broad, 32—73 flowered; 
ligules yellow; pollen grains 23-38 in diameter (mean=30,) ; achenes 
truncate-fusiform, 1.4—1.7 mm. long, 0.26—0.31 mm. wide, slightly curved, 
dark purplish-brown, sometimes straw-colored, finely 15-ribbed, with 5 
ribs more prominent, the achenes pentagonal in transverse section, the 
apex flared, bordered by a ring of about 18 white-scarious irregular teeth, 
of which the basal portions extend above the achene lip, the teeth curved 
outward, lance-deltoid, the persistent seta 1. Gametic chromosome num- 
ber, n==14. 

Type. With /dria, Yucca, Salvia, and Solanum hindsianum, 16.0 kilo- 
meters southeast of El] Rosario, Baja California, Mexico, altitude 210 
meters, 18 April 1958, Raven, Mathias & Turner 12475 (RSA). 

Specimens examined. CALIFORNIA. Santa Barbara County: Santa Cruz Island, 
T. S. Brandegee in 1888. Ventura County: Hueneme, Pezrson 5772; Hueneme Beach, 
Munz 9390. Baja CALIFORNIA, Mexico. South Todos Santos Island, Moran 2804; 
San Quintin, Bacigalupi 3020, Epling & Stewart in 1936, Raven et al. 12355; Rosario 
wash, Wiggins 5427; 3.5 miles east of Rancho San José, Wiggins 9783; Arroyo el 
Agua Marga, Wiggins 9935, 9935B; El Rancho Viejo, 7. S. Brandegee in 1889; Cedros 
Island, Anthony 434, Palmer 762. 


The chromosome number of M. similis has been determined as n=—14 
at meiosis in microsporocytes of the type collection. This species is vari- 
able and puzzling, separable only with difficulty from MW. clevelandu, but 
measurements of the pollen from the collections cited consistently have 
fallen within the tetraploid-size range. Furthermore, with the exception 
of two outlying stations in California, this species occupies a coherent 
area in Baja California south of the range of the diploid M. clevelandit. 
Much of the difficulty with respect to the delimiting of M. similis is due 
to the three collections from California. The dune habitat at Hueneme 
Beach (Ventura County), however, would be highly anomalous for M. 
cleveland, and the collections cited are morphologically distinguishable 
from that species. Plants of M. stmilis from similar beach and coastal 
plain habitats in Baja California have smaller and darker achenes than 
the Hueneme collections. Additional collections and chromosome number 
determinations from Hueneme Beach and from Santa Cruz Island, the 


1962 | DAVIS & RAVEN: MALACOTHRIX 263 


| mm 


iy b ( 
d ( f 


Fic. 2. Mature achenes of species of Malacothrix: a. M. clevelandi,; b. M. foliosa; 
c. M. similis; d. M. sonorae; e. M. stebbinsi; f. M. fendleri. 


other California station, are much to be desired. In both cases the pollen 
measurements are consistent with the range of size expected for the tetra- 
ploid. We suggest that an understanding of relationships in the complex 


264 MADRONO [ Vol. 16 


depends on a more thorough knowledge of M. foliosa and related species 
which inhabit the islands off the coast of California and Baja California, 
and which may have participated in the alloploid origin of the popula- 
tions we have named M. similis. Measurements of the pollen of different 
collections of the M. foliosa complex suggest that it contains both diploids 
and tetraploids. 


Malacothrix sonorae sp. nov. (fig. 1, 2d). Herba annua; foliis ad 
radices lanceolatis vel oblanceolatis, inaequaliter pinnatifidis; capitulis 
campanulatis, 6-9 mm. longis, 4—-6.6 mm. latis, floribus 30-61; corollis 
flavis; achaeniis columnaris 1.7—2.00 mm. longis ad basim attenuatis, sub- 
flexuosis, praesertim fuscis nunc stramineis, subtiliter aequaliterque 15— 
costatis, in sectione transversa rotundis, parte superiore dilata, achaenii 
parte superiore ad 0.2—0.3 mm. nullomodo costata, ab annula setarum 
16-18 scabriosarum circumdata; setis perstatis 2, per occasionem 1, raro 
3 vel 4. 

Annual herb 10-35 cm. tall, usually unbranched at the base but occa- 
sionally with up to 9 stems; basal leaves lanceolate to oblanceolate, ir- 
regularly and doubly dentate, the rachis broadest near the apex, nar- 
rowed below; heads campanulate, 5-109 (median, 10), 6-9 mm. high, 
46.6 mm. broad, 30—61-flowered; ligules yellow; pollen grains 20-29 pu 
in diameter (mean=—25 p); achenes cylindrical (1.6—) 1.7—2.0 mm. long, 
attenuate toward the base, slightly curved, grey-brown to straw-colored, 
finely 15-ribbed, all the ribs equal, the achene round in transverse sec- 
tion, the apex slightly expanded, the upper 0.2—0.3 mm. of the achene 
not ribbed, bordered by a ring of 16-18 white-scarious teeth, of which 
the basal portions do not extend above the achene lip, the teeth pectinate, 
straight, acicular, the persistent setae 2, occasionally 1, rarely 3 or 4. 


Type. Tucson Mountains, altitude 2600 feet, Pima County, Arizona, 
24 April 1903, Thornber 362 (ARIZ 59,491; istoypes, DS, POM, UC). 

Specimens examined. Arizona. Pima County: north base of Silver Bell Moun- 
tains, Benson 10716; Rosemont, Thornber in 1907; Sabino Canyon, Santa Catalina 
Mountains, Thornber in 1903; Tucson Mountains, Thornber 428, in 1903. Pinal 
County: between Oracle and Mammoth, Gentry 6081. Sonora, Mexico. Distrito de 
Altar: Passo San Emeterio, Keck 4135A; 4 miles west of Caborca, Keck 4040. 

The size and number of apertures of its pollen suggest that this dis- 
tinctive and rather local species may be diploid (n=7), but we have not 
yet been able to obtain living material from which to make chromosome 
counts. In achene shape (fig. 1d, a, f) it is intermediate between WM. cleve- 
landii and M. fendleri A. Gray (fig. 2), the latter a diploid® species with 
long-exserted ligules that occurs east of the range of the Malacothrix 
clevelandu complex. The range of M. sonorae likewise lies between that 
of the other two diploids. 


3 We have made two new gametic chromosome counts of M. fendleri, n=7, from 
the following collections: 1.9 miles north of Chambers, Apache County, Arizona, 
Raven 13026; 5 miles northeast of Bates Well, Pima County, Arizona, Raven 11699. 
Stebbins et al. (op. cit.) reported the same number for a collection from New Mexico. 


1962 | DAVIS & RAVEN: MALACOTHRIX 265 


We have derived the specific epithet, “‘sonorae,”’ from the Sonoran 
Desert in which the range of this taxon lies. 


Malacothrix stebbinsii sp. nov. (fig. 1, 2e). Herba annua; foliis ad 
radices lanceolatis vel oblanceolatis, dentatis, raro pinnatifidis; capitulis 
campanulatis, 7-10 mm. longis, 3.5—8 mm. latis, floribus 19-70; corollis 
flavis, raro albis; achaeniis fusiformo-columnaris, 1.7—2.3 mm. longis, ad 
basim subattenuatis, raro flexuosis, nunc cinearo-fuscis nunc stramineis, 
subtiliter aequaliterque 15-costatis, in sectione transversa rotundis, parte 
superiore subdilata, achaenii parte superiore ad 0.14—0.20 mm. non cos- 
tata, ab annulo setarum 14-17 scabriosarum circumdata; setis perstatis 1, 
per occasionem 2. 

Annual herb 6—60 cm. tall, usually unbranched at the base but occa- 
sionally with up to 9 stems; basal leaves lanceolate to oblanceolate, den- 
tate, more rarely pinnatifid, the rachis often narrowed near the base; 
heads campanulate, 5—66 (median, 20), 7-10 mm. high, 3.5—8 mm. broad, 
19--70-flowered; ligules yellow, rarely white; pollen grains 24-36 p in 
diameter (mean=30 »); achenes narrowly fusiform-columnar, tapering 
slightly to the base, 1.7—2.3 mm. long, 0.3—0.45 mm. wide, rarely curved, 
grey-brown to straw-colored, finely 15-ribbed, all the ribs equal, the 
achene round in transverse section, the apex slightly flared, the upper 
0.14-0.20 mm. of the achene not ribbed, bordered by a ring of 14-17 
white-scarious teeth, of which the basal portions rarely extend above the 
achene lip, the teeth rarely and irregularly cleft, straight, lance-linear; 
the persistent setae 1, rarely 2. 


Type. Abundant in shade of a large rock, moist soil, Mendoza Canyon, 
Coyote Mountains, Pima County, Arizona, altitude 3,800 feet, 22 April 
1945, K. F. Parker 5815 (ARIZ 32,709; isotype, UC). 


Representative specimens. NEvapA. Washoe County: hills west of Reno, Hillman 
in 1893. Clark County: Nelson, Jones in 1907. CALIFORNIA. Inyo County: Titus Can- 
yon, Eastwood & Howell 7786; 4 miles east of Aberdeen, Kerr 630; 2 miles east of 
Bradbury Wells, Howell in 1928; Slate Range, Alexander & Kellogg 1135. San Ber- 
nardino County: Turtle Mountains, Munz & Harwood 3505; Quail Springs, Little 
San Bernardino Mountains, Munz & Johnson 5227; south base of Old Dad-Granite 
Mountain Range, Wolf 10092; Kingston Mountains, Wolf 10456. Riverside County: 
Murray Canyon, Peirson 2715; 12 miles southwest of Twentynine Palms, Alexander 
& Kellogg 2129. San Diego County, Palm Canyon, Borrego Valley, Wolf 8451; San 
Felipe Hill, Jones in 1906. Arizona. Mohave County: Yucca, Jones in 1884; Cheme- 
huevis, Jones in 1903; Diamond Creek Canyon, Wilson in 1893. Yavapai County: 
Burro Creek, Crooks & Darrow in 1938; Skull Valley, Jones in 1903. Gila County: 
Pine Creek, near Roosevelt, Peebles et al. 5227; Mazatzal Mountains, Eastwood in 
1929, 17163. Pinal County: near Oracle, Peebles 6844; between Superior and Miami, 
A. & R.A. Nelson 1900; Galuro Mountains, 12 miles above Mammoth, Gentry 6051. 
Pima County: Baboquivari Peak, Goodding 4649; Florita Canyon, Knipe in 1938; 
Oracle Camp, Santa Catalina Mountains, Simon 224; Sabino Canyon, Santa Catalina 
Mountains, Thornber in 1905, in 1913. Santa Cruz County: Stone Cabin Canyon, 
Santa Rita Mountains, Thornber 5543. Sonora, Mexico. 4 miles south of Imuris, 
Abrams 13202. 


Pollen of this species is consistently larger than in M. clevelandiu and 
M. sonorae, both of which are diploids, and, like that of the tetraploid 


266 MADRONO [Vol. 16 


M. similis, is mostly tetra-aperturate. We believe that the count reported 
by Stebbins e¢ al (op. cit.) of 2n=28 for “Malacothrix clevelandi” from 
Tucson, Arizona (for which we can find no voucher) probably refers to 
M. stebbinsii. From a consideration of morphology we believe that M. 
stebbinsii may be an allotetraploid between WM. clevelandiu and M. sono- 
rae. Stebbins and his associates postulated that it might be an allotetra- 
ploid between M. clevelandiu and M. fendleri, but they were not aware 
of the probably diploid M. sonorae. As we have mentioned above, M. 
sonorae is nearly intermediate between M. clevelandiu and M. fendleri, 
both morphologically and geographically. 

Department of Botany 

University of California, Los Angeles 


Rancho Santa Ana Botanic Garden 
Claremont, California 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 
(See Madrono 9:257-258. 1948) 


SPECIES NUMBER COUNTED BY COLLECTION LOcaLiTy 
PORTULACACEAE 
Montia ne—6 P. Raven H.& M. Lewis Mather, Tuolumne 
* perfoliata LA’ in 1956, LA County, California 
(Willd.) Howell 
Pes H. Lewis H. Lewis in 1955 Mather, Tuolumne 
LA LA County, California 
n— 18 P. Raven H. Lewis in 1956 La Panza Range, San 
LA LA Luis Obispo County 
n= 1s P. Raven H. Lewis in 1956 San Juan Canyon, 
LA LA San Luis Obispo 
County, California 
n=18 P. Raven H. Lewis in 1956 Temblor Grade, 
LA LA Kern County, Calif. 
sibirica (L.) W.H. Lewis W.H. Lewis 5367 Near Sechelt, 
Howell n= 12 ASTC SMU British Columbia, 
Canada 
RANUNCULACEAE 
Delphinium n=s R. C. Jackson McGregor 14282 Douglas County, 
virescens Nutt. KANU KANU Kansas 
Trauivetteria 
grandis Nutt. ns R. Ornduff Ornduff 6262 Multorpor Moun- 
DUKE UC tain, Clackamas 
County, Oregon 
MAGNOLIACEAE 
Michelia n= 19 P. Raven Raven 14026 Cultivated, Los 
*fuscata Blume LA UC Angeles, Calif. 
SAXIFRAGACEAE 
Bolandra R. Ornduff Ornduff 6240 Elowah Falls, 
oregana S. Wats. p= DUKE UC McCord Creek, 


Multnomah County, 
Oregon 


1962 | 


SPECIES 


THY MELAEACEAE 


Daphne genkwa 
Sieb. & Zucc. 


Daphne giraldi 
Nitsche 


LENNOACEAE 


Ammobroma 
sonorae Torr. 
ex. Gray 


PRIM ULACEAE 


Hottonia 
inflata Ell. 


POLEMONIACEAE 


Phlox oklahomensis 
Wherry 


VERBENACEAE 
Verbena 


bipinnatifolia Nutt. 


perennis Wooton 


plicata Greene 


wrightii Gray 


SCROPHULARIACEAE 


Mimulus 
brevipes Benth. 


PLANTAGINACEAE 


Plantago 
insularis Eastw. 


heterophylla Nutt. 


CHROMOSOME NUMBERS 267 
NUMBER COUNTED BY COLLECTION LOcALITY 
n=) L. Nevling, Nevling 105 Cultivated, 
Jr., A AAH Arnold Arboretum, 
Weston, Mass. 
ne 9 L. Nevling, Nevling 106 Cultivated, 
rch AAH Arnold Arboretum, 


2n= 18, D.M. Moore 


n= tt 
2n — 14 
n 15 
n= 15 
n=15 
nee 15 
n— 15 
ae | 
nf 
n= 10 
n= 8 
2zn= 4, 
DT) 


LA 


O. T. Solbrig 
GH 


R. C. Jackson 
KANU 


O. T. Solbrig 
GH 


O. T. Solbrig 
GH 


O. T. Solbrig 
GH 


O. T. Solbrig 
GH 

O. T. Solbrig 
GH 

O. T. Solbrig 
GH 

O. T. Solbrig 
GH 


O. T. Solbrig 
GH 


B. B. Mukherjee. C. Hubbs in 1949 
R. K. Vickery, 


Jr., UT 


D. M. Moore 
LA 


D.M. Moore 
LA 


J. Feldner in 


1961, UCLA 


C.Wood 9426 


GH, UC 


Marsh 
KANU 


Solbrig 3168 
GH, UC 


Solbrig 3175 
GH, UC 


Solbrig 3181 
GH, UC 


Solbrig 3206 
GH, UC 


Solbrig 3213 
GH, UC 


Solbrig 3186 
GH, UC 


Solbrig 3179 
GH, UC 


Solbrig 3187 
GH Ue 


54804 UT 


Raven, Davis, 
Moore 14763 
RSA, JEPS 


Raven, Blakley, 
Ornduff 14919 


RSA 


Weston, Mass. 


5.5 mi. W. of Glamis, 
Imperial County, 
California 


Middlesex Fells 
Reservation, 
Middlesex County, 
Massachusetts 


Cowley County, 
Kansas 


10 mi. N.E. of 
Santa Fe, 
New Mexico 


32 mi. S. of 
Carizozo, 
New Mexico , 


28 mi. N. of 
Pecos, Texas 


48 mi. S. of 
Alpine, Texas 


11 mi. W. of 
Van Horn, Texas 


17 mi. S. of Fort 
Stockton, Texas 


28 mi. N. of 
Pecos, Texas 


20 mi. S. of Fort 
Stockton, Texas 


Mount Palomar, 
San Diego County, 
California 


San Felipe, 
Baja California, 
Mexico 


Santa Rosa Island, 
Santa Barbara 
County, California 


268 MADRONO [ Vol. 16 
SPECIES NUMBER COUNTED BY COLLECTION LOCALITY 
COMPOSITAE 
Bidens eZ R. C. Jackson Jackson 2955 Douglas County, 
polyle pis Blake KANU KANU Kansas 
Boltonia 
latisquama Gray n= 9 R. C. Jackson Jackson 2953 Douglas County, 
var. latisquama KANU KANU Kansas 
latisquama Gray 
var. recognita n= 18 R. C. Jackson McGregor 15865 Harvey County, 
Fern. & Grisc. KANU KANU Kansas 
Cacalia He 25 R. C. Jackson McGregor 15964 Anderson County, 
atriplicifolium L. KANU KANU Kansas 
Eupatorium n= 10 R. C. Jackson McGregor 15815 Cherokee County, 
perfoliatum L. KANU KANU Kansas 
Grindelia nie— 0 R. C. Jackson McGregor 16007. Taney County, 
lanceolata Nutt. KANU KANU Missouri 
Helenium n= [5 R. C. Jackson McGregor 15811 Cherokee County, 
tenuifolium Nutt. KANU KANU Kansas 
nudiflorum Nutt. n= 14 R. C. Jackson McGregor 15814 Cherokee County, 
KANU KANU Kansas 
Heterotheca 
latifolia Buckl. no R. C. Jackson McGregor 15849 McPherson 
var. McGregoris KANU KANU County, 
Wagenkn. Kansas 
Rudbeckia n== 19 R. C. Jackson McGregor 16001 Taney County, 
missouriensis KANU KANU Missouri 
Englm. 
Senecio n= 22 R. C. Jackson McGregor 14283 Douglas County, 
obovatus Muhl. KANU KANU Kansas 


* Prepared slide available. 


‘Symbols for institutions are those listed by Lanjouw and Stafleu, Index Her- 


bariorum, Part I. Fourth edition, 1959, Utrecht. 


REVIEW 


The Little Hill, a chronicle of the flora on a half acre at the Green Camp, Ring- 
wood, New Jersey. By ANNE OPHELIA Topp. CUAS 8, pp. 1-20. 1961. Cooper Union 
School of Art and Architecture, New York. 

There appeared on my desk a thin publication of twenty pages and four colored 
plates, much like a brochure spelling out some urgent need. The format caught my 
eye and the text, in keeping, filled me with sheer delight as it recounted the botany 


of an unkempt patch of weed. 


The Hill has been cut over, burnt over, and trampled over; 
it has been flooded in the torrential downpours of 
northeasters, hurricanes, and near-hurricanes ; 

and it has seemed to die in the droughts of many years. 


1962 | NOTES AND NEWS 269 


But each spring fresh foliage erupts, 

and each summer the growing goes on, from wild ginger to wild geranium 
to purple aster; then all returns to leaf mold for the winter. 

The Hill is romantic in the morning mist, harsh in the noonday sun, 

rich and lush in the shadows at twilight, 

and eerie in the dark of the moon. 

To the average person it is just a patch of rank weeds, 

thick matted and threatening. 

To the appreciative eye it is literally a garden 

of wild flowers, full of surprises and beauties of form and color, 

more wonderful than a suburbanite’s well nurtured backyard and often more 
rewarding, because it survives without cost, backbreak, or frustration. 


I read it and reread it and as IJ held it in my hand there came over me the sense 
of holding a priceless jewel. A professional botanist would have written three times 
as much and in his pompous style have said half as little. For there is recorded in 
simple poetic language the history of the area, a description of its physical setting, 
of its topography and its vegetation, its relation to the human occupants of the area 
today and in colonial and aboriginal times. Several years of faithful recording bring 
to light nearly 200 kinds of plants (exclusive of grasses and fungi), here presented in 
the form of a weekly almanac of blooming dates through spring, summer, and fall. 
The seasons begin with Stellaria media the first week in April and close with golden- 
rods and asters in September. Each week a new set of blooms appears. There is a 
list of the trees, of the edible plants and of the medicinal plants, the latter lists in an 
ethnological and colonial context. 


These are only a few of the secrets the Hill is waiting to disclose 

to any searching eye. 

A plant census, carried on faithfully through several years, 

revealed in our little half-acre more than 170 species of herbaceous plants, 
a dozen kinds of trees, many dozens of grasses, and a few fungi. 

For many people, each name will recall a floral acquaintance; 

other readers, who have not yet had the pleasure of an introduction, 

may find enjoyment in the poetry of the Latin names 

and in the often quaint charm of the colloquial ones. 


The text is in blank verse and the typography flawless. The color plates include 
Smilacina racemosa, Rubus orarius and R. allegheniensis, Daucus carota, Verbascum 
thapsus and V. phlomoides. The drawings are botanically accurate and lifelike and 
the color rendition is excellent. 

In a footnote we learn that the author, Anne Ophelia Todd (Mrs. Raymond 
Dowden), is artist, amateur botanist, teacher and author. The publication, known as 
CUAS, is produced by the third year students of the Cooper Union School of Art 
and Architecture of New York City. We raise our glass high in congratulations to all 
concerned.—H. L. Mason, Department of Botany, University of California, Berkeley. 


NOTES AND NEWS 


WYOMING Pinyon REvIsITED. The center of pinyon (Pznus edulis Engelm.) dis- 
tribution falls close to the geographical point, unique in the United States, where 
four states—Utah, Colorado, New Mexico, and Arizona—come together. Beyond 
these four states, pinyon extends eastward to touch Oklahoma, southward into Texas 
and northern Mexico, and westward to California. Older works on tree distribution, 
and maps copied from them, complete the symmetry of this geographic range by 
showing pinyon extending northward to southwestern Wyoming. However, a recent 
treatment of “The Gymnospermae of Wyoming” (C. L. Porter, 1957, Leaflet 28, 


270 MADRONO [Vol. 16 


Rocky Mountain Herbarium, University of Wyoming) upsets this unusual symmetry 
by stating that “reports of ... Pinyon Pine occurring naturally in southern Wyoming 
are believed to be erroneous, a thorough search for this species having failed to turn 
it up closer than about twenty miles south of the border in Larimer County, Colo- 
rado, and Daggett County, Utah.” 

In defense of the early references to the occurrence of pinyon northward into 
Wyoming, the following observations from a 1960 field trip are noted. 

1. Pinyon occurs along Sheep Creek, 5 miles south of the Wyoming border, near 
Manila, Utah (Sec. 1-2, T. 2 N., R. 19 E., Daggett County). 

2. Along the Glades, a rock outcrop nearly paralleling the State border near longi- 
tude 109 degrees 30 minutes West, scattered pinyons occur on both sides of the border, 
and are numerous at Minnie’s Gap (a break in the Glades) in Wyoming (Sec. 23, 
T. 12 N., R. 107 W., Sweetwater County). 

3. Northward from Minnie’s Gap pinyon occurs as a very minor element in the 
juniper stands. The most northerly pine found is four miles inside Wyoming and five 
miles east of the Green River (that is, four and one-half miles east of the future 
shoreline of Flaming Gorge Reservoir; southern boundary of Sec. 34, T. 13 N., 
R. 107 W., Sweetwater County). This pine is twenty-six inches in diameter at breast 
height and more than 200 years old, but too decayed for exact dating. Probably 
pinyons occur north of this old tree as well. 

Specimens of native Wyoming Pinus edulis from Minnie’s Gap (Peterson 206-60) 
have been sent to the Rocky Mountain Herbarium, Laramie, Wyoming, and to the 
United States Forest Service Herbarium, Washington, D.C. RocEer S. PETERSON, 
Rocky Mountain Forest and Range Experiment Station, United States Forest Service, 
Fort Collins, Colorado. 


A CONTROVERSIAL TREATMENT OF THE POLEMONIACEAE. The treatment of the Pole- 
moniaceae in Part 4 of the “Vascular Plants of the Pacific Northwest” (Hitchcock, 
Cronquist, Ownbey, and Thompson, 1959) evokes criticism on the part of an emu- 
lator of Polemon, the bellicose philosopher. From numerous cases of disagreement, 
a few of major importance may be selected for attention. 

Eriastrum wilcoxii, reduced to varietal status under the endemic E. sparsiflorum. 
The architecture, leaf-, calyx-, and corolla-characters of these are deemed too dis- 
similar to justify this. What is really needed is the separation of the comprehensive 
E. wilcoxii into its multiple subspecies or varieties. 

Gilia attenuata, submerged in subjective (‘“‘taxonomic’’) synonymy in G. aggre- 
gata. The corolla-characters of the two are so distinctive, corresponding to pollina- 
tion by different organisms, that they surely merit some nomenclatural recognition. 

Gilia inconspicua, sinuata, etc., brought together in a chaotic assemblage. The 
thorough morphologic studies by the Grants deserve more respectful consideration 
than this, supplemented as they are by cytotaxonomic work, one of the best pres- 
ently available means of throwing light on otherwise obscure inter-relationships. 

Leptodactylon pungens segregates, reduced to subjective synonymy under one 
comprehensive species. These are so distinctive in morphology, ecology, and range 
that they need recognition at some level. 

Phlox bryoides, reduced to subjective synonymy under P. muscoides. These are 
so unlike as to call for at least subspecies segregation. 


Phlox douglasii, reduced to subjective synonymy under P. caespitosa. This is a 
serious misunderstanding. Judging by their types, as recognized by the systematists 
of a century, they are wholly unrelated. Phlox douglasii has thin dark green acicular 
leaves covered by long gland-tipped hairs. Phlox caespitosa has thickish pale green 
linear-oblong leaves bearing coarse glandless cilia; its only glandularity consists in 
a few hairs on the inflorescence-herbage. No intergrades between them are known. 
They are surely about as distinct species as can exist among the Microphloxes. 

The type locality of P. caespitosa has been inferred from the label to lie at the 
mouth of the Flathead River, but McKelvey finds Wyeth on its collection-date to 


1962 | NOTES AND NEWS 21h. 


have been at Flathead Post, latitude 47° 35 ’N., longitude 115° 12'%4’ W.* The ele- 
vation of the “high hills” thereabouts approaches 6000 feet. Taxon caespitosa, then, 
is the ecad of moderate elevations, while taxon pulvinata is the ecad of high country. 
They do not differ in any major respect, and intergrade completely. If any reduction 
to subjective synonymy is considered desirable, taxon pulvinata is the one needing 
suppression. 


Phlox lanata, reduced to subjective synonymy under P. hoodii. This intergrades, 
however, not with P. hoodz but with P. bryoides, and the plants at some stations— 
e.g., Double Springs Pass, Idaho—can not be certainly assigned to one or the other. 


Phlox longifolia, interpreted as a grossly comprehensive species, with a host of 
subjective synonyms. Field study shows that most of these submerged taxa occur so 
frequently in pure stands and under specialized ecologic or geographic conditions as 
to merit some degree of nomenclatorial recognition. 


Phlox missoulensis, a striking endemic, reduced to varietal status under the wholly 
unrelated P. kelseyi. The latter is a succulent marsh plant with pale, coarsely ciliate 
leaves, its glandularity, if any, limited to the inflorescence-herbage. The endemic has 
thin, deep-green leaves, bearing copious long glandular hairs, and grows in dry rocky 
situations. If it must be reduced in status, then P. douglasii would be its closest 
earlier-named relative. However, students of plant geography, ecology, evolution, etc., 
find endemics of much interest, and these should be emphasized, not obscured by 
association with more or less (or un-) related taxa. 


Phlox mollis, another endemic, reduced to subjective synonymy under the mark- 
edly dissimilar P. viscida. As its name implies, the latter is one of the most glandular 
phloxes known; it has multiflorous inflorescences and is accordingly assignable to 
Brand’s subgenus Macrophlox. The endemic has the flowers solitary or in 3’s, and 
so fits into his subgenus Microphlox. Its indument is utterly different, consisting of 
copious woolly hairs on the stems and lower side of leaves, indeed resembling P. 
lanata. Most specimens of P. mollis in herbaria have indefinite localities, but in addi- 
tion to the type station, there is one at Lewis Peak, Washington (latitude 46° 314’ N., 
longitude 117° 59%’ W.). 

Phlox scleranthifolia, reduced to subjective synonymy under P. hoodii. In habit 
and measurements, except for the slightly narrower leaves, it agrees instead with 
P. diffusa. 

Last but not least, comes Polemonium. In passing, it should be emphasized that 
P. occidentale was named by Greene provisionally only, and under the current Code 
of Nomenclature is not valid. However, it is the treatment of P. pulcherrimum for 
which special criticism is called. Many of the taxa made subjective synonyms of this 
are only remotely related. And what can be gained by classing as merely varietally 
distinct two taxa as dissimilar as those figured on page 144 as var. pulcherrimum 
and var. calycinum? (The latter, by the way, is not the same as P. calycinum East- 
wood, a Californian endemic). If all the low-growing, simply-pinnate leaved, rotate- 
campanulate flowered members of the genus are deemed one species, then its name 
should be Polemonium reptans L. 

To make “species” so comprehensive that they include multiple discordant ele- 
ments releases collectors and herbarium curators from having to examine their speci- 
mens closely, but is not the way to advance our understanding of a family as com- 
plex as the Polemoniaceae. EpGAR T. WHERRY, Botanical Laboratory, University of 
Pennsylvania, Philadelphia. 


*A stupid reviewer of my book on Phlox held that my giving latitudes and longi- 
tudes made localities difficult to find. Actually the reverse is true: their positions can 
be ascertained by measurement in any atlas, however few place-names or political 
boundaries it may show. [The book on Phlox above referred to is “The Genus Phlox,” 
Morris Arboretum Monograph III, 1955. Obtainable for $4.00, 9414 Meadowbrook 
Ave., Philadelphia 18, Penna.—-Ed. | 


272 MADRONO [Vol. 16 


CNEORIDIUM DUMOSUM (NUTTALL) HOOKER F. COLLECTED MARCH 26, 1960, AT AN 
ELEVATION OF ABOUT 1450 METERS ON CERRO QUEMAZON, 15 Mites SouTH oF BAHIA 
DE LOS ANGELES, BAyA CatrFrorntA, MExiIco, APPARENTLY FOR A SOUTHEASTWARD 
RANGE EXTENSION OF SOME 140 MILEs. 

I got it there then (8068). 

I wish to express my sincere thanks to the San Diego Museum of Natural History 
and particularly to its director, Dr. George E. Lindsay, for making possible the trip 
on which this interesting specimen was collected; to my companion of the trip, Mr. 
Glen Ives, then staff artist of the Museum but functioning on the trip as collector of 
birds and mammals, for much help and encouragement during the field work; to 
Senor Ricardo Daggett of Bahia de los Angeles, majordomo of the Vermilion Sea 
Field Station of the San Diego Museum of Natural History, for help in planning and 
arranging the trip; and to Senor Pepe Smith and his 14-year-old son Favian, both 
of Bahia de los Angeles, who packed us into the mountains, for many courtesies ex- 
tended. I am very grateful to Miss Anita Carter, Principal Herbarium Botanist of the 
University of California, Berkeley, for graciously verifying my determination of the 
specimen. I also wish to extend my thanks to the editor of the publications of the 
San Diego Society of Natural History for his many helpful suggestions during the 
compilation and processing of the data and the writing of the manuscript; to Dr. 
Helen K. Sharsmith, Senora Herbarium Botanist of the University of California, 
Berkeley, for her valuable suggestions on expanding the discussion and making the 
title more precise; to Mrs. Jerry Heller of the Museum staff for her very careful and 
accurate typing and retyping of the manuscript; and to Mrs. Rosemarie Fiebig of 
the Museum staff for taking the final manuscript to the post office for mailing. I must 
also express my deep gratitude to all my former mentors, to whose excellent instruc- 
tion and training must ultimately be attributed any merit that this unworthy con- 
tribution may possess, although, needless to say, any errors are my own: in particular, 
I would name Professor Ira L. Wiggins and the late Professor LeRoy Abrams, of 
Stanford University; Professor Robert T. Clausen, of Cornell University; and Pro- 
fessors Lincoln Constance, Herbert L.Mason, and G. Ledyard Stebbins, of the Uni- 
versity of California, Berkeley. Last but not least, I cannot fail to mention my deep 
indebtedness to my parents, without whose early cooperation this work would never 
have been possible-—Re1p Moran, Museum of Natural History, San Diego, Cali- 
fornia. 


PUBLICATIONS OF Marcus E. JONES AVAILABLE.—Recent correspondence with Mrs. 
C. A. Broaddus, a daughter of the noted western botanist, Marcus E. Jones, reveals 
that several of her father’s botanical contributions are still available to those inter- 
ested in purchasing them. These papers are of considerable historical as well as botan- 
ical interest and have been virtually unattainable from bockdealers in the past few 
vears. The following works are in print: Ferns of the West (1882) ; Contributions to 
Western Botany, nos. 7, 8, 9, 13, 14, 15, 16, 17 and 18; Montana Botany Notes (1910) ; 
and Astralagus, Revision of the North American Species (1923). Correspondence re- 
garding these publications should be directed to Mrs. C. A. Broaddus, P.O. Box A-1, 
Carmel, California——RoBERT ORNDUFF, Duke University, Durham, North Carolina. 


1962 } 


INDEX 273 


INDEX TO VOLUME XVI 


Classified entries: Chromosome numbers, Reviews. New scientific names are in 
boldface type. Un-annotated taxa in floral lists are omitted from Index. 


Adiantum xX tracyi C. C. Hall, Cytolog- 
ical observations on, 158 

Aecidium: bouvardiae, 202; chamaecris- 
tae, 202 

Alderson, Rufus Davis, 224, fig. 225 

Allium: rotundum, 49, fig. 47; stami- 
neum, 50, fig. 51 

Ambrosia bryantii, 234, fig. 235; (Com- 
positae), The unique morphology of 
the spines of an armed ragweed, 233 

Aquatic Flora, of Arizona, Additions to 
the, 32 

Arbutus menziesii, Factors influencing 
survival and growth of a seedling pop- 
ulation in California, 237 

Arceuthobium: Abnormal fruits and 
seeds in, 96; americanum, figs. 97-99; 
vaginatum, Observations on, in Mex- 
ico, 31, f. cryptopodum 96, figs. 97, 99 

Arum orientale var. elongatum, 48, fig. 47 


Baker, Milo S., 155 

Bigelow, John Milton, 179 

Blake, S. F.: Edward Palmer’s visit to 
Guadalupe Island, Mexico, in 1875, 1 

Bowerman, M. L.: Review, Flora of the 
Santa Cruz Mountains of California, 
138 

Bubakia mexicana, 202 


Carex: capitata, 232; nelsonii, 232; i- 
sandra, 230 

Ceanothus: arboreus, 29; arcuatus, 29; 
cordulatus, 29; cuneatus, 29; divarica- 
tus, 29; Germination of, seeds, 23; im- 
pressus, 29; integerrimus, 29; lemmoni, 
29; megacarpus, seed germination, In- 
fluence of temperature and other fac- 
tors on, 132, 135; prostratus, 29; so- 
rediatus, 29; spinosus, 29 

Chisaki, F., and R. Ornduff: Plagio- 
bothrys austinae (Greene) Johnston: a 
new addition to the Oregon flora, 108 

Chromosome numbers: Adiantum 
tracyi, 161; Ammobroma sonorae, 267; 
Bidens polylepis, 268; Bolandra ore- 
gana, 266; Boltania latisquama var. 
latisquama, 268, var. recognita, 268; 
Cacalia atriplicifolium, 268; Cryp- 
tantha circumscissa subsp. circum- 
scissa, 170, micrantha subsp. lepida, 
170, subsp. micrantha, 170, similis, 170; 
Delphinium virescens, 266; Downin- 
gia Bacigalupii, 258, elegans, 258; Es- 
chscholzia covillei, 94, minutiflora, 94, 
parishii, 94; Eupatorium perfoliatum, 
268; Galium coloradoense, 116, gray- 


anum subsp. glabrescens, 116, subsp. 
grayanum, 116, hallii, 116, hardhamae, 
166, hypotrichium subsp. hypotrichi- 
um, 116, subsp. scabriusculum, 117, 
subsp. subalpinum, 117, subsp. tomen- 
tellum, 117, subsp. utahense, 117, mag- 
nifolium, 117, matthewsii, 117, multi- 
florum, 117, f. hirsutum, 118, munzii 
subsp. munzii, 118, var. kingstonense, 
118, parishii, 119, rothrockii subsp. 
rothrockii, 120 (wrightii subsp. roth- 
rockil, see Errata), serpenticum, 119, 
var. puberulum, 119, stellatum subsp. 
eremicum, 119; Grindelia lanceolata, 
268; Helenium nudiflorum, 268, tenui- 
folium, 261; Heterotheca latifolia, 268, 
var. McGregoris, 268; Hattonia inflata, 
267; Lepidium bourgeauanum, 80, fig. 
88, campestre, 80, densiflorum var. 
densiflorum, 80, var. elongatum, 80, 
var. macrocarpum, 80, var. pubicar- 
pum, 80, latifolium, 80, perfoliatum, 
80, ramosissimum, 80, virginicum, 80; 
Malacothrix clevelandii, 261, 264, fend- 
leri, 264, similis, 262, 266, stebbinsii, 
266; Michelia fuscata, 266; Mimulus 
arvensis, 149, aurantiacus, 105, brevi- 
pes, 267, cordatus, 149, floribundus, 
105, glabratus var. fremontii, 150, var. 
parviflorus, 150, var. utahensis, 150, 
glaucescens, 149, guttatus, 148, subsp. 
guttatus, 148, subsp. litoralis, 148, var. 
puberulus, 148, laciniatus, 149, laxus, 
149, luteus, 150, lyratus, 149, moscha- 
tus, 105, nasutus, 149, pilosiusculus, 
150, platycalyx, 149, ringens, 105, 
tilingii var. corallinus, 149, var. tilin- 
gii, 149; Montia perfoliata, 266, sibi- 
rica, 266; Phlox oklahomensis, 267; 
Plantago heterophylla, 267, insularis, 
267; Rudbeckia missouriensis, 268; 
Senecio obovatus, 268; Trautvetteria 
grandis, 266; Verbena bipinnatifolia, 
267, perennis, 267, plicata, 267, wrightii, 
267 

Clathraceae, in California, 33 

Colchicum biebersteinii, 52, fig. 51 

Cooke, W. B., and G. Nyland: Clathra- 
ceae in California, 33 

Copeland, H. F.: Review, Die Evolution 
der Angiospermen, 70 

Crocus: ancyrensis, 60, fig. 59; suteri- 
anus, 61, fig. 63 

Cryptantha: A new species (section Cir- 
cumscissae) from California and two 
recombinations (section Circumscissae 
and section Angustifoliae), 168; cir- 


274 MADRONO 


cumscissa subsp. rosulata, 170; mi- 
crantha subsp. lepida, 171; similis, 
168 

Culberson, W. L.: Discovery of the 
lichen Parmeliopsis placorodia in West- 
ern North America, 31 

Cummins, G. B., and J. W. Baxter: No- 
mencliature, life histories, and records 
of North American Uredinales, 201 

Cupressus sargentil, The Santa Lucia 
groves and their associated northern 
hydrophilous and endemic species, 173 


Davidson, J. F.: Sphenophyllum nyma- 
nensis sp. nov. from the Upper Penn- 
sylvanian, 106 

Davis, W. S., and P. H. Raven: Three 
new species related to Malacothrix 
clevelandii, 258 

Dempster, L. T.: A new species of 
Galium in California, 166 

Dowingia: A new species of, 256; Baci- 
galupii, 256, fig. 257; elegans, 257 

Draba glabella, 231 

Drouet, F.: A new name in the algal 
genus Phormidium, 108 


Ehrendorfer, F.: Evolution of the Gali- 
um multiflorum complex in western 
North America, 109 

Elatine californica, 32 

Eriodictyon: A subarborescent new, from 
San Luis Obispo County, California, 
184; altissimum, 184, fig. 185 

Eriophorum callitrix, 230 

Eschscholzia: covillei Greene, a_ tetra- 
ploid species from the Mojave Desert, 
91; minutiflora, 94; parishii, 95 


Foliar xeromorphy, of certain geophytic 
monocotyledons, 43 


Gagea arvensis var. semiglabra, 53, fig. 51 

Galium: A new species in California, 
166; coloradoense, 116; grayanum 
subsp. glabrescens, 116, subsp. gray- 
anum, 116; hallii, 116; hardhamae, 
166, fig. 167; hypotrichium subsp. hy- 
potrichium, 116, subsp. scabriuscu- 
lum, 117, subsp. subalpinum, 117, 
subsp. tomentellum, 117, subsp. uta- 
hense, 117; magnifolium, 117; mat- 
thewsii, 117; multiflorum, 117, f. hir- 
sutum, 118; multiflorum complex, dis- 
tribution of, fig. 114, evolution of, in 
western North America, 109; munzii 
approaching G. magnifolium, 118; 
munzii subsp. munzii, 118, var. king- 
stonense, 118; parishii, 119; rothrockii 
subsp. rothrockii, 120 (wrightii subsp. 
rothrockii, see Errata); serpenticum, 
119, var. puberulum, 119; stellatum 
subsp. eremicum, 119 


[Vol. 16 


Gladiolus atroviolaceus, 62, fig. 63 

Gossypium klotzschianum davidsonii and 
Stegnosperma cubense, not known in 
the Revillagigedos, 108 

Guadalupe Island, Mexico, Edward Pal- 
mer’s visit, in 1875, 1 

Gustafson, A. H.: Review, The genus 
Datura, 72 


Hadley, E. B.: Influence of temperature 

and other factors on Ceanothus mega- 
carpus seed germination, 132 

Haller, J. R.: Some recent observations 
on Ponderosa, Jeffrey and Washoe 
pines in northeastern California, 126 

Hardham, C. B.: The Santa Lucia Cu- 
pressus sargentii groves and their as- 
sociated hydrophilous and endemic 
species, 173 

Hawksworth, F. G.: Abnormal fruits and 
seeds in Arceuthobium, 96; Observa- 
tions on Arceuthobium vaginatum in 
Mexico, 31 

Herre, A. W. C. T., 102, fig. 103 

Hordeum bulbosum, 46, fig. 47 


Tris aphyllus, 63, fig. 65 


Jeffrey, Ponderosa, and Washoe pines in 
northeastern California, Some recent 
observations on, 126 

Jepson, W. L. California Botanical Ex- 
plorers—XII. John Milton Bigelow, 
179 

Johnson, P. L.: The occurrence of new 
artic-alpine species in the Beartooth 
Mountains, Wyoming-Montana, 229 


Juncus albescens, 232 


Kasapligil, B.: An anatomical study of 
the secondary tissues in roots and 
stems of Umbellularia californica Nutt. 
and Laurus nobilis L., 205; Foliar 
xeromorphy of certain geophytic mon- 
ocotyledons, 43 

Klamath Region, Vegetation history of 
the Pacific Coast states and, 5 

Kobresia: bellardii, 232; macrocarpa, 230 


Laurus nobilis L., An anatomical study 
of the secondary tissues in roots and 
stems of Umbellularia californica Nutt. 
and, 205 

Lepidium: bourgeauanum, 78, 87, fig. 
88; campestre, 78, 79; densiflorum, 78, 
84, var. densiflorum, 84, var. elongat- 
um, 87, var. macrocarpum, 86, fig. 88, 
var. pubicarpum, 87; heterophyllum, 
78, 80; latifolium, 78, 81; oxycarpum, 
78, 83; perfoliatum, 78, 79; ramosissi- 
mum, 78, 90; ruderale, 78, 83; sati- 


1962] 


vum, 78, 81; The genus in Canada, 77; 
virginicum, 78, 83 

Limodorum abortivum, 64, fig. 65 

Lycium: A new species in Nevada, 122; 
cooperi, 125; macrodon, 125; puber- 
ulum, 125; rickardii, 123, fig. 124; 
shockleyi, 125 

Lysurus: mokusin, 35, figs. 37, 39; sul- 
catus, 40, fig. 39 


Malacothrix: clevelandii, 258, Three new 
species related to, 258; fendleri, fig. 263 
foliosa, fig. 263, similis, 262, fig. 263; 
sonorae, 264, fig. 263; stebbinsii, 265, 
fig. 263. 

Mason, C. T. Jr., and R. H. Hevly: Ad- 
ditions to the aquatic flora of Arizona, 
32 

Mason, H. L.: Milo S. Baker, 155; Re- 
view, The Little Hill, a chronicle of 
the flora on a half acre at the Green 
Camp, Ringwood, New Jersey, 268 

Mathew, K. and P. H. Raven: A new 
species of Cryptantha (section Circum- 
scissae) from California and two re- 
combinations (section Circumscissae 
and section Angustifoliae), 168 

McClintock, E.: Review, Southern Cali- 
fornia Gardens, an Illustrated His- 
tory, 204 

Melampsora artica, 202 

Melicope: Burttiana, 165; grandifolia, 
165; (Solomon Islands), a new name 
for, Taxonomic and nomenclatural 
notes on Platydesma (Hawaii), and 
161; spathulata, 165 

Merendera trigyna, 54, fig. 55 

Mimulus: arvensis, 149; Chromosome 
counts in the genus, 104, on the sec- 
tion Simiolus, 141; cordatus, 149; 
glabratus var. fremontii, 150, var. par- 
viflorus, 150, var. utahensis, 150; 
glaucescens, 149; guttatus, 148, The 
chromosomal homologies of, and its 
allied species and varieties, 141, subsp. 
guttatus, 148, subsp. litoralis, 148, var. 
puberulus, 148; laciniatus, 149; laxus, 
149; luteus, 150; lyratus, 149; nasutus, 
149; pilosiusculus, 150; platycalyx, 
149; tilingii var. corallinus, 149, var. 
tilingii, 149 

Monocotyledons, Foliar xeromorphy of 
certain geophytic, 43 

Mooring, J.: Review, 
flowering plants, 171 

Moran, R.: Cneoridium dumosum, 272; 
Rufus Davis Alderson, 224; Stegno- 
sperma cubense and Gossypium klotz- 
schianum davidsonii not known in the 
Revillagigedos, 108 

Mosquin, T.; Eschscholzia covillei Greene, 
a tetraploid species from the Mojave 
Desert, 91 


Taxonomy of 


INDEX 210 


Mukherje, B. B., and R. K. Vickery, 
Jr., Chromosome counts: in the genus 
Mimulus (Scrophulariaceae), 104; in 
the section Simiolus of the genus Mi- 
mulus (Scrophulariaceae). V. The 
chromosomal homologies of M. gut- 
tatus and its allied species and vari- 
eties, 141 

Muller, C. H.: A new species of Lycium 
in Nevada, 122; A new species of 
Quercus from Baja California, Mexico, 
186 

Mulligan, G. A.: The genus Lepidium in 
Canada, 77 

Muscari: comosum, 54; fig. 55; racemo- 
sum, 56, fig. 55 


Notes and News, 31, 108, 140, 236, 269 


Oenothera, subgenus Chylismia, The sys- 
tematics of, 236 

Orchis mascula subsp. pinetorum, 66, 
fig. 65 

Ornduff, R.: Marcus Jones publications, 
272; Review, Vascular Plants of the 
Pacific Northwest, 74 

Ornithogalum: armeniacum, 57, fig. 59; 
narbonense var. pyramidale, 58, fig. 59 


Palmer, Edward, visit to Guadalupe Is- 
land, Mexico, in 1875, 1 

Parasitism in Pedicularis, 192 

Parmeliopsis placorodia, discovery of the 
lichen, in western North America, 31 

Payne, W. W.: The unique morphology 
of the spines of an armed ragweed, 
Ambrosia bryantii (Compositae), 233 

Pedicularis: attollens, 195, fig. 198; cren- 
ulata, 195; densiflora, figs. 197, 198, 
subsp. aurantiaca, 194, subsp. densi- 
flora, 194; dudleyi, 195; groenlandica, 
195; Parasitism in, 192; racemosa, 195; 
semibarbata, 195, figs. 197, 198 

Pelton, J.: Factors influencing survival 
and growth of a seedling population 
of Arbutus menziesii in California, 237 

Peterson, R. S.: Wyoming Pinyon re- 
visited, 269 

Phakopsora crotalariae, 202 

Phippsia algida, 229 

Phormidium: A new name in the algal 

genus, 108; anabaenoides, 108 

Pinus: jeffreyi, 126; ponderosa, 
washoensis, 126 

Plagiobothrys austinae (Greene) John- 
ston: A new addition to the Oregon 
flora, 108 

Platydesma: campanulata, 165, var. 
macrophylla, 165, f. coriaceum, 165, 
var. pallida, 165, var. pubescens, 165; 
(Hawaii), and a new name for a 
Melicope (Solomon Islands), taxo- 
nomic and nomenclatural notes on, 


126), 


276 MADRONO 


161; oahuensis, 165; spathulatum, 
165, figs. 162, 164, var. pallidum, 165, 
var. pubescens, 165 

Poa bulbosa, 48 

Polemoniaceae, 
ment of, 270 

Ponderosa, Jeffrey and Washoe pines in 
northeastern California, Some recent 
observations on, 126 

Potamogeton richardsonii, 32 

Puccinia: acrophila, 202; agrimoniae, 
201; bouvardiae, 201; coronata, 202; 
deschampsiae, 202; drabae, 203; es- 
clavensis, 201; eumacrospora, 201; 
monoica, 203; montanensis, 203; mu- 
senli, 203; pagana, 203; pattersoniana, 
203; wulfeniae, 203; xanthiifoliae, 201 


A controversial treat- 


Quercus: A new species from Baja Cali- 
fornia, Mexico, 186; cedrosensis, 188, 
fig. 189; vaccinifolia, 188, fig. 189 

Quick, C. R. and A. S. Quick: Germina- 
tion of Ceanothus seeds, 23 


Reviews: Allard, R. W., Principles of 
plant breeding, 140; Avery, A. G., S. 
Satina, and J. Rietsema; Blakeslee: 
the genus Datura, 72 ; Hitchcock, C. L., 
A. Cronquist, M. Ownbey, and J. W. 
Thompson: Vascular plants of the Pa- 
cific northwest, 74; Padilla, V., South- 
ern California gardens, an illustrated 
history, 204; Porter, C. L., Taxonomy 
of flowering plants, 171; Takhtajan, 
A., Die Evolution der Angiospermen, 
70; Thomas, J. H., Flora of the Santa 
Cruz Mountains of California, 138; 
Todd, A. O., The Little Hill, a chron- 
icle of the flora of a half acre at the 
Green Camp, Ringwood, New Jersey, 
268 

Rumex acetosa, 231 

Sphenophyllum: majus, characters of, 
107; nymanensis sp. nov. from the 
Upper Pennsylvanian, 106, fig. 107 


LVol. 16 


Sprague, E. F.: Parasitism in Pedicularis, 
192 

Standley, P. C.: Trees and shrubs of 
Mexico, 140 

Stegnosperma cubense and Gossypium 
klotzschianum davidsonii not known in 
the Revillagigedos, 108 

Stone, B. C.: Taxonomic and nomen- 
clatural notes on Platydesma (Hawaii) 
and a new name for a Melicope (Sol- 
omon Islands), 161 


Typha angustifolia, 32 


Umbellularia californica Nutt. and Laur- 
us nobilis L., An anatomical study of 
the secondary tissues in roots and 
stems of, 205; figs. 207, 209, 211, 213, 
216 

Uredinales, North American, nomencla- 
ture, life histories, and records of, 201 


Wagner, W. H., Jr.: Cytological obser- 
vations on Adiantum & tracyi C. C. 
Hall, 158 

Washoe, Ponderosa, and Jeffrey pines in 
northeastern California, Some recent 
observations on, 126 

Weiler, J. H.: A new species of Downin- 
gia, 256 

Wells, P. V.: A subarborescent new Erio- 
dictyon (Hydrophyllaceae) from San 
Luis Obispo County, California, 184 

Wherry, E. T.: A controversial treatment 
of the Polemoniaceae, 270 

Whitaker, T. W.: Review, Principles of 
plant breeding, 140 

Whittaker, R. H.: Vegetation history of 
the Pacific Coast states and the ‘‘cen- 
tral” signiflicance of the Klamath Re- 
gion, 5 

Wiggins, I. L.: To Albert W. C. T. 
Herre, 102 


Xeromorphy, Foliar, of certain geophytic 
monocotyledons, 43 


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Volume XV, 1959-1960 Pe) ESR “ae OLOg 
Volume XVI, 1961-1962 . . .  . 12.00 


Single numbers: 


Volumes Panda: . fio) 82 8. 50 
Volumes II through XIV. . ..... 1.00 
Volume 0Vig 2" cc! Tyan 0 a ae P25 
Voltimesevil ios ees ee ee 1.50 


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Department of Botany 
University of California, Berkeley 4, California 


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