Skip to main content

Full text of "Madroño : a West American journal of botany"

See other formats 


“Tne ont Ae erage tees oy em ae Re thee ieee % 
foe D NM tA gn ym a Rntione” 
en Menten sh ahtbe wee tie-Rabiaperone ia by MegtD Sti shame weiter iyecn,s > 


Oem we rarity 


AP pee ONS e eatw Ah wee sh gion om fie er 4 


Fe Oe mathe ee x 


Fe ale eetedlome ty 


hw te ee Set san se 


-- sR em eee 


eo 


a a 

ee ee re 
eer ees oe 

seam tenes “tempering seme ty se 
- we eee eel 


« : ; z 


TTR Re Reman te Pte ee 


bares! ews tremee ing a 
Pg OO COO TI GIs yn Smee gd CO ER ee 


eh mt ee a Nig br tee te he ep ites go 
ey ee ea. eee 


OO OO SE 
AR en rotenone nadine dhe Metmpe e herian se es mga A 
Oe Fle men aethee Sgimeser men 


oN ee Ret IA NON A hay yor Eh, 


Se eee ON Mache ys pag A el ment Be Ny 
Se att Oe ae = 


tage rhe fe Catt ne 


oe ee Mw hig mae 
wo theta tne ds the Meaceeatio 
80 = eh ear Pag 

Nea ee A Mee cee 26 eit es ete hee 
2 ae een atte ye 


et a tee em ang EE Oem the ae + oe ed 
0 AP ndite Newnan Mn Fase ae. a apt gelow ne ne 


Oe ee Ce 
A Ae ee gene ebeweretie en feng ope ety 
yO Nee Nee metie te ttemne clety bes 


oO pte en pe 


Me Oe bape Mee wes ee 


ip ree Be geny Hr he xt Sue 
SOIT eo apm nt peepee 
Pe Saget She yd ee tee Oe 
i abtaatidt nttet oe oe 
0 FOI gd AN He me ew ng toe 


ee ee oe ee 


SL te me eee re Se 


eM ne he rym 


tc eM ee nO ie ale 3 Be tg th once 


er en 


een eee 


te rene wee cate tee 


FAP She ee tere RO hd 


see Ney Re ap he rye ee de 
Ae Oye eth onion . 
ee ae 

Te eke ge eee eee 


1 meticgelonene Daete he SMe He Fie sy ee 


Pee okey Gat ate dey See 


eee Menta He shee 


tg tt ty 


ak a epee a foe ne 


a ee 


Fe ee 


Fe We thes oie far ee 


pate rene te aR dita we Meet 
eae ANE ARs AEM nm we Hho 8 
whey PE NbN ttn iwi, COS Peseta oy Mont 


El ea ip 1 Ripa gr tty dee ie Ger 
pryare tee ane Me 


he see hncia ota ee UR ven 


ee mine 


i - 


i he ee 


pl ath Ns pict tel he a Te RAE SO ee Mea arte dene ores & 


eee cab Le eT 
ABN Sete ean gg hie ws 


re actin eb gett 


fe tender 


~ en tee 


, 7 7 ue 
‘ ® 7 1 


Steet a eee ey ae 


ee ee ne ae 


a eb ees 


Ci ees Ube eo nr roo ina 
Lote Net et tee AMR ime Snes MUSEO ole ats, 
Se Re ete ee Pete enh ett Som AF NES ete 


~ mae ~ 
el ee eee ee ae 


dim tos Tbe ee ae SU oe 
. mre Hen Se tes sae am 
+ et ag ot aonaeees lg 
avis oy - . 
whee 7+ : 


£8 hah BOHR Nea en. sete etntne = 


“Yo 2 Ee a ee ee re 


ee eet dere ey etait 


Pa a OMe ene ke 
vee ae 


a en te ee 


9B et We SAS et etree een 


Tete bates: greeters te 


ne neat 8 ON ay BIG ine mee oe 


* ong ee 


oo Re Parte sbi thne ase 


ae 


ree ene) 
eo. Hage Atte Latest 


- oy oe e 


ven aD eS 


<3) Sere aM ae epee” 
Se en ere 
= Se ena Ree ethene er gale nee hee 


i dalebaghtmieanten iia catenin a ee ee 
<2 eee een lente 


tah than 


weather’ 


Heateswean 0 Oe tg tt trim neta = rel age Dw lated tie the Havens 
eee ee et a Or ee eT a 


eh ha eit et ed eel oh 


Meant mthete (nt gevmibeuie beth! 
Vag 8 he fe alg Aa eager tin tat 
HRA OL ite at g he pe eh Smet 


AV a WS NE Rte hee ag He al in TO Mc Nt che hay hen 
en ere 


Ce ra ee 
Ae AS ye ert itghe ines 


SERN eee HS Hgene eat oe oR Dette poten Mihai merge en. — ty cthe te ee tt at 


Oe oe Veen ette imei ete, 
ag Penitigars a 


Hime ee eee a Sage ee tag Sage the 
bbibiihdeedincnh a ooh a eee ee 


Nigditinitintitie, oe ee 
ne ae Hae Be ote ae Matyas ae 
ti Aele ate 

Tape eee abgiee 
TH ate ay miele ey bain EB ely. 


et dees 
wet wn eee 
aoe there tee eee 


ete nee te eee * 


Mi Aine ot 


a ee 
RR ee aie Me 


eiadiniden clint Matted tee 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


VOLUME XV 
1959-1960 


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 

JoHN F. Davipson, University of Nebraska, Lincoln 
Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts 
Mivprep E. Marutas, University of California, Los Angeles 
MARION OwWNBEY, State College of Washington, Pullman 


Tra L. Wicorns, Stanford University, Stanford, California 


SECRETARY, EDITORIAL BOARD 


ANNETTA M. Carter, Department of Botany, University of California, Berkeley 


BUSINESS MANAGER AND TREASURER 


WINSLow R. Briccs, Department of Biology, Stanford University, 
Stanford, California (1959) 


Joun H. Tuomas, Dudley Herbarium, 
Stanford University, Stanford, California (1960) 


Published quarterly by the 
California Botanical Society, Inc. 


2016 Life Sciences Building, University of California, Berkeley 


Printed by Gillick Press, Berkeley, California 


R. C. Frampton Photo, Claremont 


To PHittp ALEXANDER Munz, teacher of botany, student of the Ona- 
graceae, author of two California floras and, until his recent retirement, 
Director of the Rancho Santa Ana Botanic Garden, the California Botani- 
cal Society dedicates this, its fifteenth volume of Madrono. 

During his many years of teaching, Philip Munz has held professor- 
ships at Pomona College, Cornell University, and presently at Claremont 
Graduate School. In addition to the two floristic works, his scientific writ- 
ings have included monographic studies of both North and South Ameri- 
can plant groups. On quite another level, however, many regard his 
greatest achievement to be what he has made of the Rancho Santa Ana 
Botanic Garden of the native plants of California. Here, during the past 
fifteen years, all his talents and accomplishments have been turned to the 
purpose of creating a model botanical institution, one which has become 
an increasingly renowned center for research upon all phases of the Cali- 
fornia flora. Well-founded in his judgments, forward-looking yet moder- 
ately conservative in his administration, encouraging to achievement in 
other fields than his own, wise in his choice of personnel, his rare qualities 
of leadership have brought his institution to a point of high repute in the 
botanical world. Beloved by his colleagues and genuinely young-in-heart, 
he is looked upon by his staff as the benevolent head of his “family.” 


CONTENTS 
PAGE 
Frontispiece: Philip Alexander Munz 


Survival of transplanted Cupressus in the pygmy forests of Mendocino County, 


Bar MeN ae UCI, VC VEDI 97 areas ea ae Sole ee eee i es 1 
An interspecific cross in Cucurbita (C. lundelliana « C. moschata Duch.)— 

OM IUCUSMAUY. VU JLTE IC Of, eee ta he ee reg sc ee rong: Set Pela eaten aeanc reer aeece 4 
The reproductive structures of Schinus Molle (Anacardiaceae ) — 

TELE INA AG TAY ONG WITT. Pen eee Se neds Rie eR SA LISLE aA TT ENE EO OMe OEE eee 14 
Development of the megagametophyte in Liquidambar styraciflua L.— 

SHA as Dine) Oe ee OE a 0D. Se Ree ree Ne NTE NS Noe 25 
"RUpiig lis eee ae ea co 29, 62, 94, 158, 192, 221, 249 
ING tecmanialieINiewe- meee taste 22 oe ee 32705 200m 123, 1G Om102, 1200 
Red algal parasites occurring on members of the Gelidiales— 

Kung-Chu Fan and George F. Papenfuss....................000----20000- seen ON cece 33 
Xerophyllum tenax, squawgrass, its geographic distribution and its behaviour 

on Mount Rainier, Washington—Sue Merrick Maule_..........0000000000-02-22--- 39 
Documented chromosome numbers of plants.......00.0000000000000000 eee 40, 219 
Typification of Prosopis odorata Torr. and Frem. (Leguminosae) — 

FT PYOC eS IU S 0 1 ee 3 Aen Sd ane Se yo cs te ace age ees nw) 
Two new species of Helianthus from New Mexico—R. C. Jackson_...-....-........... 54 
Chromosome counts in the section Simiolus of the genus Mimulus 

(Scrophulariaceae). IJ1I—Barid B. Mukherjee and R. K. Vickery, Jr............. ou) 
Factors affecting the distribution of Ponderosa and Jeffrey pines in California— 

POUT sl ORCS 8 NA Slee oe a aa TOE SETAC A AT eS RO PS PO REN OP OR Tee Oe VR BD 65 
Vivipary in Cordyline australis Hook—Howard J. Arnott........0...000.00.00020000000---- = 71 
Studies on secotiaceous fungi VI. Setchelliogaster Pouzar— 

Rolf Singer and Alexander H. Smith 13 
Ceanothus seeds and seedlings on burns—Clarence R. Quick.........-..........--.----------- 79 
Chromosome numbers of California plants, with notes on some cases of 

Gylolosical mierest—_ Richard | SHOW: 2k zai te eter acest ee eee eee 81 
Chromosome counts in the genus Gayophytum—David G. Dixon...........2.0............ 90 
The grass genera Orcuttia and Neostapfia: a study in habitat and morphological 

Speclatization=—Deecher” Cram pron. calcd ekeeecteees ase ee 97 
The taxonomic relationship between Picea glauca (Moench) Voss and 

Peengelmannicbarry— J) MAG. Talon 2 a ee el Bel 
Field studies of natural hybridization in the Oregon species of Iris L. subsection 

Calitormicae Diels —Ouentin De Clar R SON icteric tones ee ee 115 
Variation patterns in four clones of Mertensia ciliata—Jeaneite S. Pelton............ 123 
New combinations in Aster—Rowxand S. Perris .........000. 0.2. 220ccc eect ee cence ee ceeeeeeeeeeeeeeeeeee 128 


The distribution of dwarf mistletoes, Arceuthobium, in California— 
BID RSSLLE GU ewe tpg eens nce oer re eee case eee cecal Wace et ee 129 


Nuclear cytology of the California mouse-tails (Myosurus)—Donald E. Stone.... 139 


Variation in section Trigonophyllae of Nicotiana—Philip V. Wells......0.000000000....... 148 


ili 


Studies on secotiaceous fungi VII. Secotium and Neosecotium— 


Rolf Singer-and Alexander Ty Sites esse see ee 152 
The reproductive structures of Fraxinus velutina (Oleaceae ) — 

Herbert F AC op clang ee ee 161 
A new Silene from northwestern California—A. R. Kruckeberg.......00......0...2000000--.. 172 
Freed Hofiman—J oli Ee MOV rts O91 cs case oe eee 178 


Cleared Cardiocarpon late-alatum Lesq., Cordaitean seeds from Michigan— 
TE DD GVUDS ON vs i ic ed OE sea a = eg ee ne ge 180 


The basic chromosome number of the genus Neptunia (Leguminosae- 
Mimosoideae)—B. L. Turner and O. S. Fearing......-.......20.20.cs0s0meetoeeeesceeeseeeses 184 


Hybridization and instability of Yucca—John Milton Webber..............0....00.00...... 187 
Sanicula deserticola, an endemic of Baja California— 

Peter. Ravenand Mildred E. Matis. 193 
A new species of Valeriana from Brazil—Frederick G. Meyer ........0.......02000000220000000--- 197 
Studies in western violets, IX. Miscellaneous species in the sections Nomimium 

and Chamaemelanium—Malo S. Barer. ....2 cto ices ct hecsel ee eee eee 199 
Chromosome numbers in Silene (Caryophyllaceae). I1—A. R. Kruckeberg............ 205 
A new species of Zinnia from Mexico—A. M. Torres .....0..0.2ccccccccccccseeecceeeeeeeeeeeseeee 215 
A comment on cold susceptibility of Ponderosa and Jeffrey pines— 

WLS WW CNC 2 scadccxccctcasemco et cacecae eee os ce ee 217 
Distributional notes on plants of the Warm Springs area, Oregon— 

Robert Ornduy and David H. French. == 225 
G. Thomas Robbins (1916—1960)—Rimo Bacigalu pi.......--0.220000200000000ccceceeeceeeeeeeee eee 231 
Studies in the perennial gentians: G. newberryi and G. tiogana— 

Gar es: TMG SOT IF rts loca eee 233 


Chromosome counts in the section Simiolus of the genus Mimulus 
(Scrophulariaceae). IV.—Barid B. Mukherjee and Robert K. Vickery, Jr..... 239 


Flowering responses in Phacelia sericea and P. idahoensis—George W. Gillett........ 245 


DNL. GOs ViO LUI Vee ese eee eee ee 253 


ERRATA 


Page 25, line 12: for Juliano, J. M., read Juliano, J. B. 
Page 48, line 34: for 1951 read 1925. 

Page 51, line 11: for Connty read County. 

Page 71, line 15: for 1928 read 1950. 

Page 138, line 7: for existes read exists. 

Page 138, line 16, for 1914 read 1913. 

Page 139, line 13: for 1914 read 1913. 

Page 188, line 1: for hybirds read hybrids. 

Page 199, line 20: for terrano read terraneo. 

Page 199, line 21: for subterreno read subterraneo. 
Page 199, line 26: for marginis read marginibus. 

Page 213, line 52: for australe read australis. 

Page 220, line 30: for phyllocephallus read phyllocephalus. 


lv 


RONO 


VOLUME 15, NUMBER 1 JANUARY, 1959 


M 


Contents 


PAGE 


SURVIVAL OF TRANSPLANTED CUPRESSUS IN THE PyGMy 
Forests OF MENDOCINO CouNTY, CALIFORNIA, 


Calvin McMillan 1 
AN INTERSPECIFIC CROSS IN CUCURBITA (C, LUNDELLI- 
ANA X C. MOSCHATA Ducu.), Thomas W. Whitaker 4 


THE REPRODUCTIVE STRUCTURES OF SCHINUS MOLLE 
(ANACARDIACEAE), Herbert F. Copeland 14 


DEVELOPMENT OF THE MEGAGAMETOPHYTE IN LIQUIDAM- 
BAR STYRACIFLUA L., Franklin F. Flint 25 


Reviews: John Thomas Howell, Peter H. Raven, and 
Peter Rubtzoff, A Flora of San Francisco, California 
(John H. Thomas); Gerd Kriissmann, Die Nadelgehdlze 
(Baki Kasapligil) 29 


NOTES AND NEws 32 


A WEST AMERICAN JOURNAL OF BOTANY > PBN 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


eae we 
AP Atiicr _ 

HZAAMVESON, 
oé : ., IN 


\ 
MAR 4 1959 


Ht 
\S ZjepaRtY 7 


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 

$4.00 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 
LymMAN BENSON, Pomona College, Claremont,. California. 
HERBERT F. COPELAND, Sacramento College, Sacramento, California. 
Joun F. Davipson, University of Nebraska, Lincoln. 

Ivan M. JounsTon, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MitpreD E. Martutas, University of California, Los Angeles 24. 
Marion OwnsBEY, State College of Washington, Pullman. 

Ira L. Wiccins, Stanford University, Stanford, California. 


Secretary, Editorial Board —ANNETTA CARTER 


Department of Botany, University of California, Berkeley. 


Business Manager and Treasurer—WINSLOW R. Briccs. 
Department of Biology, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: James R. Sweeney, San Francisco State College, San Francisco, Cali- 
fornia. First Vice-president: Baki Kasapligil, Mills College, Oakland, California. 
Second Vice-president: Henry J. Thompson, Department of Botany, University of 
California, Los Angeles, California. Recording Secretary: Mary L. Bowerman, De- 
partment of Botany, University of California, Berkeley, California. Corresponding 
Secretary: Francia Chisaki, Department of Botany, University of California, Berke- 
ley, California. Treasurer: Winslow R. Briggs, Department of Biology, Stanford 
University, Stanford, California. 


SURVIVAL OF TRANSPLANTED CUPRESSUS IN THE PYGMY 
FORESTS OF MENDOCINO COUNTY, CALIFORNIA 


CALVIN MCMILLAN 


In a recent study of the edaphic restriction of Cupressus and Pinus in 
central California (McMillan, 1956), certain species from highly acid 
soils and from serpentine soils were investigated. Soil tolerance studies 
conducted both under greenhouse conditions and in large outdoor bins 
were used in this inquiry. 

One phase of the investigation not previously reported involved the 
transplanting of seedling trees to the habitat of the pygmy forest in Men- 
docino County. At the time of the transplanting no extended analysis of 
seedling survival was anticipated. Since it has been possible to continue 
the observations, however, this report will evaluate the survival over a 
period of seven years. 

The pygmy forests have been described adequately elsewhere (McMil- 
lan, 1956), but certain features of the transplant site on the coastal 
plateau between the Little and Albion rivers are pertinent. The trans- 
plant area, approximately one mile southwest of the Mendocino County 
Airport, was on the property of Miss Jean MacGregor Boyd. it had been 
subjected to previous burnings and was covered by a low growth of trees 
and shrubs. The site was included within the narrow restriction of both 
Cupressus pygmaea (Lemm.) Sarg. and Pinus bolanderi Parl., and both 
species were represented by numerous individuals varying in height from 
25-100 cm. Vaccinium ovatum Pursh, Rhododendron californicum Hook., 
and Gaultheria shallon Pursh were common shrubs of the area. -Jrcto- 
staphylos nummularia Gray, a species which does not crown-sprout fol- 
lowing burning, was represented by a few small shrubs. In adjacent areas 
with no record of recent burning, Arctostaphylos was common with Pinus 
muricata Don. The dense Sequota-Pseudotsuga forest, common on the 
margins of the plateau, was adjacent to the transplant site. The soil in the 
transplant garden is a ground-water podsol with a pH of 3.8-4.0. This 
ashy-colored soil, which is common throughout the areas of the pygmy 
forest, is usually less than a foot in depth and is one of the most acid soils 
in California. 

Seedlings of the various strains of Cupressus, including those of C. 
pygmaea, C. goveniana Gord., C. abramsiana C. B. Wolf, and C. sargentii 
Jeps., were transplanted from the greenhouse at Berkeley to the pygmy 
forest in November, 1950. Six of each strain were planted in each of two 
series. One group was protected from browsing animals by a large wire 
screen cage, while the second series remained unprotected. The unpro- 
tected seedlings were severely damaged and could not be used in the 
study. The cage protected the other seedlings throughout the investiga- 


Maprono, Vol. 15, No. 1, pp. 1-32, February 5, 1959. 


Bal 


SONTAN 
SMisHS ITION 


FFB I Q 1958 


2 MADRONO [Vol. 15 


tion. Measurements were recorded at intervals beginning at the date of 
planting and continuing until August, 1957. 

In April, 1952, individuals of the two strains of C. pygmaea from Men- 
docino County (one from the pygmy forest area, the other from Anchor 
Bay) were dark green and vigorous. Individuals of the two strains of C. 
goveniana from Monterey County (one from Huckleberry Hill and the 
other from behind Point Lobos) and the one strain of C. abramsiana 
(from Bonny Doone in the Santa Cruz Mountains) were lighter green 
and lacked vigor. The average growth increment since transplanting 
showed clearly that the C. pygmaea strains were the most vigorous even 
though the actual increase in height for all strains was slight. Seedlings 
of C. sargenttu (from a serpentine area on Mt. Tamalpais in Marin 
County) were barely surviving in 1952. 

Over the same two-year period, seedlings were grown at Berkeley in 
large outdoor bins containing soil from the habitat of the pygmy forests. 
These bins were used until 1952 and at that time the responses of the 
strains reversed the trend indicated in the transplant garden. The vigor 
and height of the two strains of C. pygmaea was less than that of either 
C. goveniana or C. abramsiana. In contrast, seedlings of all strains grew 
well at Berkeley in bins containing an agricultural type of soil. The C. 
pygmaea strain from the pygmy forests was extremely vigorous on the 
agricultural soil and had an average height of 82 cm. after the 2-year 
growth period (McMillan, 1956). This contrasts sharply with a 3.3 cm. 
average height increase of the same strain when transplanted to the pygmy 
forest habitat. 

In August, 1954, the transplant garden was observed after an interval 
of two years. The growth differential which had been apparent in 1952 
was intensified. The trees of C. pygmaea were much more vigorous than 
those of either C. goveniana or C. abramsiana. Of the surviving strains, 
only that from Pt. Lobos failed to show an increase in height over 1952 
measurements. All of the individuals of C. sargentiz, the only strain from 
serpentine soils, had died during the interval. 

Observations at the transplant garden in August, 1957, indicated con- 
siderable development of both strains of C. pygmaea. Somewhat greater 
vigor characterized the individuals of the pygmy forest strain. Average 
growth since transplanting was 11.8 cm. The average height increase for 
the Anchor Bay strain was 9.3 cm. These strains of C. pygmaea produced 
pollen-bearing cones and only the pygmy forest strain had produced a 
mature seed-bearing cone. 

Much less growth and a reduction in the number of surviving trees 
were noted in C. goveniana and C. abramsiana. None of the trees of the 
Point Lobos strain of C. goveniana survived. The four surviving trees of 
the Huckleberry Hill strain had grown very slightly and none was vigor- 
ous. Only two of the C. abramsiana individuals were surviving and these 
showed only a trace of height increase. 

The growth pattern of dwarfed trees of C. pygmaea results from a 


1959] MC MILLAN: CUPRESSUS 3 


unique tolerance for the highly acid soils of the pygmy forest area. At the 
transplant site, small, cone-bearing trees, varying between 25 and 50 cm. 
in height, had 21 growth rings when measured in 1952. These trees were 
less than 70 cm. in 1957. Other trees, in pygmy forest areas adjacent to 
the transplant site, had over 100 growth rings, a 6-10 cm. diameter and 
a height of less than 2 m. A few larger trees (30-50 m.), such as those 
measured by Mathews (1929), indicate the type of growth which can be 
achieved under conditions which support mostly Sequoza and Pseudotsuga. 

Although the survival pattern in Cupressus indicates that the strain 
from the pygmy forests is the best fitted for growth on the highly acid 
soils in Mendocino County, it must not be inferred that the other strains 
are incapable of the growth pattern producing a pygmy forest. For ex- 
ample, at Huckleberry Hill, trees of C. goveniana closely resemble C. 
pygmaea. Although occasional tall trees of C. govenitana occur with Pinus 
radiata Don, the majority of the trees with P. muricata give an appear- 
ance of a pygmy forest. The inability of the Huckleberry Hill strain to 
grow well at the transplant garden in Mendocino County is particularly 
difficult to explain on an edaphic basis because of the marked similarity of 
the soils of the two areas. Climatic conditions of Mendocino County sug- 
gest a more likely explanation. However, trees of C. macrocar pa, a species 
also restricted to the Monterey Peninsula in its natural distribution, are 
thriving and reproducing along the Mendocino County coast. Unless the 
climatic tolerances of C. macrocarpa and C. goveniana differ markedly, 
the explanation would lie possibly in the action of the highly acid soils in 
conjunction with the cooler temperatures of the more northern location. 
Extended periods of freezing temperatures and of dry summer conditions 
occurred between 1954 and 1957. This may have provided the critical 
point for reduced survival among all of the strains from the more south- 
erly localities and, in particular, produced conditions beyond the tolerance 
of the Point Lobos strain of C. goveniana. The loss of seedlings of C. 
sargenti early in the study may indicate a low survival potential on 
highly acid soils by strains from serpentine soils.? 

Studies of the nature of restriction of Cupressus pygmaea indicate that 
the species is not confined to the pygmy forests because of an inability to 
grow elsewhere. The restriction results, in part, from the tolerances of C. 
pygmaea for conditions resulting in a dwarfed form and, in part, from 
the lack of tolerance by taller forms, such as Sequoia and Pseudotsuga, for 
growth on the shallow, highly acid soils. Furthermore, these survival 
studies indicate that C. pygmaea possesses a tolerance not shared by the 
strains of other species of Cupressus for conditions of the pygmy forest 
habitat. 


1 Preliminary studies indicate that day length factors were not critical in the dif- 
ferential survival of the Huckleberry Hill and Mendocino seedlings. In greenhouse 
studies, seedlings of the Monterey strain as well as seedlings of both Mendocino 
strains responded similarly to a range of light periods: 8-hour, 12-hour, and 16-hour. 


4 MADRONO [Vol. 15 


Selective influences on the coastal plateau of Mendocino County have 
sorted out a combination of species with unique qualities for survival in 
one of the most extreme soil situations in California. The pygmy forests 
which have resulted from this selective action include much of the natural 
distribution of both Cupressus pygmaea and Pinus bolanderi. The gigantic 
Sequota-Pseudotsuga forests which grow in adjacent portions of the 
coastal plateau present an amazing contrast in vegetational selection. 


Department of Botany 
University of Texas 
Austin, Texas 


LITERATURE CITED 


MaTHEws, W.C. 1929. Measurements of Cupressus pygmaea Sarg. on the Mendocino 
“pine barrens” or “white plains.”” Madrofio 1:216-218. 

McMittan, C. 1956. The edaphic restriction of Cupressus and Pinus in the Coast 
Ranges of central California. Ecological Monographs 26:177-212. 


AN INTERSPECIFIC CROSS IN CUCURBITA 
(C. LUNDELLIANA BAILEY x C. MOSCHATA DUCH.) 


THOMAS W. WHITAKER! 


As one aspect of a comprehensive study of the origin and relationship of 
the cultivated species of Cucurbita, C. lundelliana Bailey, a non-cultivated 
species, was crossed with C. moschata Duch., one of the five cultivated 
species of the genus. In Cucurbita successful crosses between truly wild 
species and domesticated ones have not been hitherto reported. Essentially 
the cross C. lundelliana X C. moschata combines two genotypes, the one 
(C. lundelliana) a wild species and the other (C. moschata) with a long 
history of cultivation. The hybridization experiments reported here were 
made with the idea that the compatability relations might indicate direc- 
tions in which to search for the common ancestor of the cultivated group, 
and perhaps suggest in a general way the area where the cultivated forms 
were domesticated (Whitaker, 1956). Furthermore, it was thought that 
the results would contribute to an understanding of the heritability of 
characters such as large fruit, large seed, soft rind, etc., which have value 
under cultivation. The results reported here provide partial answers to 
some of these questions. 

MATERIALS AND METHODS 


Cucurbita lundelliana, the Peten gourd, is endemic in Central America. 
It has been collected in Guatemala, British Honduras, and southern 
Mexico (Yucatan). The plants are strong, vigorous annuals, with fine, 


1 This study was aided by a grant from the American Academy of Arts and 
Sciences. I am much indebted to Professor Edgar Anderson, Curator of Useful Plants, 
Missouri Botanical Garden, for helpful suggestions and advice. My thanks are due 
to G. A. Sanderson, Agricultural Aid, for the photographs and to M. A. McClure, 
Technician, for preparing figure 4. 


1959] WHITAKER: CUCURBITA 5 


wiry stems and deeply lobed leaves having a greyish-green cast (fig. 1B). 
The flowers of both sexes are large, showy and upright (fig. 1E). The 
fruits are almost round, dark green, often striped, and have hard rinds, 
or shells (fig. 2A). The flesh is greenish white and the placenta is solid. 
The seeds are comparatively small and numerous and have characteristic 
broad, wavy margins. 

Cucurbita moschata cv. Long Genoa Queen is an old and little-known 
cultivar quite typical of the species. The modern cultivar Butternut of 
C. moschata probably originated as a selection from Long Genoa Queen. 
The plants have good vigor, large, ovoid leaves, slightly triangular lobed 
(fig. 1A); long fruits (19 to 36 inches), with an enlarged terminal por- 
tion that contains the seeds (fig. 2B). The flesh is dark-orange, moist and 
slightly stringy. 

Both C. lundelliana and C. moschata have 20 pairs of chromosomes. 
Unfortunately, it has not been possible to study cytologically the pro- 
genies of the cross between them. 

Matings of C. lundelliana and C. moschata cv. Long Genoa Queen pro- 
duced fruits with numerous fertile seeds. The F, proved to be self-fertile 
and cross-fertile with each parent. In the summer of 1954, the parents 
and the F; were grown in the experimental garden along with several Fo 
and backcross progenies. The analysis of the data obtained from the 
measurements and observations of these progenies constitutes the basis for 
this report. 

Measurements were made of leaf blade length and width, lobe depth 
and petiole diameter. For male and female flowers the following measure- 
ments were obtained: length of corolla. corolla limb, senal, style plus 
stigma. staminal column, and ovary length and diameter. Other measure- 
ments included fruit length and diameter and length and diameter of sev- 
eral typical seeds. Fruit color, rind hardness, fruit construction, flesh 
color, and placenta type were also recorded. 

The large sprawling plants of most species of Cucurbita require exten- 
sive space to mature. For this reason it is impractical to raise by conven- 
tional methods the number of plants required for a convincing genetical 
analysis. A more practical method of analysis adapted to the data obtained 
from this study was suggested by Anderson (1949, 1954), and is the 
method used herein. Using this method several key characters are chosen 
for measurement and study, exercising care to select those characters that 
best represent the differences between the species. Frequency distributions 
of the scores of each character will indicate in graphical fashion whether 
the two snecies overlap in a particular character. More effective in demon- 
strating the pattern of variation between species is the hybrid index 
which can be computed from these data. It is designed to show variation 
of several characters simultaneously. Such an analysis has the advantage 
of sharply displaving the pattern of differences and resemblances between 
snecies, and at the same time suggests how some of the characters that 
differentiate species are inherited. Stebbins and Ferlan (1956) used this 


6 MADRONO [Vol. 15 


Fic. 1. Leaves and flowers of C. lundelliana, C. moschata, and Fy hybrid. 


1959] WHITAKER: CUCURBITA 7 


method for investigating the role of hybridization in the origin of intra- 
specific polymorphism in Ophrys, where direct genetic experiments are 
not feasible for various reasons. 


RESULTS 


DESCRIPTION OF CHARACTERS USED IN DEVELOPING THE HYBRID INDEX. 
To treat the measurement and observational data in an objective manner, 
seven characters were arbitrarily selected for analysis. A hybrid index 
was computed from data similar to those recorded in Table 1 for the 
parents. In calculating the values for the hybrid index equal weight was 
arbitrarily assigned to each character. The character expressions are 
described as follows. 

1. Leaf length / lobe depth. In C. lundelliana the leaves are deeply 
lobed (fig. 1B) as contrasted with the shallow-lobed leaves of C. moschata 
(fig. 1A). This difference is reflected in proportionately larger ratios as 
the leaf approaches the unlobed condition. The ratios for C. lundelliana 
range from 1.92 to 2.67; those for C. moschata from 5.00 to 10.60; and 
those for the F, from 3.00 to 9.00. 

2. Leaf width. The leaves of C. lundelliana are generally much nar- 
rower than those of C. moschata. Leaves of C. lundelliana range in width 
from 11.5 to 15.0 cm; those of C. moschata from 15.5 to 31.5 cm; and 
those of the F, from 13.0 to 20.5 cm. 

3. Petiole. The petioles of C. lundelliana are slender and reed-like, 
while those of C. moschata are stouter and more rigid. A measure of this 
difference is the diameter of the petiole at the point of attachment to the 
stem. Petiole diameter in C. lundelliana ranges from 0.2 to 0.4 cm; in C. 
moschata from 0.7 to 1.0 cm; in the F,; from 0.4 to 0.6 cm. 

4. Length ¢ corolla / length 2 corolla. In C. lundelliana the corollas 
of the pistillate flowers are usually somewhat longer than those of com- 
parable staminate flowers on the same plant, while in C. moschata the 
corollas of the pistillate and staminate flowers are about equal in length. 
The ratios for C. lundelliana range from 0.41 to 0.80: for C. moschata 
from 0.84 to 1.45; and for the F, hybrid from 0.57 to 1.00. 

5. Fruit length / fruit width. The fruit of C. lundelliana is short and 
ellinsoidal to almost round (fig. 2A), while that of C. moschata is long, 
slender, with an enlarged terminal portion containing the seed cavity (fig. 
2B). Fruits of the F, are phenotypically verv different from those of the 
parents and have a constricted neck portion (fig. 2C). 


Fic. 1. Leaves and flowers of C. lundelliana, C. moschata, and F1 hvbrid. A, leaf 
of C. moschata with shallow lobes and stout petiole; B, leaf of C. lundelliana with 
deep finger-like lobes and slender petiole with long hairs; C, leaf of F;, which appears 
to be about intermediate between the two species; D, staminate flower of C. moschata, 
which has short, broad corolla lobes, broad flat sepals, and a stout pedicel: E, stami- 
nate flower of C. lundelliana, with long, pointed corolla lobes, slender. thread-like 
sepals, and goblet-shaped calyx tube with slender pedicel; F, staminate flower of F1, 
which has the long, lobed corolla, slender sepals and the goblet-shaved calyx of C. 
lundelliana but has a relatively stout pedicel, « 0.48. 


ERE 


Roy 
Re 


Gee 


© 
<8 
ws 
Be 
= 3S 
5 8 
—) « 
=O 
Os 
eed ge) 
D 
= 3 
8&3 
= 
3% 
Stee 
ey 
mae 
ays 
(oer 
meat 
a - 
‘a 
o 
2A 
O «ea 
on 
Ov fx, 
Mion 
ret ACO, 
So 
oS 
aos 
mS 
AS 
uw = 
° . 
nO 
ie 
> 
Ga ie 


Fic: 2. 
lundelliana 


F, backcrossed to C. lundelliana. 


1959] WHITAKER: CUCURBITA 9 


TABLE 1. DATA USED IN COMPUTING AND CONSTRUCTING THE HYBRID INDEX FROM 
S1x CHARACTERS FOR C. LUNDELLIANA AND C. MOSCHATA* 


q Leaf length ee aioe Length ¢@ corolla Fruit length Seed F 
Species an widt iameter 7, Sanne ean area Index 
plant no. Lobe depth (cm) (cm) Length Q corolla Fruit width (sq.cm) value 


C. lundelliana 


5-1 2.56 11.5 0.4 0.71 1.15 0.5 1 
5-2 227 14.0 0.3 0.64 1.08 0.5 1 
5-3 27 13.5 0.4 0.54 1.15 0.5 0 
5-4 2.36 13.5 0.4 0.52 1.13 0.5 0 
5-5 1.92 14.5 0.4 0.41 1.07 0.5 1 
5-6 2.67 12.5 0.3 0.52 1.13 0.5 1 
5-7 2.60 13.0 0.3 0.50 1.19 0.5 0 
5-8 Zech 12.0 0.3 0.60 1.00 0.5 0) 
5-9 2.50 13.0 0.3 0.63 1.08 0.5 1 
5-10 2.50 11.5 0.2 0.75 1:07 0.5 1 
5-11 2.33 15.0 0.4 0.58 1.06 0.5 1 
5-12 2.30 13.0 OS 0.80 1.07 0.5 1 
C. moschata 
24-1 6.43 £525 0.8 1.11 4.96 2.0 13 
24-2 7.40 21.0 0.7 0.96 6.35 2.0 14 
24-3 6.75 30.5 1.0 1.09 (bale 2.0 14 
24-4 6.43 25.5 0.7 0.86 4.79 2.0 116) 
24-5 5.38 25.0 0.8 0.84 4.58 2.0 13 
24-6 6.43 24.5 0.7 0.91 S71 2.0 14 
24-7 7.67 26.0 0.8 0.88 6.83 2.0 13 
24-8 7.00 235°0 O7 1.04 5.14 2.0 14 
24-9 5.78 35 1.0 1.00 7.36 2.0 14 
24-10 5.00 25.0 0.8 1.45 6.92 220 13 
24-11 10.60 30.0 0.8 1.00 5.63 1.5 13 
24-12 10.20 28.0 0.7 1.31 5.96 2.0 14 


* The seventh character used in computing the hybrid index is not included in the table, since 
all fruits of C. lundelliana have hard rinds and solid placenta, while those of C. moschata are non- 
hard and non-solid. In computing the hybrid index all characters were arbitrarily given equal weight. 
Each individual was assigned one of three scores for each of the 7 characters; O=similar to C. lun- 
delliana; 2=similar to C. moschata; 1=intermediate. Employing this scheme, ‘‘pure’’ C. lundelli- 
ana =0, while ‘‘pure’’ C. moschata =14. 


6. Hardness of rind and type of placenta. These two qualitative charac- 
ters have been combined in the hybrid index. Fruits of C. lundelliana have 
hard, woody rinds that do not decay readily and cannot be cut with an 
ordinary knife. The seeds are embedded in the solid placenta (fig. 3A, 
right). Cucurbita moschata has a relatively soft rind that can be cut easily 
even at full maturity, while the placenta is loose and stringy and col- 
lapses at maturity leaving a large cavity (fig. 3A, left). The F, hybrid 
(fig. 3A, center) has a hard rind and a more or less open placenta. 

7. Seed area. Cucurbita lundelliana has relatively small seeds with an 
area of about 0.5 sq. cm. for each seed (fig. 3B, bottom) ; C. moschata has 
much larger seeds with an area of about 2.0 sq. cm. per seed (fig. 3B, top). 
Seeds of the F, are about intermediate in area; they range from 0.65 to 
1.35 sq. cm. (fig. 3B, center). 


10 MADRONO [Vol. 15 


pay | THU 
cul 2 8 4 S$ & 7 Z 2... 
Fic. 3. A, cross sections of fruits of C. moschata (left), F; hybrid (center), C. 


lundelliana (right); B, seeds of C. moschata (top), Fy hybrid (center), C. lundel- 
liana (bottom). 


ad 
sence 
C4 


DESCRIPTION OF THE F,. The F, plants were vigorous and thrifty and 
had foliage somewhat darker green than either parent. An occasional 
plant in the F, progenies had abnormal leaves and stems. Such plants 
died before producing flowers or maturing fruits. Of the total population 
of 28 F, plants, 7 were male-sterile. The male-sterile plants produced 
neither abortive nor male flowers. The F, plants were approximately inter- 
mediate between those of the two species in most characters (see figs. 1 
and 2). The frequency distribution of hybrid index values (fig. 4) indi- 
cates a minimum overlapping of the F, into either species in the characters 
selected for analysis. There was a wide variation in pollen fertility rang- 
ing from 17 to 74%, with a mean of 42% for the 17 plants sampled. 


1959] WHITAKER: CUCURBITA 11 


TABLE 2. FERTILITY OF PARENTS, Fy, Fe, and BACKCROSSES OF CUCURBITA 
LUNDELLIANA X C. MOSCHATA 


Total Q -Sterile a -Sterile Pollen fertility 

plants plants plants Mean Range 
Parent or progeny (number) (number) (number) (percent) (percent) 
C. lundelliana 12 0 0 96 85-99 
C. moschata 12 0 0 93 80-99 
Fy 28 0) i 42 17-74 
Fo 27 1 8 39 10-74 
Backcross to C. lundelliana 72 4 0 76 6-88 
Backcross to C. moschata 67 0 ag 48 9-94 


DESCRIPTION OF THE Fs. The bulk of the F, segregates were more 
nearly comparable with C. /undelliana than with C. moschata. In fact, for 
most characters the F. scarcely exceeded the F, in the direction of C. 
moschata in our sample of 27 F. plants. This conclusion is substantiated 
by the histogram of the hybrid index values (fig. 4). The different sizes 
and shapes of fruit produced by the segregates are shown in figure 2D. 
There were 8 male-sterile plants in the population of 27 plants from 4 
progenies. The F. segregates were characterized by a wide range in pollen 
fertility (10 to 74%; mean 39%). 

DESCRIPTION OF BACKCROSSES. The most noticeable feature of the back- 
cross progenies was their phenotypic similarity to the recurrent parent. In 
general this was true of vine, flower, and fruit characters. The photo- 
graphs of the fruits (figs. 2E and 2F) illustrate this point very nicely. The 
frequency distributions of the hybrid index values (fig. 4) suggest that 
the population from the backcross to C. moschata is more variable than 
that from the backcross to C. lundelliana. Superficially the backcrosses 
looked as if they might have been slightly more variable populations of 
either parent. 


FERTILITY OF PARENTS AND PROGENIES. For the purpose of this study it 
was important to know something about the fertility of the individuals 
comprising the F,, F,, and backcrosses. Table 2 summarizes the data ob- 
tained from field observations and pollen counts. 

The data of Table 2 indicate that about one quarter of the F; and F. 
individuals from the cross are male-sterile. Plants which did not produce 
male flowers and those which produced male flowers but aborted prior to 
anthesis were lumped in this category. It is worth noting that there was 
not a single clear-cut case of male sterility in the backcross progenies. 
One plant in the F, and 4 plants in the backcross to C. lundelliana were 
males; that is, no female flowers were produced. The pollen fertility of the 
parent species is very nearly identical (mean 93 and 96%), while the 
means of the F, and Fs are not very different (42 and 39%). On the 
other hand the mean pollen fertility for the backcross to C. lundelliana 
was 76% (range 6 to 88% ) and that for the backcross to C. moschata was 
only 48% (range 9 to 94%). This sharp difference in mean fertility be- 


12 MADRONO [Vol. 15 


F, 


C. LUNDELLIANA 


BACKCROSS TO 


C. MOSCHATA C. MOSCHATA 


» 
NUMBER OF INDIVIDUALS 


BACKCROSS TO 
C. LUNDELLIANA | 


NUMBER OF INDIVIDUALS 


01234567 8 9 Ol! 12 13 14 0'123485 67 8 9 10 II l2 13 14 
INDEX VALUE INDE VALUE 


Fic. 4. Frequency distributions of hybrid index values for Cucurbita lundelliana, 
C. moschata, their Fy, Fs, and backcross progenies to both parent species (further 
explanation in the text). 


tween the backcross progenies suggests that maternal or cytoplasmic fac- 
tors are effective in restoring to a marked degree the pollen fertility of the 
progenies of the backcross to the maternal parent. 


DISCUSSION 


The method of analysis chosen for this study has many shortcomings, 
but it does illustrate certain characters by which the two species differ, 
and gives a rough quantitative measure of these differences. 

The data suggest that some fruit characters, such as large size, soft 
rind, highly colored flesh, and large seeds, which presumably have value 
under cultivation, are for the most part recessive. On the other hand, 
vegetative and flower characters, which in this instance appear to be of 
less importance under domestication, are intermediate in their expression. 

Although the backcross progenies are remarkably similar to the recur- 
rent parent in fruit characters, the frequency distribution of the hybrid- 
index values (fig. 4) suggests that genes of the other parent are present. 
The subtle nature of this hereditary contribution cannot be detected ex- 
cept by careful study. 

The frequency distribution of the hybrid-index values (fig. 4) indicates 
that while the Fy values were considerably more variable than the Fj, as 
would normally be anticipated, they did not reach values of either of 
the parent species in the population sampled. The backcross to C. lundel- 
liana extends up to the modal class of C. lundelliana. However, its own 
mode is about halfway between the C. /undelliana mode and the F, mode. 


1959] WHITAKER: CUCURBITA 13 


The backcross to C. moschata barely reaches the mode of C. moschata, 
and again its mode is approximately halfway between those of the recur- 
rent parent and the Fy. 


SUMMARY AND CONCLUSIONS 


1. Controlled pollinations demonstrated that a wild annual species of 
Cucurbita (C. lundelliana) can exchange genes with a cultivated species 
(C. moschata). The F, is fertile enough to produce F. and backcross 
progenies. Individuals of these progenies are fertile in varying degrees 
except for some male sterility in the F; and F. generations and for one 
backcross progeny that had at least 4 individuals that were female-sterile. 

2. The parent species are well separated by differences in important 
morphological characters. This separation is illustrated by frequency dis- 
tributions constructed from hybrid-index values. From the frequency dis- 
tributions it is evident that of the 7 characters selected for analysis none 
are overlapping. The F, is intermediate between the parents in the char- 
acters selected for analysis except for the qualitative characters—rind 
hardness and placenta type. An analysis shows that in most characters, 
the F. population barely exceeds the F, in the direction of C. moschata. 
This analysis combined with a similar analysis of the backcross progenies 
suggests that C. moschata has many recessive genes. 

3. Comparisons of the backcross progenies indicate that they are strik- 
ingly similar to the recurrent parent in appearance. This finding suggests 
that various isolating barriers such as linkage, selective fertilization, and 
differential viability affect the backcross. 

4. If compatibility relations are used as criteria, the isolating barriers 
that prevent crossing between C. lundelliana and C. moschata are not 
well developed in spite of the great morphological diversity between the 
species. There seems no doubt that each species is a good taxonomic entity. 
It is equally clear that the two species have a number of genes in common. 
For this reason it is not unreasonable to suggest that C. moschata may 
have been derived from C. lundelliana by isolation and subsequent selec- 
tion by man. An alternative suggestion would be that both species were 
derived from a common ancestor, but when domesticated, C. moschata 


diverged sharply by natural selection under the guidance of man. 
U.S. Deparment of Agriculture 
Agricultural Research Service 
Crops Research Division 
La Jolla, California 


LITERATURE CITED 


ANDERSON, E. 1949. Introgressive hybridization. Biol. Reviews 28:280-307. 

. 1954. Efficient and inefficient methods of measuring specific differences. 
Statistics and Mathematics in Biology. pp. 93-106. Iowa State College Press. 
1954. 

STEBBINS, G. LEDYARD AND L. FERLAN. 1956. Population variability, hybridization, 
and introgression in some species of Ophrys. Evolution 10:32—46. 

WHITAKER, THomMas W. 1956. The origin of the cultivated Cucurbita, Amer. Nat, 
90:171-176. 


14 MADRONO [Vol. 15 


THE REPRODUCTIVE STRUCTURES OF SCHINUS MOLLE 
(ANACARDIACEAE) 


HERBERT F. COPELAND 


The Vergleichende Embryologie der Angiospermen of Schnarf (1931) 
has served as the basis and stimulus for very entensive subsequent work 
in its field. Schnarf accounted for the Anacardiaceae on the basis essen- 
tially of a single paper dealing with a single species, being the account of 
Rhus Toxicodendron L. (or Toxicodendron radicans O. Kuntze) by 
Grimm (1912). Subsequent studies of floral morphology and embryology 
in this family include the following: Juliano (1932) on Spondias pur- 
purea L.; Juliano and Cuevas (1932) on Mangifera indica L.; Mahesh- 
wari (1934) on Mangifera indica; Srinivaschar (1940) on Semecarpus 
Anacardium L., Spondias Mangifera Willd., and Anacardium occidentale 
L.; Copeland and Doyel! (1940) on Toxicodendron diversilobum Greene; 
Sharma (1954) on Mangifera indica; Copeland? (1955) on Pistacia 
chinensis Bunge; Kelkar (1958a) on Rhus mysurensis Heyne; and Kel- 
kar (1958b) on Lannea coromandelica (Houtt.) Merrill. The present 
paper adds Schinus Molle L. to the list of species of Anacardiaceae in 
which the morphology of the reproductive structures is reasonably well 
known. 

Material was collected in the public parks of the City of Sacramento 
and prepared by routine microtechnical methods (a special technique ap- 
plied to pollen grains is described below). The results fall short of desired 
completeness in failing to account for the growth of the pollen tube and 
for fertilization. 


THE TREE 


Braunton and Davy (1914) tell us that Schinus Molle is a native of 
Peru; that its common name is Peruvian mastic; and that it is very 
familiar as a cultivated ornamental in southern California, where it has 
the common name of pepper tree. It is in fact familiar also in northern 
California, and in many other moderately warm countries. 

The tree is fast-growing; of deliquescent form; rarely much more than 
10 m. tall; evergreen, with alternate pinnate leaves, the branchlets and 
leaves often pendant; the small yellow flowers abundant in terminal 
clusters; dioecious; the female trees producing bunches of purplish-red 


1 Bernice Elva Doyel, afterward Mrs. Harold Strimling, my pupil, associate, and 
friend, died in Sacramento on February 24, 1954, at the age of 34 years. 

2TIt behooves me to acknowledge, and to call to the attention of readers, a ludi- 
crous error in the paper cited. The median stigma of Pistacia is said to stand above 
the large end of the ovary, in which the ovule is attached. Actually it stands above 
the small end. Referring to a passage in this paper in the second column of page 442, 
figures 17, 18, 19, 20, and 21 were drawn with the median stigma to the right; my 
other figures of the gynoecium and its parts were drawn with the median stigma to 
the left. 


1959] COPELAND: SCHINUS MOLLE 15 


Ela 
SS 


gpNOUGOL 


SS 
& 


Fics. 1-15. Schinus Molle: 1, fruiting twig, x 0.4; 2, cluster of flower buds, x 8; 
3, male flower, X 8; 4, female flower, x 8; 5, pubescence of pedicel, & 320; 6, model 
of the vascular system of a female flower, X 40; 7, cross section of young anther, 
xX 320; 8, cross section of one lobe of an older anther, x 320; 9, 10, heterotypic 
metaphase in pollen mother cell, xX 720; 11, microspore tetrad, x 720; 12, cross 
section of wall of developing anther, x 320; 13, cross section of mature anther, x 40; 
14, portion of cross section of mature anther, X 320; 15, pollen grain, x 720. 


drupes about 5 mm. in diameter (fig. 1). It was the odor and taste of 
these fruits that suggested the common name of pepper tree. 

The pepper tree has not the typical reproductive cycle of plants of 
temperate regions. It produces flowers continually from spring to fall. It 


16 MADRONO [Vol. 15 


responds to the approach of winter by dropping its flower buds, flowers, 
and young fruits, and by ceasing to grow until the following spring. 

In Sacramento, some of the minute axillary buds give rise to branches 
at about the beginning of the month of March. Each branch bears first 
two minute alternate scales, these being the covering of the bud. The axis 
of the branch elongates during a period of from six to eight weeks, pro- 
ducing from three to seven or more foliage leaves. The terminal bud then 
produces an inflorescence, or else dies: terminal buds do not normally 
serve as winter buds. Flowers appear first on a particular tree from the 
middle of May to the middle of June. They are evidently entomophilous, 
and honey bees have been seen to visit them. Fruits are mature about two 
months after flowering: the first ones of the year are found at about the 
beginning of August. 

The axis of a flower cluster, continuing that of a branch, grows in in- 
determinate fashion, bearing a series of alternate scales, but eventually 
producing a terminal flower. From the axil of each scale grows an axis 
bearing, ordinarily, two scales and a terminal flower; from the axils of 
these scales, in basipetal succession, further growth takes place, usually 
of a single flower, sometimes of a further cluster (fig. 2). The entire 
cluster of flowers as described is identifiable as a thyrse. 

The plant as a whole appears glabrous, but there is a scant pubescence 
of glandular and simple hairs on the axes and scales of the inflorescence 
(fig. 5). 

FLOWERS 


The male flowers (fig. 3) are slightly larger, and have more erect 
petals, than the female (fig. 4). Each flower has five sepals, five petals, 
ten stamens, and a pistil. A prominent disk surrounds the base of the 
ovary inside the bases of the stamens. A small number of stomata are 
present in its epidermis. The male flowers have minute and incompletely 
developed pistils; the female flower bear smaller stamens than the male. 
The sporogenous cells in the anthers of female flowers undergo abortion 
at the stage of pollen mother cells. 

The mature pistil of the female flower is dorsiventral, but not con- 
spicuously so: in the microtechnical operation of imbedding pistils with 
the intention of cutting sagittal sections, one makes a considerable num- 
ber of mistakes. The ovary is of a shape approaching that of an egg laid 
upon its side: it has a larger and a smaller end; a brief style with a ter- 
minal stigma stands in a median position toward the smaller end of the 
ovary; two lateral styles with their stigmas, not particularly different 
from the median one, stand above the larger end. As a fairly frequent ab- 
normality, pistils occur with only two styles, one above each end. The 
ovary contains a single locule almost entirely filled by the single ovule. 
This is attached to the ovary wall in the median plane of the ovary, at the 
large end, above the middle (fig. 25). 

In the phloem of every vascular bundle, but not extending to the ex- 


1959] COPELAND: SCHINUS MOLLE 17 


PHRN 
PANS 

LANs 

Bt glyN 

-Z, 7ho Weak 

6 EO ANN 
ide Al 


Fics. 16-31. Schinus Molle: 16, longitudinal section of ovary in young female 
bud, X 40; 17, beginning of development of ovule, X 320; 18, beginning of periclinal 
division in the nucellar hypodermis, X 320; 19, longitudinal section of female flower 
bud, xX 40; 20, megaspore mother cell, x 320; 21, tetrad of megaspores, x 320; 
22, 2-nucleate embryo sac, X 320; 23, 4-nucleate embryo sac, & 320; 24, an abnor- 
mality, an embryo sac developing from the spore next to the chalazal spore, 320; 
25, longitudinal section of female flower with a mature embryo sac, X 40; 26, inner 
epidermis of ovary of open flower, * 320; 27, mature embryo sac, X 320; 28, un- 
divided zygote in plurinucleate endosperm, x 320; 29, 30, young embryos, x 320; 
31, longitudinal section of fruit about one month after anthesis, « 8. h, hypostase; 
it, tubular outgrowth of outer integuments; ex, exocarp; mc, mesocarp; end, endo- 
carp. 


18 MADRONO [Vol. 15 


treme end of the bundle, there is a resin duct, conspicuous under the 
microscope, being of greater bulk than the vascular tissues. The resin 
ducts enable one easily to follow the course of the bundles. 

About five bundles, arranged in a cylinder, ascend the pedicel and 
enter the receptacle. In the base of the receptacle they fuse and fork to 
a certain extent, and send out in radial directions branches which undergo 
further forking. These are the sepal supply; about five bundles enter each 
sepal. Above their departure, the stele sends out three successive alternat- 
ing whorls of five bundles, supplying respectively the petals, the sepalad 
stamens, and the petalad stamens. In male flowers, the vascular tissue 
ascending beyond the departure of the stamen traces is scant and quickly 
fades out. In female flowers (fig. 6), much vascular tissue ascends beyond 
this point. It is of the form of a truncate cone splitting at the summit to 
form a whorl of bundles whose typical number is eight. One ascends the 
small end of the ovary; two ascend each side; at the large end there are 
typically two ovary wall bundles, and between them, or, often, fused to 
one of them, the bundle which supplies the ovule. At the summit of the 
ovary, the ovary wall bundles meet and form a scant network. From this 
network, branches go up into the styles, each style receiving a median 
bundle and two laterals. 


MALE STRUCTURES 


The stamens are in all respects of the structure typical of flowering 
plants. Within the four angles of the rudimentary anther (fig. 7), peri- 
clinal divisions occur in the hypodermal cells. The progeny of these cells, 
produced by duly numerous divisions, become differentiated into the 
following layers of the anther, in order from outward to inward: (a) the 
endothecium, the hypodermal layer of cells with ribbed walls, whose con- 
traction opens the anther through two longitudinal clefts (figs. 13, 14); 
(b) two layers of wall cells, of which the outer becomes compressed, and 
the inner crushed and absorbed (figs. 12, 14); (c) a tapetum of the secre- 
tion type. the cells becoming binucleate and then shrivelling and disap- 
pearing (fig. 12); and (d) the spore mother cells. In the male flower, 
meiosis duly takes place within these last. The haploid chromosome 
number, observed during this process, is 15 (figs. 9, 10). The pollen grains 
are separated by simultaneous furrowing. The mature pollen grain (fig. 
15) has a wall marked by numerous fine pits and three meridional grooves. 

The pollen grains contain much stainable material, and it has been 
difficult to be certain of the number of nuclei. A technique by which they 
were made visible was as follows: microtome sections, mounted, deparaf- 
fined. and hydrated, were exposed for 20 minutes to 1% NasCOs to 
which a little NaOH had been added, at 60°C.; stained briefly with 
methvlene blue; and destained in 95% alcohol. Two nuclei are present. 
The tube nucleus is the larger. It is subglobular and contains a visible 
nucleolus. The generative nucleus is smaller, fusiform, and heavily stain- 
ing. It lies within a distinct space, the generative cell 


1959] COPELAND: SCHINUS MOLLE 19 


THE OVULE 


The parts of the flower originate in the bud in acropetal succession. The 
pistil, in the female bud, originates as a cycle of three knobs, evidently 
rudimentary carpels, which, as they grow, become coalescent at the base 
(fig. 16). Somewhat above the bottom of the cavity in the resulting three- 
pointed cone, the nucellus begins to grow into it from an area below one 
of the notches between the points (fig. 17). By further growth, the ovary 
becomes closed above the locule containing the developing ovule (fig. 19). 
The points above develop, of course, into the styles and stigmas. 

The direction of the axis of the nucellus is at first horizontal or slightly 
upward. It soon bends and grows diagonally downward. At about this 
time, the inner integument originates as a collar of tissue surrounding the 
tip of the nucellus and growing forward with it. The hypodermal cells of 
a small area near the tip of the nucellus begin to undergo periclinal divi- 
sions. One of these cells, near the middle of the area, cuts off to the inside 
a cell, slightly larger than the others, which is to become the megaspore 
mother cell (fig. 18). By further periclinal divisions, the megaspore 
mother cell is carried well to the interior of the nucellus (fig. 20); by 
growth in other parts, an outer integument is produced; both integuments 
extend beyond the nucellus and close over it, leaving a small micropyle. 
At the same time that the growth of the ovule produces these changes, it 
continues to have a bending or coiling effect: the mature ovule is so bent 
that its axis points to the ovary wall below its insertion (fig. 25). 

Meanwhile, the spore mother cell has produced a tetrad of spores (fig. 
21), among which the chalazal spore is functional. The nucleus of the 
functional spore undergoes three divisions (figs. 22, 23). Six of the re- 
sulting eight nuclei are set apart in an egg, two synergids, and three anti- 
podal cells, leaving two polar nuclei in the endosperm mother cell (fig. 27). 
Thus the embryo sac is of the type which Schnarf designated as normal. 

As the embryo sac approaches maturity, there appears in the chalaza, 
between the end of the bundle traversing the raphe and the antinodal cells, 
a hypostase (figs. 25, 32, 35), being a body of thick-walled cells of mori- 
bund appearance. 

A few examples of abnormal development have been noted. An oc- 
casional ovule is distorted, twisted otherwise than in the normal down- 
wardly coiled fashion. Sometimes a megaspore other than the chalazal 
one undergoes development (fig. 24). 


DEVELOPMENT OF FRUIT AND SEED 


The ovary grows during about four weeks after anthesis approximately 
to the full size of the fruit (fig. 31). 

At anthesis, the cells of the inner epidermis of the ovary show periclinal 
divisions (fig. 26). By these divisions, the epidermis gives rise to three 
regular layers of cells. The cells of the innermost layer become radially 
elongate to a length of about 0.2 mm., after which they develop thick 


20 MADRONO [Vol. 15 


a tsmny-f--- 
~ 


Fics. 32-39. Schinus Molle: 32, longitudinal section of ovule about one month 
after anthesis, & 40; 33, 34, developing embryos, x 320; 35, longitudinal section 
of nearly mature fruit, x 8; 36, section of endocarp, area x of fig. 35, XX 320; 
37, mature seed, X 8; 38, mature embryo, X 8; 39, cross section of mature fruit, 
x 8. h, hypostase; it, tubular outgrowth of outer integument; ex, exocarp; mc, meso- 
carp; end, endocarp. 


walls. The middle layer becomes a palisade of thin-walled cells, very 
much smaller than the preceding. The outer becomes a palisade of thick- 
walled cells, of about the diameter of those of the inner layer, but only 
about 0.05 mm. long (fig. 36). These three layers, all derived from the 


1959] COPELAND: SCHINUS MOLLE 21 


inner epidermis of the ovary, make up the stony endocarp of the fruit. 
Until about a month after anthesis, it is possible to cut on the microtome 
paraffin sections of the endocarp; later, the thickness and hardness of the 
walls make this impossible. The endocarp is deeply impressed by the meri- 
dional bundles, with their large resin ducts, in the ovary and fruit. As 
compared with the more or less egg-shaped ovary and fruit, the endocarp 
is more strongly compressed in the lateral dimension, being approximately 
lenticular. 

During the first month after anthesis, the ovule grows principally in its 
micropylar-chalazal dimension. The shape of its axis becomes that of a 
horizontal U: the end of the nucellus points to an area directly below the 
insertion of the ovule. The outer integument grows forth as a prominent 
tube, directed downward in the large end of the ovary at right angles to 
the rest of the ovule. This structure is presumably without function (figs. 
31, 32). It persists in a shrivelled condition in the mature fruit (figs. 35, 
37). 

The endosperm is of nuclear type. It contains a considerable number of 
free nuclei before the zygote divides (fig. 28). The nuclei remain free when 
the embryo is 4-celled (fig. 29); when the embryo is of about a dozen 
cells (at the stage shown in fig. 30), cellular endosperm has begun to 
form about it. Further uninucleate cells are cut out, first throughout the 
surface of the endosperm, and then throughout its interior. 

The first division of the zygote has not been seen. It occurs about three 
weeks after anthesis, and is believed usually to be by an oblique wall. The 
epibasal cell divides, by walls which are usually oblique, two or three 
more times. Longitudinal divisions then begin, first in the more distal cells, 
then in the more proximal, but not usually in the basal cell. Thus when a 
globular embryo is formed (figs. 33, 34), it has a suspensor which is brief 
and not sharply set apart, consisting of a basal cell which is not enlarged 
and of one or two pairs of cells beyond this. 

After the fruit and its locule have reached nearly their full size, the 
seed and embryo enter their phase of most rapid enlargement. The seed 
fills the locule laterally; its end toward the large end of the fruit becomes 
vertically elongate; it bulges greatly on the side away from the raphe. The 
impressions in the endocarp, produced by resin ducts, produce impres- 
sions on the sides of the seed. 

The vertical elongation of the end of the seed toward the large end of 
the fruit is accompanied by elongation of the hypocotyl and radicle as a 
stout column in this end of the seed. The root tip points upward, and re- 
mains for a long time near the micropyle, but eventually grows past and 
curves away from it (fig. 38). 

The cotyledons grow forth at right angles to the direction of the hypo- 
cotyl. They lie in planes parallel to the median plane of the fruit and 
seed. The raphe increases only moderately in length; the seed coat and 
endosperm grow forth beyond its end, and the sharp points of the cotyle- 
dons extend beyond the hypostase into the pocket thus formed. 


22 MADRONO [Vol. 15 


The plumule consists of two small knobs placed decussately with re- 
spect to the cotyledons. 

After the fruit and seed have reached apparent structural maturity, 
the mesocarp, containing the bundles and resin ducts, becomes shrivelled. 
A pigmented filmy exocarp, consisting of the external epidermis of the 
fruit together with one hypodermal layer, encloses a vacant space. Within 
this, the mesocarp, of gummy consistency, forms a layer on the surface of 
the stone. A considerable body of endosperm survives in the mature seed 
(fig. 39). 

DISCUSSION 


Here one attempts to characterize the Anacardiaceae by their reproduc- 
tive structures, microscopic as well as macroscopic. Since the microscopic 
structures are definitely known only of a moderate number of species, 
most of which belong to the single tribe Rhoideae, the characterization is 
acknowledgedly tentative. 

The Anacardiaceae bear flowers in cymes which are usually gathered 
into thyrses. They produce imperfect flowers, some species being dioecious, 
others polygamous. The imperfect flowers usually appear complete 
through the presence of rudimentary organs. The flowers are pentamer- 
ous, except that in most species the gynoecium is of fewer than five carpels. 
The ovary or ovaries are superior, surrounded at the base by a prominent 
disk. (To several of these statements, Pistacia offers exceptions, evidently 
through loss of parts.) In the tribe Rhoideae, the pistil is compound, con- 
sisting of one fertile and two sterile carpels, with separate styles and 
stigmas. There is a single locule. A single ovule is attached basally or else 
to the wall of the locule, on the side away from the mid-plane of the fertile 
carpel. 

Except in the gynoecium, the receptacular vascular system is undis- 
tinguished, a matter of the stele emitting alternating whorls of bundles. 
The sepalad whorl of stamens is the lower. With the reduction of some of 
the carpels is associated the fact that the ovary wall bundles are not 
definitely identifiable as carpel dorsals, carpel laterals, and so forth. When 
the ovule is basifixed, the bundle supplying it is formed by the anasto- 
mosis of several or many bundles running together from all sides. This 
evidently derived character has been noted outside of tribe Rhoideae in 
Mangifera (Sharma, 1954). 

Stamens and pollen show no characters distinguishing Anacardiaceae 
from flowering plants in general. The four pollen sacs open, by the action 
of a ribbed endothecium, through two slits. The tapetum is of the secretion 
type: the cells become and remain binucleate, and remain separate until 
they are absorbed. The most frequently observed haploid chromosome 
number is 15; Srinivasachar found 14 in Spondias. The pollen grains are 
separated by simultaneous furrowing. They are binucleate. The exine is 
in many examples marked by numerous fine pits and by three meridional 
grooves, and contain two nucle}. 


1959] COPELAND: SCHINUS MOLLE ws: 


Ovule and seed grow in such fashion as to become more or less strongly 
apotropously coiled in the median plane of the pistil. Basically, the ovule 
is bitegmous, but various genera exhibit peculiarities in the development 
of the integuments, particularly the outer. One suggests for future study 
the possibility that the obturator, the outgrowth at the base of the funi- 
culus which occurs in various genera but not in Schinus, may be morpho- 
logically a part of the outer integument. 

The ovule is crassinucellate. In the area of the tip of the nucellus, there 
is a hypodermal archesporium, a layer of cells which undergo periclinal 
divisions. Only one cell of this layer is functionally archesporial: only one 
megaspore mother cell is produced (these points, not established in our 
paper on Toxicodendron diversiloba, have been observed in subsequently 
prepared slides of this species). In the chalaza there is present a hypostase, 
a body of thick-walled cells of moribund appearance. 

The embryo sac is of the type which Schnarf designated as normal. 
Current usage designates this the Polygonum type, a term which fails to 
express what we believe to be its significance. 

The peculiar types of development in the integuments are believed to 
be associated with the occurence of aporogamy, which is definitely known 
in Toxicodendron and Pistacia. Double fertilization is presumed to occur, 
but has been observed only in Spondias (Srinivasachar, 1940). 

The fruit grows nearly to its full size before the ovule grows consider- 
ably, and the ovule grows considerably before the embryo does so. The 
occurrence of these three distinguishable cycles of growth was noted in 
Mangifera by Kennard (1955). 

The fruit is a drupe. In tribe Rhoideae, the endocarp is derived entirely 
from the inner epidermis of the ovary: it appears regularly to consist of 
three definite layers of cells; its stony character is produced by the ma- 
turation of one or more of these layers as dense palisades of fiber-like 
cells. These things appear not to be true of Mangifera and Spondias. 

The endosperm is of nuclear type. It becomes cellular first about the 
embryo, then throughout its periphery, then in all parts. 

The first divisions of the zygote are more or less oblique. A brief sus- 
pensor, not definitely set apart, or none, is produced. 


Enlargement of the ovule in the course of becoming a seed involves 
considerable bulging on the side describable as the lower, away from the 
raphe. The raphe does not elongate considerably, but the enlarging seed 
coat and endosperm push forth beyond the hypostase which marks the 
end of the raphe. As the growing seed bulges downward, elongation of 
the hypocotyl-radicle pushes its proximal end downward, while the root 
tip remains in the neighborhood of the micropyle. The cotyledons lie in 
planes parallel to the mid-plane of the gynoecium. They grow forth ap- 
proximately at right angles to the hypocotyl, and their tips extend beyond 
the hypostase. A certain amount of endosperm survives in the mature 
seed. 


24 MADRONO [Vol. 15 


We will be able to recognize other groups of plants as immediate allies 
of Anacardiaceae whenever we discover in them assemblages of characters 
showing a family resemblance to the foregoing. Observations of the same 
nature as these upon further Anacardiaceae should lead to improvements 
in the system of the family. In the present state of knowledge, it appears 
that Schinus is relatively primitive, and Pistacia relatively advanced. 


SUMMARY 


Schinus Molle, a South American tree popular as an ornamental in 
California, where it is called the pepper tree, is dioecious. Flowers of both 
sexes, produced throughout the summer in thyrses, appear complete, but 
in each the organs of opposite sex are imperfectly developed. 

The male parts exhibit no notable peculiarities: endothecium ribbed; 
tapetum of secretion type, the cells becoming binucleate; haploid chrom- 
osome number 15; pollen grains 3-grooved, binucleate. 

The pistil is compound, of one fertile and two sterile carpels, with sep- 
arate styles and stigmas. The single locule contains a single ovule, at- 
tached to the wall on the side opposite the style of the fertile carpel. It is 
apotropous, bitegmous, crassinucellate. A hypodermal layer of archespo- 
rial cells in the distal end of the nucellus produces a single megaspore 
mother cell. The embryo sac is of normal type. 

After anthesis, the ovary grows in about four weeks nearly to the size 
of the mature fruit; ripening requires another month. The fruit is a drupe 
whose exocarp, of only two layers of cells, encloses, in an empty space, a 
gummy mesocarp about a stony endocarp. The endocarp is derived en- 
tirely from the inner epidermis of the ovary, and consists of three layers 
of radially elongate cells, two of them with greatly thickened walls. 

Considerable enlargement of the ovule is delayed until the fruit is nearly 
of its full size, and the embryo grows but little until the ovule is in full 
course of enlargement. The enlarging ovule is marked by a peculiar tube- 
like extension of the outer integument. The endosperm is nuclear. The first 
divisions of the zygote are oblique. The suspensor is brief and not sharply 
set apart from the embryo proper. In the mature seed the hypocotyl] points 
vertically upward in the end of the seed toward the larger end of the fruit; 
the cotyledons project at right angles from its lower end; a certain amount 
of endosperm remains present. 

Most of these characters are found, by comparison with those of vari- 
ous other Anacardiaceae, to be characters of family Anacardiaceae, or 
at least of tribe Rhoideae. Groups other than Anacardiaceae will confi- 
dently be recognized as related to this family when sets of characters show- 
ing a family similarity to these are observed in them. 


Sacramento Junior College, 
Sacramento, California 


1959] FLINT: LIQUIDAMBAR STYRACIFLUA 25 


LITERATURE CITED 


Braunton, E., and J. B. Davy. 1914. Schinus. In Bailey, L.H., Standard Cyclopedia 
of Horticulture 6:3108-3109. 

CopELanD, H. F., and B. E. Dover. 1940. Some features of the structure of Toxico- 
dendron diversiloba. Am. Jour. Bot. 27:932—939. 

CopELAND, H. F. 1955. The reproductive structures of Pistacia chinensis (Anacardia- 
ceae). Phytomorphology 5:440-449. 

GrimM, J. 1912. Entwicklungsgeschichtliche Untersuchungen an Rhus und Coriaria. 
Flora 104:309-334. 

Juriano, J. B. 1932. The cause of sterility in Spondias purpurea Linn. Philippine 
Agriculturist 21:15-24. 

Juxiano, J. M., and N. L. Cuevas. 1932. Floral morphology of the mango (Mangifera 
indica L.) with special reference to the Pico variety from the Philippines. Philip- 
pine Agriculturist 26:443-472. 

Kerxkar, S.S. 1958a. Embryology of Rhus mysurensis Heyne. Jour. Indian Bot. Soc. 
37:114-122. 

KeLkar, S. S. 1958b. A contribution to the embryology of Lannea coromandelica 
(Houtt.). Merr. Jour. Univ. Bombay 26:152-159. 

KENNARD, W. C. 1955. Development of the fruit, seed, and embryo of the Paheri 
mango. Bot. Gaz. 117:28-32. 

MauesuHwakrlI, P. 1934. The Indian mango. Current Sci. 3:97—98. 

ScunarF, K. 1931. Vergleichende Embryologie der Angiospermen. Berlin. 

SHarmMa, M. R. 1954. Studies in the family Anacardiaceae. I. Vascular anatomy of 
the flower of Mangifera indica L. Phytomorphology 4:201-208. 

SRINIVASACHAR, D. 1940. Morphological studies in the family Anacardiaceae. Jour. 
Mysore Univ. n.s. B 1:83-91. 


DEVELOPMENT OF THE MEGAGAMETOPHYTE 
IN LIQUIDAMBAR STYRACIFLDUA L. 


FRANKLIN F. FLINT 


No work has been published on the megagametophyte of Liquidambar. 
Within the family (Hamamelidaceae) only two other genera have been 
studied in regard to megasporogenesis and megagametogenesis (Flint, 
1957, 1957A). 

Pistillate floral heads of Liquidambar styraciflua were collected in Din- 
widdie County, Virginia, from March 28 until July 10, 1953. They were 
fixed in formalin-aceto-alcohol in the proportions of 90 cc. of 70 percent 
ethyl alcohol to 5 cc. formalin and 5 cc. of glacial acetic acid. Tertiary 
butyl alcohol, as used by Johansen (1940), was found most satisfactory 
for dehydration. The heads were then embedded in tissuemat, sectioned 
at 10-30 microns, and stained with Harris’ Hemotaxylin, Safranin O, and 
Fast Green FCF. 

Around 245 ovules are included in this study. Ovules which showed 
signs of unusual plasmolysis, nuclear degeneration, or had unusually dark 
staining cytoplasm were eliminated and are not included in the above 
count. 


26 MADRONO [Vol. 15 


Fic. 1. Young ovule with one megaspore mother cell and one other enlarged 
nucellar cell. Fic. 2. Young ovule with five potential mother cells. Only one will form 
megaspores. 


MEGASPOROGENESIS 


The ovules are formed in two rows within each locule and develop from 
its base toward the apex. As many as twelve to eighteen ovules are initiated 
in each locule although only two to six usually form gametophytes and 
these are most often located near the base of the locule. The ovules may 
cease development and become degenerate during any stage of megasporo- 
genesis or megagametogenesis. 

The megaspore mother cell is first distinguishable within the young 
ovule, embedded beneath seven to nine layers of nucellar cells. This cell 
is large, nearly oval in shape, with dense cytoplasm, and a large vesicular 
nucleus (fig. 1). Frequently as many as five large cells are present within 
the nucellus and have the appearance of megaspore mother cells (fig. 2). 
Only one of these cells in an ovule undergoes the meiotic divisions, and it 
is then that the functional megaspore mother cell can first be determined. 
In such ovules the functional mother cell is most often located near the 
center of the group of enlarged cells. After meiosis I is completed, cyto- 
kinesis takes place, and a dyad of cells of approximately the same size, 
each with dense cytoplasm and a large, deeply staining nucleus containing 
a prominent nucleolus, is formed (fig. 3). The chalazal dyad cell com- 
pletes meiosis II and cytokinesis takes place, but the micropylar dyad 


Fics. 3-5. Megasporogenesis. 3, dyad cells formed from meiosis I of megaspore 
mother cell; 4, micropylar dyad cell disintegrating while chalazal dyad cell undergoes 
meiosis II and forms two megaspores; 5, enlarging chalazal megaspore with upper 
megaspore and micropylar dyad cell disintegrating. 

Fics. 6-16. Megagametogenesis. 6, young 2-nucleate megagametophyte; 7, begin- 
ning of vacuoles in 2-nucleate megagametophyte; 8, vacuoles have coalesced in center 


1959] FLINT: LIQUIDAMBAR STYRACIFLUA 


of 2-nucleate megagametophyte; 9, 4-nucleate megagametophyte; 10, nuclear divi- 
sions to form 8-nucleate megagametophyte; 11, egg apparatus formed, also antipodals, 
but before migration of polar nuclei; 12, polar nuclei have migrated to center of 
megagametophyte; 13, polar nuclei have fused and antipodals are disintegrating ; 
14, fusion of two polar nuclei; 15, fusion of two polar nuclei and sperm nucleus; 
16, enlarged egg apparatus of mature megagametophyte. 


28 MADRONO [Vol. 15 


cell degenerates without undergoing this division (fig. 4). The linear 
triad of cells which results from this activity consists of the degenerating 
micropylar dyad cell and the two megaspores formed by the chalazal dyad 
cell (fig. 4). In all cases observed the chalazal megaspore enlarged further 
and became the only functional megaspore (fig. 5). The other two cells 
of the triad are soon pushed aside by the developing megagametophyte 
which presses them against the surrounding nucellar cells where they 
gradually disintegrate (figs. 5, 6, 7, 8, 9). 


MEGAGAMETOGENESIS 


In the ovules studied the chalazal megaspore increases in size, the 
nucleus divides, and gradually a 2-nucleate megagametophyte is formed 
(fig. 6). As the cell continues to enlarge, two small lateral vacuoles appear 
in the cytoplasm, one to either side of the nuclei (fig. 7). Growth of the 
megagametophyte continues and the two lateral vacuoles coalesce into a 
single large central vacuole as one of the nuclei and part of the cytoplasm 
move toward the micropylar end and the other nucleus and remainder of 
the cytoplasm move toward the chalazal end of the cell (fig. 8). Rapid 
growth of the megagametophyte continues and each of the nuclei under- 
goes another division forming a 4-nucleate cell with a pair of nuclei at 
either end oriented along the major axis of the megagametophyte (fig. 9). 
The four nuclei undergo a division with one spindle at each end of the 
megagametophyte oriented along the major axis of the cell and one spindle 
at each end oriented perpendicular to the major axis (fig. 10). This divi- 
sion results in an 8-nucleate megagametophyte with four nuclei located at 
either end. The megagametophyte elongates further and the egg ap- 
paratus, consisting of three cells, is soon formed at the micropylar end 
of the megagametophyte (fig. 11). There is no discernible differentiation 
into an egg cell and two synergids at this time. Three distinct, compara- 
tively large, antipodal cells are formed at the chalazal end of the mega- 
gametonhyte (fig. 11). The two polar nuclei, one located beneath the egg 
annaratus and the other directly above the antipodal cells, are immersed 
in thin cytoplasm and separated by a central vacuole. The polar nuclei 
soon migrate toward the center of the megagametophyte and lie next to 
each other (fig. 12). The antipodal cells begin to degenerate (fig. 12) and 
then separate from each other and the primary endosperm cell (fig. 13). 
At the same time the micropylar end, containing the egg apparatus, in- 
creases in size (fig. 13). The egg cell is soon distinguished from the two 
synergids by its larger nucleus (fig. 16). The polar nuclei may fuse to 
form a secondary nucleus (fig. 14) or fusion may be delayed until the 
sperm nucleus enters, in which case all three nuclei fuse at once to form 
the primary endosperm nucleus (fig. 15). With the disintegration of the 
antipodal cells the megagametophyte becomes broader, the single second- 
ary nucleus (or the primary endosperm nucleus, as the case may be) and 
the egg apparatus enlarge (fig. 16). The cytoplasm of the two synergid 


1959] REVIEWS 29 


cells becomes vacuolated and a small vacuole often appears above the egg 
nucleus within the greatly enlarged egg cell. 


SUMMARY 


The megaspore mother cell is embedded beneath seven to nine nucellar 
cell layers. There are frequently more than one and sometimes as many 
as five large cells which resemble megaspore mother cells in a single ovule, 
but only one of these cells functions. A linear triad of cells is formed as 
the chalazal dyad cell undergoes meiosis II and the micropylar dyad cell 
does not. In previously studied species of the Hamamelidaceae only one 
potential megaspore mother cell forms in each ovule and both dyad cells 
undergo meiosis IT to form a linear tetrad of megaspores. The chalazal 
megaspore develops into the megagametophyte in all species studied. The 
pattern of development is essentially that of the Polygonum type listed 
by Maheshwari (1950). 


Department of Biology 
Randolph-Macon Woman’s College 
Lynchburg, Virginia 


LITERATURE CITED 


Fuint, F. F. 1957. Megasporogenesis and megagametogenesis in Hamamelis virgini- 
ana L. Va. Jour. Sci. 8(3) :185-189. 
. 1957A. Megasporogenesis and megagametogenesis in Fothergilla gardeni 
Murr. and Fothergilla major Lodd. Trans. Am. Micro. Soc. LX XVI(3) :307-311. 
JoHaNnsEN, D.A. 1940. Plant microtechnique. McGraw-Hill, N.Y. 
Manesuwar], P. 1950. An introduction to the embryology of the angiosperms. Mc- 
Graw-Hill, N.Y. 


REVIEWS 


A Flora of San Francisco, California. By JOHN THomMas HOWELL, PETER H. Raven, 
AND PETER RuBTZOFF. 157 pp., 20 figs., 1 map. The Wasmann Journal of Biology, 
Vol. 16, No. 1. 1958. Published in book form by the University of San Francisco. 
Paper. $3.00. 

In spite of the many people now living within the city limits of San Francisco, an 
amazing number of native plants are still to be found. This is largely due to the hilli- 
ness of the city. The hilltops, bluffs, and cliffs provide refuges, many of which have 
not as yet been commercially exploited. Some habitats, notably the marshes and 
sand dunes, are rapidly disappearing. On Bernal Heights, however, Dodecatheon 
patulum var. bernalinum still grows at its type locality and Aristolochia californica 
is still to be found in ravines near Lake Merced. 

In 1892, Katharine Brandegee listed 605 taxa of vascular plants in her “Catalogue 
of the Flowering Plants and Ferns Growing Spontaneously in the City of San Fran- 
cisco.” Howell, Raven, and Rubtzoff list 1132 species, subspecies, varieties, forms, 
and hybrids as growing spontaneously in San Francisco now or in the past. Of these, 
about 42 per cent are introduced plants and about 19 per cent have not been collected 
since 1940 and are probably now extinct locally. A number of garden escapes are 
included. Among them are: Pseudosasa japonica, Pittosporum crassifolium, 2 species 
of Cotoneaster, 3 species of Acacia, Albizzia lophantha, Cassia tomentosa, Ruta chale- 
pensis, and Buddleia davidii. The main criterion used for inclusion of a particular 
garden escape is “whether or not the plant could survive in San Francisco without 


30 MADRONO [Vol. 15 


the aid of summer irrigation.” Ample attention also has been paid to weeds. The 
senior author in particular has collected in disturbed areas in San Francisco, along 
railroad tracks and in the vicinity of stockyards, and has found such weeds as Kochia 
scoparia, a plant not previously reported from central California. The introduction 
to the flora contains a short discussion of the vegetation and the geology of San 
Francisco and a short history of past botanical work on the flora of San Francisco. 
Descriptions and keys are not provided, but detailed distributional notes are included, 
and specimens are cited for most species. Authors’ names are spelled out in full. The 
photographs, most of them original, picture typical plant communities, and some 
of them show the inroads which civilization is making into the environment. 

This work is a most welcome addition to the existing local floras of various parts 
of the San Francisco Bay region and will be especially useful to those interested in 
the distribution and future migration of weeds and garden escapes. It may be obtained 
at the University of San Francisco Bookstore, San Francisco—JoHn H. Tuomas, 
Dudley Herbarium, Stanford University. 


Die Nadelgeholze. By GERD KRUSSMANN. 304 pp., 362 figs. Paul Parey, Berlin 
and Hamburg. 1955. D.M. 39.60. 

The author of “Die Laubgeholze” (Broadleaved Trees) by the same publisher has 
now made a contribution to the literature on gymnosperms. The term ‘“Nadelgeholze”’ 
(Needle-leaved Trees) is used by the author as a traditional convenient term for 
all gymnosperms, including broad-leaved ones such as Ginkgo biloba, Agathis and 
Podocar pus. 

Gerd Kriissmann, dendrologist of the Botanical Garden in Dortmund, Germany, 
is the General Secretary of the International Dendrology Union. A revised edition 
of Beissner-Fitschen’s ‘Handbuch der Nadelholzkunde” (Handbook of Gymno- 
sperms) was expected after its third edition which was published in 1930. Although 
the present book does not represent a revision of this well-known “Handbook of 
Gymnosperms,” it answers a great need in dendrology. 

Kriissman’s book not only emphasizes the gymnosperms hardy in Central Europe, 
but it also attempts to cover very many other species, varieties and forms that are 
not hardy in Central Europe. The book is illustrated with excellent line drawings 
and black and white photographs. The source of each illustration is cited carefully 
by the author. 

The introductory chapter gives a good popular account on the gross morphology 
of gymnosperms and explains the botanical terminology used for the descriptions 
throughout the text. This certainly increases the practical value of the book among 
gardeners, beginners and amateurs. However, some may expect a more proper and 
careful usage of such terms as “flower” and “fruit” which are generally applied only 
to angiosperms. The author points out the absence of perianth and ovaries among 
gymnosperms, but he nevertheless uses these terms freely in the text. “Male” micro- 
sporangiate and “female” (ovulate or megasporangiate) cones (strobili) would be the 
proper terms to apply to the reproductive structures of gymnosperms. Three types 
of strobili are distinguished under the heading of “Fruits and seeds’: 1) Zapfen 
(cones), 2) Beerenzapfen (berry-like cones), 3) Friichte (fruits). Indeed it is pleas- 
ing to note the proper application of the term “berry-like cones” to designate the 
ovulate cones of Juniperus instead of the term “berries” which is a common mistake 
repeated in various manuals. On the other hand, the term “Friichte” (fruits) is used 
improperly to designate the fleshy seeds of Cephalotaxus, Torreya, Taxus, Podocarpus 
and Ginkgo. A fruit is a matured ovary of an angiosperm, and the usage of the term 
should be restricted to angiosperms. 


The systematic arrangement follows the classification scheme proposed by Pro- 
fessor R. Florin, Stockholm. According to this scheme the Gymnospermae is divided 
into four classes: Class 1, Cycadopsida, comprises the orders Pteridospermae, Cay- 
toniales, Cycadales, Nilssoniales, Bennettitales, Pentoxylales, Ginkgoales; Class 2, 


1959] REVIEWS 31 


Coniferopsida, covers Cordaitales and Coniferae; Class 3, Taxopsida, includes only 
one order, Taxales; Class 4, Chlamydospermae, includes one order, Gnetales. 

It is interesting to note that this system of classification segregates Podocarpus, 
Cephalotaxus and their allies from the Taxaceae and treats them as separate families 
such as Podocarpaceae and Cephalotaxaceae under the order Coniferae. The Taxaceae 
comprises the genera Amentotaxus, Torreya, Austrotaxus, Nothotaxus and Taxus. 

The book gives the descriptions of the orders, families and genera of recent gym- 
nosperms, which are summarized nicely without unnecessary details. The lack of 
keys seems to be a disadvantage which would render the use of the book somewhat 
difficult among beginners and amateurs. However, this disadvantage is highly reduced 
by the comparison of specific differences under each genus, these being presented in 
beautifully prepared tabular form. 

The special part of the book dealing with the descriptions of the genera and species 
of gymnosperms covers 261 pages in which the genera as well as the species of each 
genus are arranged alphabetically. The generic descriptions cover gross morphological 
features of the vegetative and reproductive structures, but also provide some limited 
information at the anatomical level such as the number and position of resin ducts, 
vascular bundles, stomatal bands, pollen grains, basic chromosome numbers, etc. 

The approximate number of species under each genus is cited in all cases, but I 
believe some of these figures represent out-of-date information. For instance the 
approximate number of the species of the genus Pinus is cited as “80 species,” which 
agrees with the citation of Engler and Gilg’s “Syllabus der Pflanzenfamilien” (cf. 3rd 
edition, 1919). Most recent publications accept 90 or more species under this genus. 
Likewise 40 species are cited for Juntperus while other recent publications cite as 
many as 70 species for this genus. Naturally all newly established species are subject 
to acceptance or rejection by taxonomists, but I believe that the remarkable increase 
in the number of species of gymnosperms in recent years is the result of more careful 
and detailed studies. Therefore this increase is to be expected and should be taken 
into consideration. 

The distribution maps of genera are quite practical and more or less of the same 
nature as the distribution maps of the genera and families presented in J. Hutchin- 
son’s “Families of Flowering Plants.” Ranges of distribution and places of main 
occurrences are also given following the descriptions of genera and species. In some 
instances the distribution maps disagree with cited ranges and sometimes incomplete 
data are also obvious. For example the distribution map of the European species of 
Abies taken after Mattfeld is somewhat different from Kriissman’s distribution map 
of this genus. The latter’s figure 4 excludes the distribution of Abies nordmanniana 
and fails to indicate the western extension of A. alba in Pyrenees. Likewise the dis- 
tribution of Torreya californica is neglected in figure 37 although it is cited under the 
description of this species on page 285. 

Consistent spelling of the locality names would be desirable. “Cilicischen Taurus’’ 
appears intermittently with the germanized spelling, “Zilizischen Taurus.” Further- 
more the usage of ancient, discontinued names of localities should be avoided wher- 
ever possible. For example, “Cilicia” of ancient ages has now been replaced by the 
Turkish name, “Adana.” 

Rather a large number of gymnosperms of the Pacific Coast of the United States 
and Canada is included in the book, and therefore it can serve as a good reference to 
the botanists, students and amateurs of this area. All species of the genus Abies along 
the Pacific Coast are included in the work. With the exception of Cupressus pygmaea 
Sarg. and C. sargenti Jepson, all other Cupressus species native to the Pacific Coast 
are taken into account. Furthermore the book comprises all Pacific Coast pines except 
two recently described species, Pinus remorata Mason (1930) and P. washoensis 
Mason et Stockwell (1945). Three-needle pines such as Pinus attenuata, P. coulteri 
and P. jeffreyi are listed under the two-needle pines on page 209 erroneously although 
the cross sectional outlines of their needles on the following page show clearly that 
they belong to the three-needle section of the genus. 


32 MADRONO [Vol. 15 


Sequoia sempervirens (Lamb.) Endl. and Sequoiadendron giganteum (Lindl.) 
Buchholz are treated under the same genus although the proposal of J.T. Buchholz 
regarding the generic segregation of the sequoias is accepted by many in modern lit- 
erature. The reviewer believes that there are more than sufficient morphological, 
geographic and ecological differences between the two sequoias to justify their seg- 
regation. . 

The book contains very valuable information on the horticultural uses of different 
varieties and garden forms of gymnosperms. Those who can correlate the hardiness to 
conditions of Central Europe with the hardiness to conditions of other geographic 
areas can make a good use of the knowledge given in the book for judging the suit- 
ability of various gymnosperms. 

Toward the end of the book a small chapter enumerates the most important pineta 
in the world. Seventy-seven institutions or localities are listed for eighteen countries 
in Europe and North America. The author admits the incompleteness of his list and 
expresses his desire for receiving the recommendations of his readers for the exten- 
sion of the list. In this connection I would like to suggest the addition of the follow- 
ing arboreta in the United States: Eddy Arboretum at the Institute of Forest Genetics 
in Placerville, California, which has the most complete living collection of Pinus in 
the world; Westtown School Arboretum in Philadelphia for Abies and Picea; Morton 
Arboretum in Lisle, Illinois, for Picea, Taxus and Juniperus. 

The list of literature is short, but it comprises the principal standard references 
on gymnosperms. 

Finally, an index of invalid synonyms of gymnosperms concludes the book. This 
index is extremely useful to the reader since it clears many confusions in nomen- 
clature. 

In general the book is a treasury of information about gymnosperms in spite of 
its relatively small size. Both the author as well as the publisher deserve congratula- 
tions for this valuable publication ——BAkr KaAsapricit, Department of Biology, Mills 
College, Oakland 13, California. 


NOTES AND NEWS 


The second issue of the Index to Plant Chromosome Numbers, compiled from 
nearly 300 journals published in 1957, is now ready for distribution. There are around 
2000 listings of original chromosome counts from the entire plant kingdom and a 
bibliography of 196 papers from which the listings were taken. Preparation of the 
Index has been supported in part by a grant from the National Science Foundation 
of the U.S.A. The price of each issue is $1. Orders for subscriptious may be sent to: 

Dr. C. Ritchie Bell 

Department of Botany 

University of North Carolina 
Chapel Hill, North Carolina, U.S.A. 


Information on the location and history of any trees of Cedrus or Sequoia (includ- 
ing Sequozadendron) which were planted in the Pacific coastal states prior to 1900 
will be appreciated by E. E. Stanford, Department of Botany, College of the Pacific, 
Stockton, California. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree to bear the cost of the ad- 
ditional pages at the rate of $15 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. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


VolumeI,1916-1929 . . . . . $5.00 
Volume IT, 1930-1934. . . . . 5.00 
Volume ITT, 1935-1936 . . . . 5.00 
Volume IV, 1937-1938 . . . . 5.00 
Volume V, 1939-1940. . . . . 5.00 
Volume VI, 1941-1942 . . . . 5.00 
Volume VII, 1943-1944 . . . . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 . . . . 6.00 
Volume X,1949-1950. . . . . 7.00 
Volume XI, 1951-1952 . . . . 8.00 
Volume XII, 1953-1954 . . . . 8.00 
Volume XIII, 1955-1956 . . . 8.00 
Volume XIV, 1957-1958 . . . . 8.00 


Single numbers, 
Volumesland2 . ... . .50 


Succeeding numbers, 

(except Volume XI,No.2) . . . 1.00 

Volume X,'No:.2. 0%... Ge) ee 1.50 
The subscription price of MapRONo is $4.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


MADRONO 


VOLUME 15, NUMBER 2 APRIL, 1959 


PAGE 


Rep ALGAL PARASITES OB@URERING OM PPEMBERS OF THE 
GELIDIALES, Kung-Chu Fan and George F. Papenfuss 33 


XEROPHYLLUM TENAX, SQUAWGRASS, ITS GEOGRAPHIC 
ities 
DISTRIBUTION AND ITS BEHAVIOUR ON Mount RAIN- 
IER, WASHINGTON, Sue Merrick Maule 39 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 49 


TYPIFICATION OF PROSOPIS ODORATA ToRR. & FREM. 
(LEGUMINOSAE), Lyman Benson 53 


Two NEW SPECIES OF HELIANTHUS FROM NEw MEXxIco, 
R.C. Jackson 54 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF 
THE GENUS MIMULUS (SCROPHULARIACEAE), IIT, 


Barid B. Mukherjee and R. K. Vickery, Jr. 57 
Review: G. Erdtman, Pollen and Spore Morphology/ 
Plant Taxonomy (Jane Gray) 62 


NOTES AND NEws: THE OCCURRENCE OF PILOSTYLES 
THURBERI (RAFFLESIACEAE) IN CALIFORNIA, 
Job Kutjt; NOTES ON THE FLORA OF ARIZONA, 
Charles T. Mason, Jr. 63 


A WEST AMERICAN JOURNAL OF BOTANY 


PUBLISHED QUARTERLY BY THE ae 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 

$4.00 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. 
JouNn F. Davipson, University of Nebraska, Lincoln. 

Ivan M. JoHnstTon, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MiLpreD E. MATHIAS, University of California, Los Angeles 24. 
Marion OwnsBEY, State College of Washington, Pullman. 

Tra L. Wiccins, Stanford University, Stanford, California. 


Secretary, Editorial Board —ANNETTA CARTER 


Department of Botany, University of California, Berkeley. 


Business Manager and Treasurer—WINSLowW R. Briccs. 
Department of Biology, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: James R. Sweeney, San Francisco State College, San Francisco, Cali- 
fornia. First Vice-president: Baki Kasapligil, Mills College, Oakland, California. 
Second Vice-president: Henry J. Thompson, Department of Botany, University of 
California, Los Angeles, California. Recording Secretary: Mary L. Bowerman, De- 
partment of Botany, University of California, Berkeley, California. Corresponding 
Secretary: Francia Chisaki, Department of Botany, University of California, Berke- 
ley, California. Treasurer: Winslow R. Briggs, Department of Biology, Stanford 
University, Stanford, California. 


1959] FAN AND PAPENFUSS: ALGAL PARASITES 35 


RED ALGAL PARASITES OCCURRING ON MEMBERS OF THE 
GELIDIALES 


Kunc-CHu FAN AND GEORGE F. PAPENFUss! 


Three genera and four species of red algae have been reported as para- 
sitic on the various members of the Gelidiales. They are Syringocolax 
macroblepharis Reinsch (1875; Martin and Pocock, 1953), Gelidiocolax 
microsphaerica Gardner (1927), Choreocolax suhriae Martin et Pocock 
(1953), and C. margaritoides Martin et Pocock (1953). These species are 
reviewed and two parasitic red algae occurring on Pterocladia, also a 
member of the Gelidiales, collected by Setchell in New Zealand in 1904 
and by Papenfuss in New Zealand in 1949 and in Hawaii in 1942, are 
described as new. 

Gelidiocolax microsphaerica was obtained in 1911 by Gardner on Geli- 
dium pulchrum growing at Balboa Beach, Orange County, California. 
Dangeard in 1952 reported Gelidiocolax microsphaerica (in error as G. 
hemisphaerica) from Dakar. Subsequently, the species was reported by 
Dawson (1952, 1953) from the shores of Isla San Martin, Baja California. 
Careful examination of specimens of Gelidium pulchrum (collected at 
Punta Descanso, Baja California, Mexico, Dawson 131-45, April 8, 1945, 
UC 694021) has resulted in the finding of additional specimens of Geli- 
diocolax micros phaerica. The tubercle produced by this species is spheri- 
cal in form, about 175—225v. in diameter, and difficult to detect without 
the aid of a handlens; it is mostly composed of the reproductive tissue of 
the parasite. The carpogonial branch is composed of two cells, the sperma- 
tangia are produced in chains, and the tetrasporangium is cruciately 
divided. Although currently placed in the Gelidiaceae (cf. J. & G. Feld- 
man 1958), it appears more likely that Gelidiocolax belongs in the Chore- 
ocolacaceae. 

Choreocolax suhriae is parasitic on Suhria vittata in South Africa; the 
wart-like swelling is composed of a dense mixture of parasite and host 
tissues; it is about 2-3 mm. broad and about 1 mm. high when mature. 
During maturation, many or sometimes most of the surface cells of the 
parasite produce reproductive organs. Its carpogonial branch is two-celled 
like that of Gelidiocolax micros phaerica and not four-celled as in Choreo- 
colax polysiphoniae, which is the type species of Choreocolax (Sturch 
1926). In fact C. suhriae resembles G. microsphaerica so much that it 
should be placed in Gelidiocolax instead of in Choreocolax. The following 
combination is therefore proposed: Gelidiocolax suhriae (Martin et 
Pocock) Fan et Papenfuss, comb. nov. (Choreocolax suhriae Martin et 
Pocock 1953, p. 48). 

Choreocolax margaritoides is parasitic on Beckerella pinnatifida in 


1 This study was aided by a grant-in-aid from the National Science Foundation 
to the second author. Dr. Mary A. Pocock kindly furnished the material of Choreo- 
colax suhriae and C. margaritoides. 


Maprono, Vol. 15, No. 2, pp. 33-64, May 15, 1959. 


NIAN aay 2 6 1958 
BS TION MAY 2 


34 MADRONO [Vol. 15 


South Africa. Tetrasporic specimens of this species were examined. The 
pustule is roundish, 200—290y. in diameter, and is mostly composed of 
parasite tissue. This species is closely related to Gelidiocolax micros phae- 
vica. However, the tetrasporic pustule is slightly larger than that of G. 
microsphaerica, which is 175—225y. in diameter. The tetraspores of C. 
margaritoides are 12-18y in length; those of G. microsphaerica are 22— 
28u. in length. In our opinion C. margaritoides is a species of Gelidio- 
colax. The following combination is therefore proposed: Gelidiocolax 
margaritoides (Martin et Pocock) Fan et Papenfuss, comb. nov. (Chore- 
ocolax margaritoides Martin et Pocock 1953, p. 50). 


Gelidiocolax mammillata Fan et Papenfuss sp. nov. Planta in Ptero- 
cladia sp. parasitica, constans e filamentis multis irregulariter ramosis, in 
telam hospitis profunde penetrantibus, tuberculum solidum verruciforme 
efficientibus; tuberculum maturum 0.3—1 mm. lat., 0.3-0.5 mm. alt., e tela 
et hospitis et parasitici compositum, proiectiones mammillatas multas in 
superficie habens; cystocarpi, spermatangia, tetrasporangiaque in proiec- 
tionibus mammillatis praecipue producta, et in cubiculis per telas hospitis 
effectis reperta; ramus carpogonialis bicellularis; spermatangia in catenis 
formata, per septa transversa ab extremitatibus terminalibus cellularum- 
matrum spermatangialium successive abscissa; tetrasporangia cruciate 
divisa. 

Plant parasitic on Pterocladia sp., consisting of many irregularly 
branched filaments which penetrate deeply into the host tissue, producing 
a wart-like solid tubercle; tubercle when mature 0.3-1 mm. wide and 
0.3-0.5 mm. high, composed of both host and parasite tissues, with many 
mammillate projections on the surface; cystocarps, spermatangia and 
tetrasporangia produced primarily in the mammillate projections and 
occurring in chambers produced by host tissue; carpogonial branch two- 
celled; spermatangia formed in chains, successively cut off by transverse 
septa from the terminal ends of the spermatangial mother cells; tetra- 
sporangia cruciately divided. 

Type. Hanauma Bay, Oahu, Hawaiian Islands, March 1, 1942. G. FP. 
Papenfuss (UC 1058497). Figs. 3, 6, 7, 9, 10. 


Pterocladiophila hemisphaerica Fan et Papenfuss gen. et sp. nov. 
Planta in Pterocladia lucida parasitica, constans e filamentis multis ir- 
regulariter ramosis in telam hospitis profunde penetrantibus, tuberculum 
solidum plus minusve hemisphericum efficientibus; tuberculum maturum 
0.4-1 mm. lat., 0.4-0.8 mm. alt. e telis et hospitis et parasitici composi- 
tum, in superficie inaequale; cystocarpi, spermatangia, tetrasporangiaque 
in concepticulis formata; ramus carpogonialis bicellularis; spermatangia 
in catenis producta, per septa transvera ab extremitatibus terminalibus 
cellularum-matrum spermatangialium successive abscissa; tetrasporangia 
zonate divisa, pavimento lateribusque conceptaculi introrsus obducentia. 

Plant parasitic on Pterocladia lucida, consisting of many irregularly 
branched filaments which penetrate deeply into the host tissue, producing 


1959] FAN AND PAPENFUSS: ALGAL PARASITES 35 


SN 
l, mp thy fed 
A SELF po EN 
aS SEX 7 


1 nis phaerica: 1, tetrasporangia within the concep- 
rpogonial branches. Fic. 3. Gelidiocolax mammillata (marginal stippling) 
with spermatangia (solid stippling). 


= 
— 
to 
as, 
~~ 
~~ 
i=) 
& 
v 
= 
™— 
= 
v 


MADRONO 


Fics. 4, 5, 8. Pterocladiophila hemisphaerica: 4, spermatangia; 5, part of goni- 

porangia; 8, habit of parasite on Pterocladia lucida. Fics. 6, 7. 

Gelidiocolax mammillata (stippled cells): 6, gonimoblast with carposporangia; 7, 
Pterocladia sp. 


1959 | FAN AND PAPENFUSS: ALGAL PARASITES OW 


10p 


Fics. 9, 10. Gelidiocolax mammillata (marginal and solid stippling); 9, tetra- 
porangia; 10, carpogonial branches. 


a more or less hemispherical solid tubercle; tubercle when mature 0.4—1 
mm. wide and 0.4—0.8 mm. high, composed of both host and parasite tis- 
sues, uneven on surface; cystocarps, spermatangia and tetrasporangia 
formed in conceptacles; carpogonial branch two-celled; spermatangia 
produced in chains, successively cut off by transverse septa from the ter- 
minal ends of the spermatangial mother cells; tetrasporangia zonately 
divided, lining floor and sides of the conceptacle. 

Type. Island Bay near Wellington, New Zealand, June 1904. W. A. 
Setchell 6098 (UC 1141475) [separated from host plant, Pterocladia 
lucida (UC 95977) |. Figs. 1, 2, 4, 5, 8. 

Additional material. New Zealand: Kaikoura, 17 February 1949, 
Moore and Papenfuss (UC 1058496); Goose Bay 10 miles south of Kai- 
koura, 18 February 1949, Afoore and Papenfuss (UC 1058495). 

Pterocladiophila is readily distinguished by its zonately divided tetra- 
sporangia (fig. 1), formed within conceptacles, from all the known genera 
of parasitic red algae except Chaetolithon Foslie (1898; Kylin 1956), 
Choreonema Schmitz (1889; Suneson 1937), and Polvporolithon Mason 
(1953) in which three genera the sporangia are also zonately divided and 
localized in conceptacles. Chaetolithon, Choreonema and Polyporolithon 
belong to the family Corallinaceae, which family is almost exclusively 
composed of calcareous algae. Also, as far as known, the spermatangia 
are not produced in chains in the Corallinaceae as they are in Pterocla- 
diophila. 

Pterocladiophila cannot be assigned to the Choreocolacaceae owing to 
its zonately divided tetrasporangia which are formed in deeply sunken 
conceptacles (fig. 1). Nor can it be assigned to the Corallinaceae (although 
it resembles some members of this family by its simple, two-celled carpo- 


38 MADRONO [Vol. 15 


gonial branch and its zonately-divided tetrasporangia produced in con- 
ceptacles) because it is not calcified as all the Corallinaceae are with the 
exception of Schmitziella (Batters 1892, Suneson 1944); it also differs 
from members of this family in having the spermatangia produced in 
chains (fig. 4). A new family Pterocladiophilaceae is therefore suggested 
here, and this family is tentatively assigned to the order Cryptonemiales. 
This family is characterized by the following features. 

Pterocladiophilaceae Fan et Papenfuss fam. nov. Thallus haud calci- 
factus ramo carpogonio bicellulato spermatangia catenulata septis trans- 
versis termino cellulae maternae spermatangialis enata tetrasporangia 
zonatim divisa in conceptaculis circumscripta. 

Thallus not calcified; carpogonial branch two-celled; spermatangia 
produced in chains, succesively cut off by transverse septa at the terminal 
end of the spermatangial mother cell; tetrasporangia zonately divided, 
formed in conceptacles. 

Department of Botany, 


University of California, 
Berkeley, California 


LITERATURE CITED 


Batters, E. A. L. 1892. On Schmitziella; a new genus of endophytic algae, belonging 
to order Corallinaceae. Ann. Bot. 6:185-194, pl. 10. 

DANGEARD, P. 1952. Algues de la presqu’ ile du Cap-Vert (Dakar) et de ses environs. 
Botaniste 36:195-329, 15 figs., pls. 14-21. 

Dawson, E. Y. 1952 [1953]. Marine red algae of Pacific Mexico. Part I. Bangiales to 
Corallinaceae subf. Corallinoideae. Allan Hancock Pac. Exped. 17:1-239, pls. 
1-33. 

1953. [1954]. Notes on Pacific Coast marine algae. VI. Wasmann Jour. 
Biol. 11:323-351, pls. 1-6. 
FELDMANN, J. and GENEVIEVE FELDMANN. 1958. Recherches sur quelques Floridees 
parasites. Revue Gen. Bot. 65:49-127, figs. 1-30, pls. 1-2. 

Fosiiz, M. 1898. List of species of the Lithothamnia. K. Norske Vidersk. Selsk. Skr. 
1898(3). 11 pp. 

Garpn_er, N. L. 1927. New Rhodophyceae from the Pacific coast of North America. 
III. Univ. Calif. Publ. Bot. 13:333-369, pls. 59-71. 

Kyun, H. 1956. Die Gattungen der Rhodophyceen. xv-+673 pp., 458 figs. Lund. 

Martin, Marcaret T. and Mary A. Pocock. 1953. South African parasitic Florideae 
and their hosts. 2. Some South African parasitic Florideae. Jour. Linn. Soc. 
London Bot. 55:48—64, figs. 1-7, pls. 10-12. 

Mason, Lucite R. 1953. The cructaceous coralline algae of the Pacific coast of the 
United States, Canada, and Alaska. Univ. Calif. Publ. Bot. 26:313-390, pls. 27-46. 

Reinscu, P. F. 1875. Contributions ad algologiam et fungologiam. xii+104 pp., 130 
pls. Norimbergae. 

SturcH, H.H. 1926. Choreocolax polysiphoniae, Reinsch. Ann. Bot. 40:585-605, figs. 
1-15. 

SUNESON, S. 1937. Studien tiber die Entwicklungsgeschichte der Corallinaceen. Lunds 
Univ. Arsskr. N. F. Avd. 2, 33(2). 102 pp., figs. 1-42, pls. 1-4. 

. 1944. Notes on Schmitziella endophloea. K. Fysiogr. Sallskap. Lund 
Forhandl. 14:239-245, figs. 1-2. 

Scumitz, F. 1889. Systematische Ubersicht der bisher bekannten Gattungen der Flori- 

deen. Flora 72:435-456, pl. 21. 


1959] MAULE: XEROPHYLLUM TENAX 39 


XEROPHYLLUM TENAX, SQUAWGRASS, ITS GEOGRAPHIC 
DISTRIBUTION AND ITS BEHAVIOUR ON 
MOUNT RAINIER, WASHINGTON! 


SuE MERRICK MAULE” 


The objectives of the present study of Xerophyllum tenax are twofold; 
to compile information regarding its geographic distribution and to study 
the specific environmental requirements necessary for its vegetative 
growth and blossoming on Mount Rainier, Washington. Since the field 
observations have been confined to a single season, the results presented 
must be considered as tentative. In order to study the environmental 
requirements fully, stations with permanent protected plots would have 
to be established throughout the range of the species and observations 
made over a period of years. 

Xerophyllum tenax (Pursh) Nutt. (fig. 1) is a tufted, herbaceous, 
graminoid, perennial hemicryptophyte having a tuber-like woody root- 
stock bearing cord-like roots. It bears numerous grass-like, keeled, rigid 
leaves 5—10 dm. long, 5--10 mm. wide at the base and gradually tapering 
to a narrow, stiff and wiry tip, the margins rigid and serrulate (Peck, 
1941; Jepson, 1951). According to Dr. D. B. Lawrence (personal com- 
munication ), the young seedlings can be distinguished from those of Carex 
species by a grey-white bloom which covers the leaves of the Xerophyl- 
lum. Flowering occurs between May and September. The inflorescence 
stalks, 3-15 dm. high, are covered by leaf-like bracts which are reduced 
toward the top, and the raceme, 1—2 dm. long, is very dense, with slender 
pedicels 2—5 cm. long. The perianth is cream-colored, with lanceolate 
segments 6—10 mm. long, the stamens surpass the perianth, and the cap- 
sule is broadly ovoid, acute, 5—7 mm. long. 

Xerophyllum tenax is found from west-central California northward 
to northwestern Washington, and from Yellowstone National Park north- 
westward to southwestern Alberta and southeastern British Columbia. 
Along the coast, at least in northwestern Washington where habitat con- 
ditions are specifically recorded, it is found at sea level on bogs, and in 
the rain-shadow of the Olympic Mountains on gravelly “prairies” (Jones, 
1936). It occurs again high in the coast ranges, and from approximately 
2000 feet to 7000 feet in the Sierra-Cascade and Rocky Mountain ranges. 
Thus it is widely separated geographically from the eastern North Ameri- 
can species, X. asphodeloides (L.) Nutt., which grows at low altitudes on 
sandy acid pineland of the Atlantic Coastal Plain from North Carolina 


1 Requests for reprints should be addressed to Prof. D. B. Lawrence, Department 
of Botany, University of Minnesota, Minneapolis 14. 


* Sincere thanks to the following, without whose help this contribution would not 
have been possible: Robert William Maule, for his indispensible work in the field; 
Dr. Donald B. Lawrence, for help in planning and in organization of results; Dr. 
Gerald Ownbey, for help with taxonomic problems. 


40 MADRONO [ Vol. 15 


Fic. 1. Xerophyllum tenax with an abnormal stunted inflorescence, beside trail, 
Station 2, Mount Rainier. Normal inflorescence from another plant held in hand. 


1959] MAULE: XEROPHYLLUM TENAX 41 


to New Jersey, and again in the Blue Ridge Mountains and Appalachian 
Plateau in acid woods of Georgia, North Carolina, Tennessee, and Vir- 
ginia (Fernald, 1950; Small, 1933). 

The geographic distribution of Xerophyllum tenax is shown in a map 
(fig. 2), data for which was compiled from information supplied by staff 
members of various herbaria (Univ. Calif., Univ. Idaho, Univ. Oreg., 
Oreg. St. Coll., Univ. Wash., Wash. St. Coll., Nat. Mus. Canada, Univ. 
Alberta, Univ. Brit. Col.). The species occurs very sparingly in the coastal 
region near sea level from west-central California to northwestern Wash- 
ington, and again just below the summits of the coast mountains over 
almost the same latitudinal range. In the Sierra-Cascade range, it is found 
from Placer County, California, northward 700 miles to Stampede Pass, 
Washington, ranging in altitude from approximately 2000 to 6000 feet. 
It is not known to occur on Mount Shasta. In the Rocky Mountains com- 
plex it occurs from 2000 to 7000 feet, with the southernmost limit of its 
range along the southern boundary of Yellowstone National Park, whence 
it extends northwestward about 450 miles to Crow’s Nest Pass on the 
British Columbia-Alberta boundary. Westward it extends from the Rocky 
Mountain divide approximately 200 miles to northern Idaho, reaching its 
western limit on the summit of Mount Spokane, Washington, at 5800 
feet. In Idaho it ranges from southern Lemhi and Valley counties north- 
ward to the International Boundary and on in British Columbia to 
Kootenay Lake, a northwestward extent of 380 miles. 

Judging from the differences in habitat that have been reported in the 
literature, and the broadness of the geographic distribution pattern, it 
would be reasonable to expect that several races exist within the species. 
There appear to be definite distributional gaps between the plants grow- 
ing at sea level along the immediate coastal strip and those growing high 
in the coast range; similarly a distributional gap exists between the coast 
ranges and Sierra-Cascades, and another large one between the latter and 
the Rocky Mountains complex. Although the distribution map (fig. 2) 
would suggest continuity within the groups running north and south, it is 
quite possible that the plants growing in the north belong to quite different 
races from those growing in the outposts along the California coast, the 
southern Sierras, and the Rocky Mountains in northern Wyoming. It 
would be very interesting to take individuals from widely different geo- 
graphical areas and grow them adjacent to each other in experimental 
plots at several places within the distribution range. Dissimilarities would 
almost certainly be found. 


XEROPHYLLUM TENAX ON MOUNT RAINIER 


Between July 13 and August 23, 1955, field work was carried on at 
Mount Rainier National Park, Washington, and thirty-four stations were 
established where the species was studied. Figure 3 and Table 1 present 
the data for these stations and show through description and symbol those 
characteristics of the plants and of their local environments which were 


42 MADRONO [Vol. 15 


ALBERTA 


BRITISH COLUMBIA 


' : 
122° Mn rs [QP e ° 
; 120 a ery We W6 (14° "2° | 


Fic. 2. Distribution map of Yerophyllum tenax. 


1959] MAULE: XEROPHYLLUM TENAX 43 


considered significant in determining causes of local distribution. At thirty- 
three field stations detailed data were recorded as follows: date, time of 
day, location, elevation, degree and direction of slope, density of spacing 
measured as minimum distance between Xerophyllum clumps, presence 
of new or old flower stalks, composition of the adjacent vegetation, and 
history of fire. Where flowering was in progress the following were also 
recorded: root temperatures in degrees centigrade of flowering and of 
non-flowering plants three inches below ground surface, height in centi- 
meters of the flower stalk up to base of the raceme, and diameter of flow- 
ering stalk in millimeters measured just above basal leaves. 

From the data accumulated at these field stations, it appears that the 
distribution of X. tenax within Mount Rainier National Park seems to be 
influenced by a number of factors, including soil temperature, elevation, 
and direction and angle of slope. 

There are some factors, for example soil water content, which seem to 
have no apparent influence on its distribution. Individuals were found 
thriving equally well on dry sunny hillsides and on moisture saturated soil 
immediately below rapidly disappearing snow banks. This latter observa- 
tion seems related to the occurrence of X. tenax on bogs near sea level. 
The amount of shading by an overstory also does not seem to be a limit- 
ing factor for vegetative growth. Although all the plants found blooming 
in 1955 were growing in open meadows, light woods, or shrubby areas, and 
not in dense forests, the ability of the adult plants to survive did not seem 
to be affected by the amount of shading they received. Plants were found 
growing well vegetatively in dense forests where little or no direct sun- 
light filtered through the canopy and also on slopes exposed to the sun 
for as long as twelve hours a day. 

Most of the stations were visited between 10 a.m. and 3:30 P.M. on days 
of roughly equivalent fair meteorological conditions over a period of about 
forty days between July 13 and August 25. Recorded air temperatures 
ranged from 13°C. to 30°C., and soil temperatures three inches below 
the surface beneath the leaf crowns of X. tenax ranged from 9°—18°C. 

Within the Park, only the upper limit of distribution could be ascer- 
tained, as plants were found growing at the lowest easily accessible bound- 
aries of the Park at 2000 feet and also somewhat lower outside. Although 
most of the plants were found below 6000 feet, one group of plants was 
found near Panhandle Gap, Station 22, at 6800 feet. This elevation was 
the upper limit of all vascular plant growth here, with only lichens and 
mosses occurring higher. Within this area there were numerous perennial 
snow fields. 

The chief factor that did appear to affect the distribution was direction 
of slope, and this in turn influences length of snow-free growing season 
and soil temperature. Xerophyllum tenax was found growing on south- 
facing slopes, one as steep as 55 degrees, at ten stations, on southeast 
slopes at seven, on southwest slopes at four, on west slopes at three, on 
east slopes at two, and on north slopes at only one station. In this last 


44 


MADRONO [Vol. 15 


GLACIER 


COLUMBIA CREST 
ray 
14408 FEET 


UN aX 


\ 
\ 


\ 
— 
4 AS 
— 


d} 


MN 33 fu Kw ( 
32 


Fic. 3. Map of Mount Rainier National Park showing distribu- 
tion of Yerophyllum tenax (based on observations and reports in 
1955). The stations, numbered clockwise beginning in the northwest, 
are represented by circles connected to black dots (the actual sites). 


1959] MAULE: XEROPHYLLUM TENAX 


Thee 00° 
12P 30° KEY 
ELEVATION 


2. DIRECTION OF SLOPE 
3. OLD BURN 


4 DENSITY 

CLUMPS TOUCHING 
CLUMPS 0-2’ APART 
CLUMPS 2-4' APART 
CLUMPS 4-8° APART 


mon DOD 


CLUMPS OVER 8° APART 
5 DIAMETER OF INFLORESCENCE STALK, mM 
6 INFLORESCENCE HEIGHT TO 
LOWEST FLOWER, CM 
7. AIR TEMP, °C 
@ ROOT TEMP FL PLANT, °C 
9. ROOT TEMP NON-FL PLANT, °c 


QOG 6 


55'BLOOMING OLD STALKS NO BLOOMING 
OBSERVED OBSERVED OBSEAVED 

| O ©) ©) 
f BLOOMING OLD STALKS NQ BLOOMING 
| REPORTED REPORTED REPORTED 
I 
) 
é 
i \ 
4X oS 
7 OPEN MEADOW = MEADOW AND TREES SHRUBS 


LIGHT OENSE 
FOREST FOREST 


_ One or more study sites was established at each station. Traced from 
USCG topographic map, Mount Rainier National Park, Washing- 
ton, 1954. 


44 


MADRONO. [Vol. 15 7 1959] MAULE: XEROPHYLLUM TENAX 


snare 00! 
ir'so! KEY 


1b etevation 


tae 33 


ZoOMECTION OF 5LOFE 
3.019 aves 
A ocusity 
A chuues Tou: 
8 ces 0-2 


© cluurs 2-4" 
© clues aoe apaar 
1 chuwps ovta a! arnt 
DIAMETER OF IMFLOSESCEMCE STALK, mae 
6 ImrLomescemce HEIGHT 7 
Lowest Flomem, cM 
TAIR TEAR, © 
moot TEMP FL PLANT. °C 
moot Thur womerL PLAST, °C 


QO8 6 


ss'BLeomne uy stares 50 BLCOMING 


coesaves) Ons CAWES: opstaveo 
= = c 
6) (an 7 
© KF 
SJ 

pisos 1D STALKS no momane 
merontes —scronTep REPORTED 


vate 


Fic. 3. Map of Mount Rainier National Park showing distribu- 
tion of Xerophyllum tenax (based on observations and reports in 
1955). The stations, numbered clockwise beginning in the northwest, 


are represented by circles connected to black dots (the actual sites). | ton, 1954. 


One or more study sites was established at each station. Traced from 
USCG topographic map, Mount Rainier National Park, Washing- 


46 


MADRONO 


[Vol. 15 


TABLE 1. DESCRIPTIONS OF FIELD STATIONS WHERE STUDY PLOTS OF XEROPHYLLUM 
TENAX WERE ESTABLISHED ON Mount RAINIER IN 1955 


STATION 

No. DATE 

1 Aug. 2 
2 Aug. 2 
Sm UES Z 
4 Aug. 2 
5 (See Fig. 3° 
6 Aug. 1 
7 July 25 
8 July 25 
9 July 25 
10 July 25 
11 Aug. 25 
12. = Aug. 22 
13. Aug. 22 
14 Aug. 23 
15 Aug. 23 
16 Aug. 23 
17. —=— Aug. 23 
18 Aug. 23 
19 Aug. 23 
20 ~=Aug. 23 
21 ~Aug, 5 
22 + Aug. 17 
23 Aug. 16 
24 Aug. 16 
25 Aug. 25 


TIME 


3:30 p.m. 


3:15 p.m: 
2:45 p.m. 


11:30 a.m. 


3:45 p.m. 


12:15 p.m. 


12:00 m. 


11:00 a.m. 


12:00 m. 


2:20 p.m. 


9:00 a.m. 
10:45 a.m. 
12:30 p.m. 

1:00 p.m. 

2215 pam: 
4:30 p.m. 

2:45 p.m. 


12200;m: 


10:45 a.m. 


2:45 p.m. 


10:15 a.m. 


SLOPE 


15° south 


45° south 


10° west 


15° south 
and also 


gentle north 


10° south 


18° south 


25° west 


18° east 


south 

south 

5° south 
45° east 


southwest 


40° west 
south 
west 
40° west 
5° south 
18° south 


10° south 


15° west 


south 


LocaTION, SNOW AND SOIL 
CONDITIONS 


1 mile west Yellowstone Cliffs, 
North Loop trail; soil wet. 
Yellowstone cliffs, North Loop trail. 


Windy Gap, North Loop trail; large 
snow fields adjacent. 


Lake James, North Loop trail; snow 
areas present. 


West slope Grand Park, North Loop 
trail; numerous snow fields nearby. 


South edge of Grand Park, North 
Loop trail; no snow. 


South edge of Grand Park, North Loop 
trail; snow fields present. 


1 mile north Berkeley Park shelter, 
North Loop trail; snow fields present. 


Yakima Park. 

White River Entrance Station. 
Deadwood Lakes; very wet. 
Ghost Lake. 


Cascade Crest trail. 


Three Lakes-Ohanapecosh trail. 
Cascade Crest trail. 
Shriner’s Peak Lookout. 


0.5 mile south Panhandle gap, Won- 
derland trail; highest elevation at 
which flowering plants were found. 


Cowlitz Divide, 1 mile south Indian 
Bar, Wonderland trail. 


2 miles east Nickel Creek patrol cabin, 
Cowlitz Divide, Wonderland trail. 


Paradise Valley. 


1959 | MAULE: XEROPHYLLUM TENAX 47 


STATION LocaTION, SNOW AND SOIL 
No. Date TIME SLOPE CONDITIONS 


26 July 13 10:35a.m. 12° southeast 0.1 mile below Canyon Rim; snow 
present; dry soil. 


27. July 13. «10:15am. 30° south Narada Falls. 

28 July 12) 10:30a.m. 13° south 0.7 mile below Narada Falls; melting 
snow fields nearby, soil wet. 

29. July 13 10:40a.m._ 12° east 1 mile below Christine Falls; 
soil dry. 

30 July 13) 10:50am. 38° south Christine Falls; only a few plants in 
bloom, all against rock cliff. 

31 Aug. 31 Nisqually entrance. 

82 july 17 3:30p.m. 38° south 2 miles south of Round Pass, 
West Side road. 

33. July 17 3:40p.m. 55° south 2 miles south Round Pass, 
West Side road. 

34 Aug. 17 Golden Lakes. 


instance the surface was nearly level, and the plants had evidently spread 
over the crest of a small hill from a south-facing slope; they indicated 
no sign of present or past flowering. 

It has been noted in former years by Mount Rainier Park rangers that 
X. tenax increases greatly in numbers two to three years after a forest fire. 
Several competent observers related having seen it growing densely in old 
burns; these observations agree with those of the present writer. 

A few factors that seemed to have relatively little effect on distribution 
definitely did appear to inhibit flowering. The one most obviously im- 
portant in bringing about flowering during the 1955 season was the 
amount of light reaching the plant. Although old flowering stalks were 
found in dense forest as well as in open meadows in 1955, no plants were 
observed to bloom in dense shade. Instead, all flowering plants were con- 
fined to open meadows, light forests, or shrub-covered areas. Other limit- 
ing factors controlling flowering are probably temperature, especially of 
the soil, and, as will be evident later, the length and warmth of the grow- 
ing season the previous year when the flower buds were being formed. 

Soil temperatures about the roots at a three-inch depth beneath the 
clumps of flowering plants were between 0.5°—2.5°C. higher than those 
beneath non-flowering plants in the same vicinity. At only two stations 
were the root temperatures of flowering and non-flowering plants the 
same, and in no case was root temperature of the non-flowering plant 
found higher than the flowering one. Great care was taken to insure that 
the two plants studied at each station were on terrain of approximately 
the same amount of moisture, sunlight, etc. Perhaps the differences in 
recorded soil temperatures were due to slight differences in topography, 


48 MADRONO [Vol. 15 


humus content of the soil, and qualitative and quantitative differences in 
the ground cover. 

At most stations, no plants were seen flowering in 1955, although most 
plants retained old flowering stalks. At no time or place during the entire 
season were great displays of blossoms found, as had been reported in 
previous years. At the few stations where blossoms were present, only a few 
plants were involved and incidence of flowering did not seem especially 
related to density of spacing of clumps. Jepson (1901) has suggested 
that X. tenax blooms only once every seven years on Mount Tamalpais, 
Marin County, California. It is not likely that a similar flowering cycle 
exists on Mount Rainier. The 1955 season seemed to be the only season 
in several years in which X. tenax had not flowered heavily. 

Inasmuch as flower-stalk elongation in this species begins in early sum- 
mer, the 1955 flowering stalks were quite surely initiated during the 1954 
growing season. Dr. A. W. Harrison of the University of Washington has 
observed the weather conditions on Mount Rainier for several summers. 
He reported (personal communication) that the summer of 1954 was one 
of the “‘poorest”’ during the last ten years, and that the last ten years have 
been below average in temperature, rainfall, and sunny days. In 1954 the 
snow disappeared at an elevation of 4500 feet on the south side of Mount 
Rainier one week later than in 1953 and four weeks later than in 1952. 
Probably the effectiveness of the growing season for Xerophvllum in 1954 
was reduced by cooler temperature during the summer, as well as by the 
later start due to the persistence of the snow cover. It is quite probable 
that the length of the 1954 growing season was not long enough or warm 
enough to initiate many flower buds. 


Department of Botany, 
University of Minnesota, Minneapolis 


REFERENCES 


FERNALD, M. L. 1950. Gray’s manual of botany. 8th ed. American Book Co., New 
York. 
Jepson, W.L. 1901. A flora of western middle California. Encina Publ. Co., Berkeley. 
. 1922. A flora of California. Vol. 1. Univ. Calif. Press, Berkeley. 
. 1951. A manual of the flowering plants of California. Univ. Calif. Press, 
Berkeley. 
Jones, G. N. 1936. A botanical survey of the Olympic Peninsula, Washington. Univ. 
Wash. Press, Seattle. 
Macoun, J. 1888. Catalogue of Canadian plants. Vol. 2. Dawson, Montreal. 
Peck, M. E. 1941. A manual of the higher plants of Oregon. Binfords and Mort, 
Portland, Oregon. 
RypsBerc, P. A. 1900. Flora of Montana and Yellowstone National Park. New Era, 
Lancaster, Pennsylvania. 
SMALL, J. K. 1933. Manual of the southeastern Flora. Printed by the author, New 
York. 
SMILEY, F. J. 1921. A report upon the boreal flora of the Sierra Nevada of California. 
Vol. 9. Univ. Calif. Publ. Bot. 


1959 | CHROMOSOME NUMBERS 49 
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 
(See Madrono 9:257-258. 1948.) 
SPECIES NUMBER COUNTED BY COLLECTION LocaLitTy 
LILIACEAE 
*Hesperocallis 
undulata 2n==24 H. Lewis, LA’ |Lewis in 1952, Borrego Valley, 
A. Gray ul LA San Diego County, 
California 
*Nolina 
parryi 2n=20 H. Lewis, LA |O’Donnell Pinon Flats, 
S. Watson II in 1953, Riverside County, 
LA California 
*Zigadenus 
brevibracteatus nd H. Lewis, LA |Lewis in 1950, Red Rock Canyon, 
H. M. Hall I LA Kern County, 
California 
RANUNCULACEAE 
*Ranunculus 
occidentalis 2n—14 H. Lewis, LA |Lewis in 1955, Mather, Tuol- 
var. eisenii II LA umne County, 
A. Gray California 
CRUCIFERAE 
*Dithyrea H. Lewis, LA |Lewis in 1955, Borrego Valley, 
californica 2n=10 LA San Diego County, 
Harv. at California 
*Erysimum 
capitatum 2n==16 P. Raven, LA |Mathias 3027, Kern County, 
var. bealianum Bt LA California 
(Jeps.) Rossbach 
*Lyrocar pa 
palmeri n=20 P. Raven, LA |Lewis in 1957, Sentenac Canyon, 
S. Watson LA San Diego County, 
California 
*Streptanthus 
inflatus 2014 H. Lewis, LA |Lewis in 1957, Temblor Range, 
(S. Watson) II LA Kern County, 
Greene California 
LINACEAE 
*Cathartolinum 
digyvnum 2n=8 P. Raven, LA |Lewis in 1955, Mather, Tuol- 
(A. Gray) Small II LA 


‘umne County, 
California 


* Prepared slide available. 
1 Symbols for institutions are those listed by Lanjouw and Stafleu, Index Her- 
bariorum, Part I. Third Edition, 1956, Utrecht. 


(continued on p. 50) 


50 MADRONO [Vol. 15 
SPECIES NUMBER COUNTED BY COLLECTION LocaLitTy 
APOCYNACEAE 
F ; Quail Springs, 
Amsonia — 222 C. Epling, LA |Zewis in 1940, San Bernardino 
Mi qaaee LA County, 
y California 
*Amsonia — 222 C. Epling, Lewis in 1940, Quail Springs, 
aU LA LA San Bernardino 
Var. mentvosa County, 
ae Frem.) California 
BORAGINACEAE 
*Coldenia n=9 M. S. Cave Raven 13122, Kaibab Trail, 
canescens DC. UC UC so. rim of Grand 
Canyon, Coconino 
County, Arizona 
* palmeri A. Gray n=8 H.F.Chisaki, |Alava 1831, 27 miles south of 
UC JEPS Vidal, Riverside 
County, California 
* plicata (Torr.) n=8 H.F.Chisaki, |Alava 1809, Junction of highways 
Coville UC JEPS 80-98 and road from 
Ogilby, Imperial 
County, California 
* pur pus n==9 H.F.Chisaki, |Moran 6320, South of Jose Maria 
Brandegee UC UC Aguirre, Nuevo Leon, 
Mexico 
*Cryptantha ; : 
pterocarya n=12 H.F.Chisaki, |Raven 11723, Gila Mountains, 
(Torr.) Greene we LUKE Yuma County, 
Arizona 
*Heliotropium 
parviflorum (eas 6} H.F.Chisaki, |Chisaki 1168, San Blas, 
L. UC Nayarit, 
Mexico 
*M yosotis oe 
versicolor ne? H.F.Chisaki, |Alava 2295, Pitkin Marsh, 
(Pers.) Smith UC 1Ofe Pion oen 
alifornia 
VERBENACEAE 
Glandularia 
perackii n= 5 O. T.Solbrig, |Solbrig 2839, Sierra de San Luis, 
Covas et Schnack UC UE San Luis Province, 
Argentina (grown 
at the U.C. Bot. 
Garden, Berke- 
ley, Calif.) 
gooddingi n=16 O.T.Solbrig, (Solbrig 2803, Highway 66, 19.8 
(Briq.) LOG UC miles wect of 


O. T. Solbrig2 


| 


Seligman, 
Arizona 


2 Glandularia gooddingii (Briq.) O.T. Solbrig comb. nov. (Verbena gooddingii 
Briq., Ann. Conserv. & Jard. Bot. Genéve 10:103. 1907.) 


1959 | CHROMOSOME NUMBERS 51 
SPECIES NUMBER | COUNTED BY COLLECTION LOCALITY 
Glandularia 
canadensis (L.) n=15 O. T. Solbrig, |Solbrig 2840, Cultivated at the 
Small UC UC U.C. Bot. Garden, 
Berkeley, Calif. 
Verbena . 
bracteata n=—14 O.T.Solbrig, |Solbrig 2909, Highway 20, 
Lag. & Rodr. UC UC 12 miles west of 
Niles, Oregon 
prostrata O.T.Solbrig, |Chisaki, Hernandez, 
R. Br. ee UC Sharsmith & San Benito Connty, 
Solbrig 2823, California 
VE 
LABIATAE 
Agastache 
urticifolia 2n=9 H. L. Wedberg, |Lewis in 1956, Mather, Tuol- 
(Benth.) Ktze. I LA LA umne County, 
California 
*Monardella San Gabriel Moun- 
cinerea 2n=21 P. Raven, LA |Raven 11197, tains, Los Angeles 
Abrams I CAS County, California 
*Monardella Mather, Tuol- 
lanceolata 22 H. Lewis, LA |Lewis in 1955, umne County, 
Bentham I LA California 
*Monardella Piute Mountains, 
linoides 2071 H. L. Wedberg, |Raven 9327, Kern County, 
A. Gray a LA California 
SOLANACEAE 
*Chamaesaracha Silver Lake, 
nana 2n==12 P. Raven, LA |Lewis in 1956, Mono County, 
A. Gray II LA California 
*Solanum West Los Angeles, 
xantit 2n=12 H. Lewis, LA |Levene in 1949, Los Angeles County, 
A. Gray II California 
SCROPHULARIACEAE 
Orthocar pus Near Arvin, 
attenuatus Pn) H. Lewis, LA |Lewis in 1956, Kern County, 
A. Gray II LA California 
Orthocar pus Near Arvin, 
linearilobus n—24 H. Lewis, LA |Lewis in 1956, Kern County, 
Benth. LA California 
Orthocar pus Near Arvin, 
purpurascens oD ere 192 H. Lewis, LA |Lewzs in 1956, Kern County, 
A. Gray II LA California 


(continued on p. 52) 


By MADRONO [Vol. 15 
SPECIES NUMBER COUNTED BY COLLECTION LOCALITY 
CUCURBITACEAE 
* Marah Santa Monica 
macrocar pus 2n==32 H. Lewis, LA |Lewis in 1950, Mountains, Los 
(Greene) Greene I LA Angeles County, 
California 
COMPOSITAE 
*Chaenactis Douglasii 
var. rubricaulis n=6 R. O. Alava, Bacigalupi & Converse Mountain, 
(Stockwell) Ferris UC Alava 6497, Fresno County, 
JEPS California 
*Chryso psis 
foliosa Nutt. n=18 R.C. Jackson, |Jackson 2464, Sandoval County, 
KANU KANU New Mexico 
Haplopappus 
ciliatus n= 12 R.C. Jackson, | Richards 2648, Barber County, 
(Nutt.) DC. KANU KANU Kansas 
*Haplopap pus 
divaricatus 2n=10 R.C. Jackson, | Richards 2055, Barber County, 
(Nutt.) Gray KANU KANU Kansas 
*Haplopap pus 
hartwegti n=6 R.C. Jackson, |Jackson 2465, San Juan County, 
(Gray) Blake KANU KANU New Mexico 
*Haplopap pus 
pleuriflorus n==12 R.C. Jackson, | Jackson 2507, Rio Arriba 
(Gray) Hall KANU KANU County, 
New Mexico 
* Machaeranthera 
gymnocephala (DC.) n==5 R.C. Jackson, |Jackson 2516-1, |Otero County, 
Shinners KANU KANU New Mexico 
Machaeranthera 
tanacetifolia n=4 R.C. Jackson, |Jackson 2556, Socorro County, 
(H.B.K.) Nees KANU KANU New Mexico 
*Ratibida Bernalillo 
tagetes (James) N16 R.C. Jackson, |Jackson 2565, County, 
Barnhart KANU KANU New Mexico 
*Ratibida 
columnaris n=14 R.C. Jackson, |Jackson 2133, Otero County, 
(Pursh) Raf. KANU KANU New Mexico 
*Ratibida 
peduncularis n=14 R.C. Jackson, | Richards 564, Wilson County, 
var. picta Gray KANU KANU Texas 
* Senecio Bernalillo 
longilobus n=20 R.C. Jackson, |Jackson 2037, County, 
Benth. KANU UNM New Mexico 
Zinnia Bernalillo 
grandiflora n=24 R.C. Jackson, |/ackson 2002, County, 
Nutt. KANU UNM New Mexico 


1959] BENSON: PROSOPIS 53 


TYPIFICATION OF PROSOPIS ODORATA TORR. & FREM. 
(LEGUMINOSAE) 


LYMAN BENSON 


Clarification of the concept of nomina ambigua and nomina confusa in 
the 1956 edition of the International Code of Botanical Nomenclature 
makes unnecessary my proposal (1941) that Prosopis odorata be included 
in the list of Nomina confusa. Article 66 (Lanjouw, et al., 1956) declares 
nomina ambigua to be illegitimate unless a satisfactory separation of the 
ambiguous elements can be made. If separation can be made clearly, one 
constituent must be designated as a lectotype. With this in mind, re- 
examination of the nomenclatural status of Prosopis odorata Torr. & Frem. 
(in Frem. 2nd. Rept. Expl. Exped. Rocky Mts., Ore., Calif. 313, pl. 1, 
1845) is in order. 

Three specimen sheets of the Fremont collection upon which Prosopis 
odorata is based are in the Torrey Herbarium, New York Botanical Gar- 
den. On each sheet there is a fallen fruiting spike with a cluster of fruits 
from the screw-bean, Prosopis pubescens Benth. The bulk of the material 
on each sheet consists of vegetative branches with spines, young leaves, 
and spikes of flowers of the western honey mesquite, Prosopis juliflora 
(Swartz) DC. var. Torreyana L. Benson. This mixture of material from 
two species resulted from collection of specimens in April when the mes- 
quite was developing new leaves and was in flower. Probably the peculiar 
spiral pods were added from the ground. The two species often grow to- 
gether along washes or in springy areas, and confusion may have arisen 
either in collecting specimens or in sorting them. The ambiguity of the 
type specimen of Prosopis odorata has been discussed as follows (Benson, 
1941, pp. 753-754): 


“Type collections: (1.) P. odorata, ‘A characteristic tree in the mountainous part 
of northern [Alta] California, particularly along the Mohave [Mojave] and Virgen 
rivers [Virgin River in Nevada, Arizona, and Utah] the latter part of April.’ Accord- 
ing to Torrey, Pac. R.R. Rept. 4:82. 1855, the plant ‘is P. glandulosa (in flower 
only), with the pods of Strombocarpa pubescens. The error arose from the mixing 
of specimens in Fremont’s collections.’ Torrey appended the following note to one 
of the three type sheets now in the New York Botanical Garden, ‘I have scarcely a 
doubt that the leaves belong to P. (Algarobia) glandulosa—the fruit to Stromb. pu- 
bescens! I was led astray by Fremont placing the pods and the leafy specimens 
together—.’ Interpretations of P. odorata have varied. Standley, Contr. U.S. Nat. 
Herb. 23: 353. 1922, in discussing the nomenclature of P. juliflora var. velutina, 
argued as follows: ‘In case the plant should receive such recognition [as a species 
instead of a variety], the proper name for it is Prosopis odorata Torr. That name 
was based upon a flowering specimen of the present plant and fruit of P. pubescens, 
and for that reason has been discarded by most writers. Taking into consideration 
the specific name, ‘odorata,’ it seems reasonable to typify the name by the flowering 
specimen.’ The branches with leaves and young flowers included in the type of P. 
odorata are neither P. glandulosa (2.e., P. juliflora var. glandulosa) as supposed by 
Torrey nor, despite the presence of a few hairs on the rachilla and secondary leaflets, 
P. juliflora var. velutina as supposed by Standley. Instead, they are P. juliflora var. 
Torreyana. Britton and Rose, N. Amer. Fl. 23: 183. 1928, took up the specific epithet 
odorata under Strombocarpa for the screw-bean. According to Article 64 of the In- 


54 MADRONO [Vol. 15 


ternational Rules of Botanical Nomenclature, ‘A name of a taxonomic group must be 
rejected if the characters of that group were derived trom two or more entirely dis- 
cordant elements, especially if those elements were erroneously supposed to form part 
of the same individual. A list of names to be abandoned for this reason (Nomina 
confusa) will form Appendix V’ [not published]. The writer proposes that Prosopis 
odorata Torr. should be included in this list.” 


In the light of the 1956 rule, it is fortunate that in this instance seg- 
regation is clear and unmistakable, and the choice is easy. Prosopis pubes- 
cens Benth. (in Hook. Lond. Jour. Bot. 5:82. 1846) is established clearly 
in nearly all recent literature for the screw-bean, and changing the name 
by substituting Prosopis odorata, published one year earlier, is not desir- 
able, even though this was done by Britton and Rose (loc. cit.). Further- 
more, selection of the fruits alone for a lectotype would be less satisfac- 
tory than choice of the combination of twigs, leaves, and flowers. So long 
as Prosopis juliflora var. Torrevana is considered to be a variety, restric- 
tion of the type specimen of Prosopis odorata to include only the material 
from that population system will cause no nomenclatural upset. There- 
fore, the three sheets in the Torrey Herbarium of the New York Botanical 
Garden, the fruits excluded, are designated together as a lectotype of 
Prosopis odorata Torr. & Frem. 

If the lectotype rule had been adopted before 1941, the writer would 
have recombined the epithet odorata in varietal rank under Prosopis 
juliflora (Swartz) DC. rather than to add the new epithet var. Torrevana 
to the nomenclature. Should var. Torrevana be elevated to specific rank, 
however, it must be known as Prosopis odorata Torr. & Frem. 


Department of Botany, 
Pomona College 
Claremont, California 


LITERATURE CITED 
Benson, LyMAN. 1941. The mesquites and screw-beans of the United States. Am. 
Jour. of Bot. 28:748-754. 


Lanjouw, J., ET AL. 1956. International Code of Botanical Nomenclature adopted by 
the Eighth International Botanical Congress, Paris, July 1954. Utrecht. 


TWO NEW SPECIES OF HELIANTHUS FROM NEW MEXICO! 
R. C. JACKSON 


During a field study of the New Mexican species of Helianthus, two 
new species were discovered. Morphologically they appear to be related 
to Helianthus ciliaris DC. but differ from this species in several diagnos- 
tic characteristics. 


Helianthus heiseri sp. nov. Herba perennis 5—12 dm. alta; caulibus 
pluribus (vel unica) sparse strigosis, flavo-viridibus, striatis; foliis oppo- 


1 Field work for this study was supported by faculty research grants from the 
University of New Mexico. 


1959| JACKSON: HELIANTHUS 55 


sitis, sessilibus; laminis ad 7 cm. longis 2.3 cm. latis, cuneatis vel obtusis, 
acuminatis, serratis vel dentato-crenatis, ambis paginis strigosis resino- 
sisque, nervis tribus conspicuis instructis; capitulis 1-3 in pedunculis 
4-10 cm. longis; disco diam. 1.5—2.0 cm.; phyllariis lanceolatis, ciliatis, 
dorso levibus viridibusque, 2.6—3 mm. latis, 8-10 m. longis; radiis 16—20, 
10-12 mm. longis; disci corollis 4—4.5 longis, basi flavis puberulentisque, 
lobis purpureis puberulentisque; receptaculi paleis acutis purpureis disci 
corollas aequantibus, earum apicibus acutis dorso plus minusve villosis: 
achaeniis ca. 3 mm. longis in maturitate nigris; pappis florium discorum 
aristis 2 lanceolatis instructis; pappis eorum radiorum aristis 1—3 inaequa- 
libus lanceolatis item instructis. 

Perennial herb, 5—12 dm. high; stems one or several, sparingly strigose, 
yellowish-green, striate; leaves opposite, sessile, the blades up to 7 cm. 
long and 2.3 cm. wide, cuneate or obtuse at the base, acuminate at the tip, 
serrate or dentate-crenate on the margins, strigose and resin-dotted above 
and below, conspicuously 3-nerved; heads 1-3 on peduncles 4-10 cm. 
long; disc 1.5—2.0 cm. in diameter; phyllaries lanceolate, ciliate, smooth 
and green on the backs, 2.6-3.3 mm. wide, 8-10 mm. long; rays 16—20, 
10-12 mm. long; disc corollas 4—4.5 mm. long, yellow and puberulent at 
the base, the lobes purple and puberulent; pales of the receptacle acute, 
purple, about equal with the disc corollas, the tips moderately villous on 
the backs; achenes about 3 mm. long, black at maturity; pappus of the 
disc of 2 lanceolate awns, the pappus of the rays of 1-3 unequal, lanceo- 
late awns. 

Type. New Mexico. Grant County: shallow ditch near the Mimbres 
River, elevation ca. 5400 feet, September 30, 1957, Jackson 2521-1 
(IND.) Isotype. Jackson 2521-2 (UNM). 

Additional specimens from the type locality are deposited in the her- 
baria listed above. 

Thus far H. keisert is known only from the Mimbres River Valley 
where it has been observed at several sites other than the type locality. 
This entity was probably responsible for Torrey’s (1859) report of H. 
grosseserratus in “‘the valley of the Mimbres.” The two species have sev- 
eral characteristics in common. 


Helianthus crenatus sp. nov. Herba perennis 5—12 dm. alta; caulibus 
pluribus (vel unica), sparse strigosis, flavo-viridibus, striatis; foliis op- 
positis, sessilibus vel brevipetiolatis; laminis ad 9 cm. longis et 2.3 cm. 
latis, oblanceolatis, acutis, attenuatis, inaequaliter crenatis vel crenato- 
lobatis insuper strigosis resinosisque, subtus strigosis vel strigoso-pilosis 
resinosisque, nervis tribus conspicuis instructis; capitulis 1—6 in peduncu- 
lis 6-13 cm. longis; disco diam. 1.8—2.4 cm.; phyllariis anguste lanceolatis, 
ciliatis, dorso in maturitate strigosis, aliquantulum patentibus, 2—3 mm. 
latis, 10-12 mm. longis; radiis 16-20, 10-12 mm. longis; disci corollis 
5—5.7 mm. longis, basi flavis puberulentisque, lobis purpureis puberulen- 
tisque; receptaculi paleis quam disci corollis brevioribus, purpureis, dorso 


56 MADRONO [Vol. 15 


ad apicem versus moderate villosis; achaeniis ca. 3 mm. longis, in maturi- 
tate nigris; pappis florium discorum aristis 2 lanceolatis instructis; pappis 
eorum radiorum aristis 1—3 inaequalibus lanceolatis item instructis. 

Perennial herb, 5-12 dm. high; stems one or several, sparingly strigose, 
yellowish-green, striate; leaves opposite, sessile or short petioled, the 
blades up to 9 cm. long and 2.3 cm..wide, oblanceolate, acute, attenuate 
at the base, unequally crenate or crenate-lobed on the margins, strigose 
and resin-dotted above, strigose or strigose-pilose and resin-dotted below; 
conspicuously 3-nerved; heads 1—6 on peduncles 6—13 cm. long; disc 1.8— 
2.4 cm. in diameter; phyllaries narrowly lanceolate, ciliate, strigose on 
the backs, somewhat spreading at maturity, 2-3 mm. wide, 10-12 mm. 
long; rays 16-20, 10-12 mm. long; disc corollas 5—5.7 mm. long, the 
base yellow and puberulent, the lobes purple and puberulent; pales of the 
receptacle shorter than the disc corollas, purple, acute, the tips moderately 
villous on the backs toward the apices; achenes about 3 mm. long, black 
at maturity; pappus of the disc of 2 lanceolate awns; the pappus of the 
rays of 1-3 uneven lanceolate awns. 

Type. New Mexico. Sierra County: low area on the south side of Truth 
or Consequences, June 22, 1957, Jackson 2504-1 (IND). Isotype. Jack- 
son 2504-2 (UNM). 

Additional specimens are deposited in the herbaria listed above. 

Helianthus heiseri and H. crenatus are closely related morphologically. 
The main diagnostic differences separating the two are apparent in the 
descriptions. In addition, the two species are separated by flowering dates. 
Helianthus crenatus reaches its maximum flowering period during the 
latter part of July, whereas H. heiseri reaches its peak from the middle 
to the latter part of September. Geographically the two species are sepa- 
rated by the Black Mountain Range. 

It is quite possible that H. crenatus and H. heiseri may have been 
lumped with H. ciliaris DC. in the past. Some of the distinguishing char- 
acteristics are listed below in a comparison of the two new species with 
H. ciliaris as it occurs in the Rio Grande Valley of New Mexico. 


H. ciliaris H. heiseri H. crenatus 
Stems Mostly glabrous, Sparingly Sparingly 
glaucous strigose strigose 


Leaves Glabrous, bluish-green Strigose, light- or yel- Strigose to strigose- 


and glaucous lowish-green, resin- pilose, light- or yellow- 
dotted ish-green, resin-dotted 
Phyl- Ovate, obtuse, erect ap- Lanceolate, acute, loose, Narrowly lanceolate 
laries pressed, glabrous or glabrous on the back, somewhat spreading, 
subglabrous on the ciliate, 2.6-3.3 mm. strigose on the back, 
back, ciliate, about 3.5. wide, 8-10 mm. long ciliate, 2-3 mm. wide, 
mm. wide, 5-7 mm. long 10-12 mm. long 


In the population of H. crenatus, several individuals not typical of the 
population as a whole were found. These plants were different from the 


1959 | MUKHERJEE AND VICKERY: MIMULUS a7 


type in that some had long trichomes on the stem, the leaves were ashy 
grey, more heavily pubescent, and they apparently flowered earlier. Hy- 
bridization between /7. crenatus and H. ciliaris may have been responsible 
for some of these variations, but generally they represent combinations 
not present in either species. Furthermore, Heiser and Smith (1955) have 
reported the chromosome number of H. ciliaris as n—51 (also n=—34, 
Heiser unpublished) while the writer has found n=17 in H. crenatus. It 
may well be that these variations resulted from past hybridization with 


an unknown or now extinct species. 
Department of Botany, 
University of Kansas, 
Lawrence, Kansas 


LITERATURE CITED 


Heiser, C. B. ano D. M. SmitH. 1955. New chromosome numbers in Helianthus. 
Proc. Indiana Acad. 64:250-253. 

Torrey, J. 1859. United States and Mexican Boundary survey. Part I, Botany of the 
Boundary. Ex. Doc. No. 135, 34th Congress, Ist Session, 27-259. 

Watson, E. E. 1929. Contribution to a monograph of the genus Helianthus. Pap. 
Mich. Acad. 9:305-475. 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF 
THE GENUS MIMULUS (SCROPHULARIACEAE). III. 


Barip B. MUKHERJEE AND R. K. VICKERY, JR. 


This report? on the determination of chromosome numbers in the sec- 
tion Simiolus of the genus Mimulus is an integral part of a long range 
investigation into the taxonomy, genetics, and evolution of species in 
Mimulus (Vickery, 1951). Taken in conjunction with the previous counts 
(Vickery, 1955 and Mukherjee, Wiens, and Vickery, 1957), the counts re- 
ported here reveal a pattern of evolution in section Szmolus that involves 
both polyploidy and aneuploidy. 

A slightly modified version of the technical method of Swaminathan, 
Magoon, and Mehra (1954) was found to produce better results than the 
methods previously used (Vickery, 1955 and Mukherjee, Wiens, and Vick- 
ery, 1957). Buds expected to contain anthers at the desired stages of 
microsporogenesis were killed and fixed for 24 hours in a mixture of two 
parts absolute ethanol and one part glacial acetic acid saturated with 
ferric acetate. Acetic acid was substituted for the propionic acid called 


| This work was supported by the National Science Foundation. It forms a portion 
of the dissertation of the senior author submitted to the faculty of the University of 
Utah in partial fulfillment of the requirements of the Ph.D. degree. The authors wish 
to thank Drs. W.W. Newby and C. M. Woolf for their helpful criticisms of the 
manuscript. 


8 MADRONO LVol. 15 


on 


for in Swaminathan, Magoon, and Mehra’s schedule. Also, the buds were 
transferred after 24 hours to 70 per cent ethanol whereas their schedule 
called for leaving the buds in the fixative until used. The anthers were dis- 
sected out of the buds, smeared and stained in iron-aceto-carmine. Camera 
lucida drawings were made for each count and, in addition, photomicro- 
graphs were taken of the more intricate configurations. Each chromosome 
number reported is based on counts from an average of approximately ten 
microsporocytes. Herbarium specimens were prepared for each of the 
cultures studied. They will be deposited for future reference in the Garrett 
Herbarium of the University of Utah. 

A total of eleven cultures was studied during the present investigation 
(table 1). The cultures include representatives of seven species and varie- 
ties of the section Simiolus: M. guttatus DC., M. tilingii Regel var. tilingit, 
M. tilingi var. corallinus (Greene) Grant, M. glaucescens Greene, M. 
glabratus var. parviflorus (Lindl.) Grant, M. pilosiusculus HBK., and 
M. tigrinus hort. 

Of the five cultures of W/. guttatus examined, three, (5003, 5007, and 
5839) showed n=—14 chromosomes. The configurations were regular and 
similar to those observed previously for other cultures of M. guttatus 
(Vickery, 1955, Mukherjee, Wiens, and Vickery, 1957). However, the two 
cultures of MW. guttatus from Mather, California, exhibited frequent lag- 
ging chromosomes during the anaphase stage of the first meiotic division. 
Eight cells from three different plants of culture 5009 were observed at 
this stage of division. Two pairs of lagging chromosomes were found in 
each of two cells, one pair in each of three cells and no lagging chromo- 
somes in the remaining three cells. In addition, two cells from two differ- 
ent plants of the other Mather culture, 5010, were observed at the first 
anaphase stage of meiosis. These cells each contained a single pair of 
lagging chromosomes. The cause of the lagging chromosomes was not clear 
from the configurations studied. Observations of five cells in the first telo- 
phase stage of division revealed that in one case the lagging pair of 
chromosomes was not being included in either nucleus whereas in the 
other four cases, both members of the pair were being included in one 
nucleus producing 15 to 13 segregations of the chromosomes. Observa- 
tions of cells in the second metaphase stage of division confirmed the real- 
ity of these irregular segregations and indicated that, apparently, in a few 
cases, two pairs of lagging chromosomes had been included in the same 
daughter nucleus. Of such 16 to 12 segregations, only cells with 16 chro- 
mosomes were actually observed. Perhaps the number of cells studied was 
too small a sample to detect cells with 12 chromosomes which may be less 
viable than cells with 13 or more chromosomes. Of the 22 configurations 
of second metaphase chromosomes observed in microsporocytes of three 
plants of culture 5009, three contained 14 chromosomes, twelve contained 
15, five contained 13, and two contained 16. Of twelve configurations of 
second metaphase chromosomes observed in microsporocytes of plants of 
culture 5010 five contained the normal number of 14 chromosomes, four 


Un 
Ne) 


1959] MUKHERJEE AND VICKERY: MIMULUS 


TABLE 1. CHROMOSOME CounNTS IN MIMULUS, SECTION SIMIOLUS 


n=14 M. guttatus DC. 
Pescadero, San Mateo County, California, altitude 20 feet, Clausen 2083 
(5003), 
Yosemite Junction (marsh), Tuolumne County, California, altitude 1,350 
feet, Hzesey 559 (5007). 
Big Cottonwood Canyon, Salt Lake County, Utah, altitude 7,100 feet, 
Vickery 334 (5839). 
M. glaucescens Greene 
Richardson Springs, Butte County, California, altitude 600 feet, 
F.W. Pennell & A.A. Heller 25667 (5653). 

n—1W4+1o6r2 M. guttatus DC. 
Mather (meadow), Tuolumne County, California, altitude 4,600 feet, 
Hiesey 571 (5009). 
Mather (spring area), Tuolumne County, California, altitude 4,800 feet, 
Hiesey 569 (5010). 

n—15 M. tilingit Regel tilingii 
Mount Timpanogos, Utah County, Utah, elevation 7,800 feet, Del Wiens, 
Aug. 6, 1956 (5967). 

n= 24 M. tilingii var. corallinus (Greene) Grant 
Porcupine Flat, Mariposa County, California, altitude 8,000 feet, 
Hiesey 576 (5011). 

n—32 M. tigrinus hort. 
Commercial seeds from F. Kirchhoff and Co., Johannesburg, South Africa 
(5056). 

n—45 WM. glabratus var. parviflorus (Lindl.) Grant 
Ilapel, Coquimbo, Chile, altitude 4,000 feet; Plant Introduction Service 
no. 144534, USDA (5041). 

n=46 M. pilosiusculus HBK. 
Botanic Garden, Copenhagen, Denmark (wild in Argentina, Chile, and 
Peru) ; Plant Introduction Service no. 181130, USDA (5320). 


contained 15, one contained 13 and two contained 16. Thus, in the Mather 
cultures irregular meioses occurred in more than 50 per cent of the micro- 
sporocytes and a comparable proportion of aneuploid microspores was 
produced. If the resulting aneuploid gametes are functional, even occa- 
sionally, they might lead to the formation of aneuploid plants or popula- 
tions. If such gametes are generally non-functional, they might help to 
explain the marked self and cross sterility observed in the Mather cultures 
in comparison to the relatively high self and cross fertility of other cul- 
tures of WM. guttatus (Vickery, in press). 

Mimulus glaucescens (5653) is morphologically closely related to M. 
guttatus (Pennell, 1951). It has n=14 chromosomes which are indistin- 
guishable in appearance from those of M. guttatus (fig. 1). This investi- 
gation revealed no cytological basis for the strong crossing barrier (Vick- 
ery, 1956) that separates M. glaucescens from M. guttatus and its related 
species. 

Mimulus tilingit var. tilingii (culture 5967) from a population growing 
on Mount Timpanogos of the Wasatch Mountains, Utah, generally has 
n=15 chromosomes in contrast to other Utah and California populations 


60 MADRONO [ Vol. 15 


2005 Spisn os) 5 01019 5011.0 

oN OTs S87 SF @@39 SOU 

9 67 5 Ot 5009 50110 
| ean [aE | 


O lO 2@OMICRA 


Fic. 1. Meiotic chromosomes of North American Mimulus: M. guttatus, 5003, 
5007, 5009, 5010, 5839; M. glaucescens, 5653; M. tilingit var. tilingii, 5967, var. coral- 
linus, 5011. All cells are in or near second metaphase except 5007, 5011, and 5967 
which are in first metaphase. (Camera lucida drawings, x 645.) 


of that variety which have n—14 (Vickery, 1955 and unpublished). Vari- 
ous stages of meioses were examined in 16 microsporocytes from three 
different plants of culture 5967. Two of the cells contained 13 and 14 
chromomoses instead of the more prevalent 15. However, M. tilingii var. 
tilingu did not show irregular numbers nearly as frequently as did the 
Mather populations of M. guttatus. 

Mimulus tilingti var. corallinus (culture 5011) is closely related mor- 
phologically to M. guttatus and to M. tilingii var. tilingu (Grant, 1924) 
but is effectively separated from them by genetic barriers (Vickery, 1956). 
Culture 5011 forms sterile hybrids with W/. guttatus and will not hybri- 
dize, despite numerous attempts, with W. tilingi var. tilingu. Mimulus 
tilingu var. corallinus has n=24 chromosomes in contrast to the n=14 
of M. guttatus and the n=14 and n=15 of M. tilingi var. tilingi. Fur- 
ther work is in progress to try to establish the chromosome homologies 


1959] MUKHERJEE AND VICKERY: MIMULUS 61 


5056 5041 5320 


a . 
ea 
8 
or) 


e® 
ead » 20e8 
e® 


& 


Fic. 2. Meiotic chromosomes of South American Mimulus: M. tigrinus, 5056 (first 
metaphase) ; M. glabratus var. parviflorus, 5041 (second metaphase) ; M. pilosius- 
culus, 5320 (second metaphase). (Camera lucida drawings, x 1134.) 


and the genetic relationships of the various entities within the Mimulus 
tilingu complex. 

The three species from South America used in this study have higher 
chromosome numbers than any of the North American forms determined 
thus far. Mimulus glabratus var. parviflorus (culture 5041) from the 
Andes Mountains of Chile has n=45 chromosomes and M. pilosiusculus 
(culture 5320), originally from southern South America, has n=46 (fig. 
2). Mimulus tigrinus hort., a cultivated derivative of M. luteus L. (Miller 
and Bailey, 1947) has n=32 chromosomes (fig. 2) which agrees with the 
report of Brozek (1932). 

In conclusion, this survey of chromosome numbers and behavior in sec- 
tion Szmiolus indicates that in general the chromosome number for M. 
guttatus is n=14. However, aneuploid microspores are produced with a 
frequency of greater than 50 per cent in the two cultures of M. guttatus 
from Mather. Such microspores, if functional even occasionally, might 
lead to the production of aneuploid plants or populations of M. guttatus 
similar to the aneuploid populations found in MM. tilingi var. tilingu, M. 
glabratus var. utahensis and in the M. glabratus var. parviflorus— 
M. pilosiusculus group. The high chromosome numbers found in the 
South American species indicate that polyploidy, as well as aneuploidy, 
plays an important role in the evolution of species in section Szmtolus. 
Further work is in progress to elucidate the questions of the cytogenetic 
relationships and taxonomic status of several entities in section Szmiolus 
which have been raised by the results of this investigation. 


Department of Genetics and Cytology, 
University of Utah, 
Salt Lake City 12, Utah 


LITERATURE CITED 


BrozEK, A. 1932. Mendelian analysis of the “red-orange-yellow” group of flower 
colors in Mimulus cardinalis Hort. Preslia 11:1—10. 

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


62 MADRONO LVol. 15 


Miter, W. ann L.H. Battery. 1947. in The standard cyclopedia of horticulture by 
L.H. Bailey. MacMillan. Vol. 2, pp. 2054-2055. 

MUKHERJEE, B.B., D. Wiens, AND R.K. VicKEry, JR. 1957. Chromosome counts in 
section Simiolus of the genus Mimulus (Scrophulariaceae). II. Madrofio 14: 
128-131. 

PENNELL, F.W. 1951. im Illustrated Flora of the Pacific States by Leroy Abrams. 
Stanford University Press. Vol. 3, pp. 688-731. 

SWAMINATHAN, M.S., M. L. Macoon, anp K.L. Menra. 1954. A simple propio- 
carmine PMC smear method for plants with small chromosomes. Indian Jour. 
Gentics and Plant Breeding 14:87-88. 

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

1955. Chromosome counts in section Simiolus of the genus Mimulus 
(Scrophulariaceae). Madrono 13:107-110. 

1956. Data on interracial and interspecific hybridization in the section 
Simiolus of the genus Mimulus (Scrophulariaceae). Proc. Utah. Acad. Sci., Arts 
and Letters 32:45-64. 

(in press). Barriers to gene exchange within Mimulus guttatus (Scrophu- 
lariaceae). 


REVIEW 


Pollen and Spore Morphology/Plant Taxonomy. Gymnospermae, Pteridophyta, 
Bryophyta (Illustrations). (An Introduction to Palynology. II). By Gunnar Erpt- 
MAN. 151 pp., frontispiece, 5 plates, 265 figs. Almquist & Wilsell, Stockholm. 1957. 
$8.00. 

Over a period of more than 13 years, Gunnar Erdtman has undertaken the volu- 
minous task of describing and illustrating representative pollen and spores of the 
world’s plants. His contribution toward better understanding of the fundamentals of 
microspore morphology has given world-wide impetus to the development of this 
aspect of plant morphology and to palynology. 

Erdtman’s earlier publications were largely concerned with the pollen morphology 
of the more common angiosperms and gymnosperms in the experience of the Pleis- 
tocene pollen-analyst. Pollen workers, 10-15 years ago, were generally satisfied with 
knowing the gross morphologic features and key characteristics of pollen of the 
common wind-pollinated genera. Within the past few years, however, the boundaries 
of pollen work have been vastly expanded. The need for a thorough understanding 
of the pollen morphology of living plants has become increasingly apparent in identi- 
fication and interpretation of fossil pollen, as well as a basis for the application of 
pollen morphology to systematic studies. Need for clarification of many of the details 
of microspore morphology and knowledge of pollen and spores of increasing num- 
bers of plants has been answered in part by two volumes recently published by Erdt- 
man. “An Introduction to Palynology. I. Pollen Morphology and Plant Taxonomy. 
Angiosperms” appeared in 1952. The volume being reviewed, ‘An Introduction to 
Palynology. II.”, comprises the illustrations to the text of a treatise (Vol. III) on the 
morphology of microspores of the gymnosperms, pteridophytes and bryophytes which 
will be published at a later date. 

Volume II includes “palynograms” (diagrammatic drawings showing the gross 
morphology of the grains as well as details of the surface pattern and exine strati- 
fication), a few photomicrographs and some electron micrographs of thin sections 
through spore walls of representatives of 12 gymnosperm families, 29 pteridophytic 
and 63 bryophytic families (23 Hepaticae and 40 Musci). Also included are similar 
illustrations for the surface pattern of, and optical sections through, the megaspore 


1959 | NOTES AND NEWS 63 


membrane of a few members of the Cupressaceae, Pinaceae and Podocarpaceae among 
the gymnosperms and the Isoetaceae, Pilulariaceae and Selaginellaceae among the 
pteridophytes. The illustrations are arranged in sections dealing with each of the 
major groups: Gymnospermae, Pteridophyta and Bryophtya. Within each section the 
figures are placed alphabetically by genus. Reference to the family to which each 
genus belongs is made only at the beginning of the section. Spore palynograms of the 
Hepaticae and Musci are ordered alphabetically without regard to their class within 
the Bryophyta. This treatment makes for much clumsiness in use of the book by other 
than one well informed with the taxonomy of each group, and is the most serious 
fault one may find with this publication. Part of the difficulty could have been over- 
come by cross-referencing all genera to family. An arrangement in conformity with 
an acceptable taxonomic heirarchy would have been far more satisfactory, however, 
in gaining understanding of the overall microspore morphology of a family in terms 
of the representatives illustrated, or in attempting to compare the spore morphology 
of related families. One other less serious criticism which might be leveled at this 
publication is that concerning the number of figures and plates (3 out of 5) which 
have appeared already in other publications which most palynologists and others 
interested in microspore morphology would have seen. Some 17 figures and plates, 
in large part, or the only illustrations for Abzes, Cedrus, Ephedra, Keteleeria, Picea, 
Pinus, Pseudolarix, Lycopodium, Ophioglossum, members of the Hedwigiaceae, 
Schizaeaceae and Marattiales, have already appeared in one or both of two periodi- 
cals in 1954 and 1956: the Svensk Botanisk Tidskrift and Grana Palynologica. We 
might suggest that this material could have been supplemented by additional data 
for these genera or groups in the present publication which purports to be a survey 
of a large group of plants. While not serious, numerous other obvious errors impair 
slightly the usefulness of this publication. There is no reference to illustrations of 
members of the Podocarpaceae (Gymnospermae) other than to one figure of the 
megaspore membrane of Dacrydium cupressinum. Microspores of six podocarpaceous 
genera are figured. Family references have also been omitted from the lists prefacing 
sections on Pteridophyta and Bryophyta for Negripteris incana (fig. 148, p. 81); 
Oleandra neriiformis (fig. 150, p. 81); Athalamia nana (fig. 196, p. 101); Brachio- 
lejeunia sandwicensis (fig. 198, p.102) ; and Southbya stillicidiorum (fig. 245, p. 121). 
Other errors include absence of page numbers for some figures, or mistakes in page 
numbers for figures and plates. 

A brief Introduction discusses exine morphology and the nature of the bladder 
among the winged (or saccate) gymnosperms (members of the Pinaceae and Podocar- 
paceae). Some of the terminology used was introduced and defined in the earlier, 
1952, publication. Other terms (mesosaccia, aposaccia, cristae marginales, etc.) are 
apparently newly introduced further to confuse the already over-termed pollen mor- 
phologist. A twenty-two page supplement, containing technical articles by B. M. 
Afzelius and J. Radwan Praglowski on electron microscopy and cutting ultra-thin 
sections as an aid to study of exine stratification, completes this publication. Praglow- 
ski’s article is simply and well presented and well illustrated, and is very welcome 
to those desiring to undertake the sectioning of pollen grain exines. 

It is unfortunate that Vol. II has been published without the proposed accom- 
panying text. The appearance of Vol. III, hopefully in the near future, will be awaited 
with interest——JANE Gray, Geochronology Laboratories, University of Arizona, 
Tucson. 


NOTES AND NEWS 


THE OCCURRENCE OF PILOSTYLES THURBERI (RAFFLESIACEAE) IN CALIFORNIA. In 
various articles and manuals relating to the vegetation of Arizona and California the 
suggestion has been made that Pélostyles thurberi Gray may occur in the desert areas 
of southern California. Never, as far as I am aware, has a documentation of this 


64 MADRONO [Vol. 15 


range extension been published. The plant, parasitic on the branches of Dalea emoryi 
Gray, seems to be an exceedingly elusive one, as it is rarely collected. Only the minute 
flowers erupt from the host bark, the vegetative body being completely internal. 

It is therefore worthwhile to confirm its occurrence in California. The following 
specimens, all from the Colorado Desert region of California, leave no doubt as to 
the presence of the parasite in the state: Goodding 43-19, 57 miles south of Indio 
(UC 690661), Goodding, 50 miles south of Indio (UCLA), Wilson, 30 miles west of 
El Centro (POM 273347), Wilson, near Ocotillo (RSA 43067). All these collections 
were on Dalea emoryi. On a field trip in 1958 I was unable to locate the parasite in 
any of the California localities. Because of agricultural developments it may not have 
survived in some of its known California and Arizona sites (perhaps including the 
type locality in Yuma County, Arizona). It still exists at Wellton, Yuma County, 
where it grows along the highway (Kuwijt 1490, UC). 

Pilostyles thurberi is the only species of the Rafflesiaceae in California. The other 
species of Pilostyles show an extraordinary geographical distribution. Some are found 
in Texas and northern Mexico, others in Chile, in Iraq, in Ethiopia, or in southwest 
Australia. The hosts are invariably leguminous shrubs growing in desert areas—J OB 
KuiytT, Department of Botany, University of California, Berkeley. 


NOTES ON THE FLORA OF ARIZONA.—Four interesting collections have come to the 
herbarium of the University of Arizona for identification during the past year. Of 
these two are genera new to the state, and two represent second collections of species 
rare in Arizona. Only one is a foreign weed. 

Echinodorus berteroi (Spreng.) Fassett (Alismaceae) was collected 1% miles 
north of Imperial Dam in the Imperial National Wildlife Refuge, Yuma County, by 
Gale Monson. It is a new generic record for Arizona, but one that might be expected. 
Echinodorus berteroi ranges from southern California eastward to the West Indies, 
but, according to Fassett’s citation of specimens (Fassett, N.C. 1955. Echinodorus 
in the American Tropics. Rhodora 57:133-212), this is the first collection from the 
Colorado River as well as the first from Arizona. 

Cardiospermum halicacabum L. (Sapindaceae), also a genus new to Arizona, was 
collected by L. N. Goodding and Richard Hevly on a rocky slope in the Coyote 
Mountains, Pima County. This new locality is about 50 miles north of the Inter- 
national Boundary, while the closest previous collection represented in the University 
of Arizona Herbarium is near Hermosillo, Sonora, Mexico, about 160 miles south of 
the border. Although C. halicacabum is cultivated as the “balloon vine,” the rough, 
rocky area in which this specimen was growing precludes the possibility that it was 
an escape. 

Beckmannia syzigachne (Steud.) Fernald (Gramineae), collected near Alpine, 
Apache County, was sent to the University of Arizona for identification by C. L. 
Isaacson, the county agricultural agent. This collection is the second for the state; 
the first having been made in 1951 by John Merkle at Greenland Lake, Grand Canyon 
National Park, Coconino County (Merkle, J. 1953. Beckmannia Syzigachne (Steud.) 
Fernald, New Record for Arizona. Madrono 12:32). The new collection is about 200 
air miles southeast of the first. 

Bupleurum rotundifolium L. (Umbelliferae) was brought to the herbarium for 
identification by a resident of Tucson, Pima County. It was growing in an unseeded, 
unfertilized section of a yard under an established elm tree (Ulmus sp.). This, also, 
is a second collection, the first having been made in Tucson by Toumey in 1892. Un- 
doubtedly the new collection is a re-introduction of this Mediterranean weed, but 
the seed source is unknown. 

Representatives of all of these collections are on file in the Herbarium of the Uni- 
versity of Arizona—CHarLes T. Mason, Jr., Department of Botany, University of 
Arizona, Tucson. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree to bear the cost of the ad- 
ditional pages at the rate of $20 per page. Illustrative materials (includ- 
ing “‘typographically difficult”’ matter) in excess of 30 per cent for papers 
up to 10 pages and 20 per cent for longer papers are chargeable to the 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
the author assume the complete cost of publication. 

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

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


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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume I,1916-1929 . . . . . $5.00 
Volume IT, 1930-1934. . . .~ . 5.00 
Volume ITI, 1935-1936 . . .~ . 5.00 
Volume IV, 1937-1938 . . .. . 5.00 
Volume V, 1939-1940. . . .) . 5.00 
Volume VI, 1941-1942 . . . . 5.00 
Volume VII, 1943-1944 . . .. ., 5.00 
Volume VIII, 1945-1946 . . ..) . 5.00 
Volume IX, 1947-1948 Ry ES 6.00 
Volume X, 1949-1950. . . .. . 7.00 
Volume XJI,1951-1952 .  . wi, 8.00 
Volume XIT, 1953-1954 . . . 8.00 
Volume XIII, 1955-1956. . . . 8.00 
Volume XIV, 1957-1958 . . .. . 8.00 
Single numbers, 

Volumes I and 2.2) 3.000.) 0 2510) 
Succeeding numbers, 3 

(except Volume XI, No.2). . . 1.00 

Volume XI, No. 2.0.0 7s) ees 1.50 


The subscription price of MADRONo is $4.00 per year. If your 
institution does not now subscribe to MApDRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


ew 
VOLUME 15, NUMBER 3 JULY, 1959 


Contents 


PAGE 


Factors AFFECTING THE DISTRIBUTION OF PONDEROSA 
AND JEFFREY PINES IN CALIFORNIA, John R. Haller 65 


VIVIPARY IN CORDYLINE AUSTRALIS HOoK., 
Howard J. Arnott 71 


STUDIES ON SECOTIACEOUS FUNGI VI. SETCHELLIO- 
GASTER Pouzar, Rolf Singer and Alexander H.Smith 73 


CEANOTHUS SEEDS AND SEEDLINGS ON BURNS, 
Clarence R. Quick 79 


CHROMOSOME NUMBERS OF CALIFORNIA PLANTS, WITH 
NoTES ON SOME CASES OF CYTOLOGICAL INTEREST, 


Richard Snow 81 
CHROMOSOME COUNTS IN THE GENUS GAYOPHYTUM, 
David G. Dixon 90 


Reviews: Clara Chapman Hill, Spring Flowers of the 
Lower Columbia Valley (Robert Ornduff) ; Lloyd H. 
Shinners, Spring Flora of the Dallas-Fort Worth Area, 


Texas (Rogers McVaugh) 94 
Notes AND NEws: ALLIARIA OFFICINALIS ANDRZ. IN 
OrEGON, Robert Ornduff 96 


A WEST AMERICAN JOURNAL OF BOTANY > Pak 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


ANY HSONgs 


FL) Ses) 


— LIBRARY 


~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 

$4.00 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. Davipson, University of Nebraska, Lincoln. 

IvAN M. JounstTon, Arnold Arboretum, Jamaica Plain, Massachusetts. 
Mitprep E. Martuias, University of California, Los Angeles 24. 
MARION OWNBEY, State College of Washington, Pullman. 

Tra L. Wiccrns, Stanford University, Stanford, California. 


Secretary, Editorial Board—ANNETTA CARTER 


Department of Botany, University of California, Berkeley. 


Business Manager and Treasurer—WINSLOW R. Briccs. 
Department of Biology, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: James R. Sweeney, San Francisco State College, San Francisco, Cali- 
fornia. First Vice-president: Baki Kasapligil, Mills College, Oakland, California. 
Second Vice-president: Henry J. Thompson, Department of Botany, University of 
California, Los Angeles, California. Recording Secretary: Mary L. Bowerman, De- 
partment of Botany, University of California, Berkeley, California. Corresponding 
Secretary: Francia Chisaki, Department of Botany, University of California, Berke- 
ley, California. Treasurer: Winslow R. Briggs, Department of Biology, Stanford 
University, Stanford, California. 


1959] HALLER: PONDEROSA AND JEFFREY PINES 65 


FACTORS AFFECTING THE DISTRIBUTION OF PONDEROSA 
AND JEFFREY PINES IN CALIFORNIA 


JoHN R. HALLER 


Pinus ponderosa Doug). and P. jeffreyi Murr. are among the most wide- 
ly distributed forest trees in California and are well known to western 
botanists. They are very closely related, and various authors have treated 
them either as species (Sudworth, 1908; McMinn, 1951) or as varieties 
of a single species (Shaw, 1914; Jepson, 1925). However, after extensive 
field investigations (Haller, 1957) the details of which will be published 
elsewhere, I am convinced that P. ponderosa and P. jeffreyi are well de- 
fined species, and I shall treat them as such in this paper. All of the obser- 
vations which follow are my own, unless otherwise noted, and were made 
in connection with the taxonomic study referred to above. 

In California, Pinus ponderosa occurs from the Oregon border south 
along the Cascade Range and the western slope of the Sierra Nevada in 
an uninterrupted belt over 400 miles long and about 25 miles in width. 
Throughout this belt P. ponderosa is a conspicuous element in a forest 
rich in coniferous species. It is also common on the higher coastal-facing 
slopes of the mountains of southern California as far south as Cuyamaca 
Lake in San Diego County, and occurs sporadically in the Coast Ranges, 
especially north of San Francisco Bay. 

The distribution of P. jeffreyi roughly parallels that of P. ponderosa 
in California, but P. jeffrevi is relatively more abundant in the south, and 
extends beyond the range of P. ponderosa into the higher mountains of 
northern Baja California, Mexico. 

Although the ranges of P. ponderosa and P. je ffreyi in California near- 
ly coincide in the broadest geographical sense, and although both are 
often found at the same locality, the two species characteristically occupy 
different habitats. Pinus ponderosa occupies the lower coastal-facing 
slopes of the mountains, whereas P. jeffreyi is usually found on the higher 
coastal or desert-facing slopes. The altitudinal ranges of P. ponderosa and 
P. jeffreyi shift gradually higher from north to south in California, just as 
do vegetation zones in general. However, the degree of shifting is not the 
same in both species. The lower limit of the P. ponderosa zone rises steep- 
ly from north to south. In the north, near Mount Shasta, the lower limit 
of P. ponderosa is about 1000 feet, but in the south, near Barton Flats in 
the San Bernardino Mountains, it is at 5000 feet (fig. 1). The upper limit 
of the P. ponderosa zone does not rise as steeply. It changes from about 
5000 feet in the north to 7000 or 7500 feet in the south. Therefore, the 
altitudinal range of P. ponderosa contracts from about 4000 feet near 
Mount Shasta to little more than 2000 feet at Barton Flats. South of the 
Barton Flats area P. ponderosa becomes rapidly more infrequent in the 


Maprono, Vol. 15, No. 3, pp. 65-96. July 20, 1959. 


66 MADRONO [Vol. 15 


montane forest, until the most southerly stand is reached at Cuyamaca 
Lake, at the unusually low altitude of 4600 feet. 

The lower edge of the P. jeffrey zone rises from about 5000 feet near 
Mount Shasta to only a little over 6000 feet near Barton Flats (fig. 1). 
The zone in which P. ponderosa and P. jeffreyi overlap is consequently 
broader in the south than in the north, and stands comprised of both 
species are extensive near Barton Flats but highly restricted near Mount 
Shasta (Wiggins, 1940 and my own observations).! However, there are 
numerous localities throughout the state such as Shasta Valley in Siskiyou 
County and Pine Valley in southern San Diego County where P. jeffreyi 
occurs much lower, either with or without P. ponderosa, in what would 
usually be the lower portion of the P. ponderosa zone. The upper limit of 
P. jeffreyi rises from about 7500 feet near Mount Shasta to 9500 or 10,000 
feet near Barton Flats. Thus the normal altitudinal range of P. jeffreyi 
increases from about 2500 feet in the north to 3500 or 4000 feet in the 
south. 

What are the factors that cause this narrowing of the altitudinal range 
of P. ponderosa and broadening of the altitudinal range of P. jeffrey in 
the south? There is both experimental and observational evidence which 
indicates that moisture at the lower altitudinal limit and temperature at 
the upper limit are usually the critical factors. Daubenmire (1943) has 
shown experimentally that seedlings of several important species of Rocky 
Mountain conifers, including P. ponderosa, are capable of withstanding 
much higher soil temperatures than those in their natural environments, 
but that they quickly succumb to drought conditions more severe than 
those in their natural habitats. In most areas of California also, P. pon- 
derosa is probably restricted at its lower limit by moisture rather than 
temperature or some other factor. Pinus ponderosa is rarely found in areas 
with less than 25 inches of annual precipitation except near permanent 
sources of water. Of course moisture alone may not always be the limit- 
ing factor at low elevations. For example, in some localities where precipi- 
tation is still adequate but almost limiting, P. ponderosa might be held in 
check by its inability to compete with the more xeric chaparral vegeta- 
tion. Billings (1950) has described the interesting occurrence of P. pon- 
derosa, P. jeffreyi and other montane coniferous species on chemically 
altered andesitic soils in western Nevada. The climate of the area is too 
arid for these species under normal soil conditions, and supports only 
sagebrush or pifon-juniper vegetation. However, the sagebrush and most 
other shrubs cannot tolerate the chemically altered soils, and Billings has 
concluded that the absence of shrubby competitors enables the pines to 
persist. 

The upward migration of P. ponderosa is, in all probability, usually 


1 Protessor G. Ledyard Stebbins (personal communication) has observed exten- 
sive mixed stands of P. ponderosa and P. jeffreyi in the North Coast Ranges. The 
stands in this area apparently comprise an exception to the usual pattern of restricted 
mixed stands in the north and extensive ones in the south. 


1959] HALLER: PONDEROSA AND JEFFREY PINES 67 


is 
KOSS 
42s 


cI Sirgen CA Z 


10,000 ft. 


8,000 ft. 


6,000 ft. 


4,000 ft. 


2,000 ft. NN Ae 
As 
S 
SEA ed 
MT.SHASTA 0 100Miles YOSEMITE BARTON FLATS MEXICAN 
(41°N.) Horizontaliscale (37°45’N) (34°N.) BOUNDARY 


Fic. 1. Altitudinal distribution of Pinus ponderosa and P. jeffreyi along a north- 
south transect in California. 


checked by low temperature. Californian P. ponderosa is killed by freez- 
ing when grown in areas with low winter temperatures, such as the 
Rocky Mountains (Weidman, 1939). I have observed frost damage on 
young individuals of P. ponderosa in California after periods of severe 
cold. The limiting effect of the low temperature undoubtedly works on the 
seedlings or young trees, because mature stands of P. ponderosa at its 
upper altitudinal limit are vigorous and show no evidence of being stunted. 

Assuming that moisture and low temperature are the principal limiting 
factors for P. ponderosa at its lower and upper limits respectively, it is 
apparent from climatic data why P. ponderosa has a narrower altitudinal 
range in the south than in the north. As one travels south in California, 
precipitation generally decreases more rapidly than temperature increases 
at any given altitude (U.S. Weather Bureau, 1958). In other words, the 
isohyets gain altitude faster than the isotherms toward the south, leaving 
an ever shrinking zone that is favorable to the growth of P. ponderosa. It 
might be wondered, if the above is true, why P. ponderosa stops so 
abruptly in San Diego County rather than continuing toward the south 
over a decreased altitudinal range. There is no abrupt environmental shift 
at Cuyamaca Lake, and there are sites farther south that appear to be 
capable of supporting P. ponderosa. The reasons for the relatively sharp 
southern limit of the species may be historical. For example, a severe 
drought could have eliminated P. ponderosa south of where it occurs now, 
and the species may not yet have had sufficient opportunity to expand 
and reach its former extent. 

Pinus jeffreyi is probably limited at the upper margin of its altitudinal 
range by a complex of factors all basically caused by low temperatures. 
Individuals growing near the upper limit of the species are often severely 
stunted in appearance, and obviously are showing the effects of the harsh 


68 MADRONO [Vol. 15 


climate. In addition, the upper limit of P. jeffrevz almost exactly parallels 
that of . ponderosa from north to south in California, suggesting that 
different degrees of the same factors are limiting both species at the upper 
margins of their altitudinal range (fig. 1). 

The lower altitudinal limit of P. jeffreyi does not appear to be deter- 
mined by any obvious physical factors, but rather by competition with 
P. ponderosa or other species. Pinus ponderosa grows faster under culti- 
vation than P. jeffreyi in many diverse localities, from the Institute of 
Forest Genetics at Placerville, California (within the range of P. ponde- 
rosa), to England (Hooker, 1884). Near its upper limit P. ponderosa may 
not grow much faster than P. jeffreyi and thus permit the occurrence of 
mixed stands, but lower down, the more vigorous growth of P. ponderosa 
could exclude P. jeffreyi since both species have about equally high light 
requirements (Sudworth, 1908). The much greater extent of stands con- 
taining both species in southern California than farther north might be 
due to a slightly lowered vigor and subsequent lessening of the competi- 
tive ability of P. ponderosa in the south. In relatively dry southern Cali- 
fornia, P. ponderosa is in an environment where moisture is more likely 
to be limiting than in the north, and therefore might be expected to be 
less vigorous. The occurrence of P. jeffreyz in areas where P. ponderosa 
is lacking is further evidence of the limitation of P. jeffreyi by competi- 
tion. In southern San Diego County, beyond the southern limit of P. pon- 
derosa, P. jeffreyi often occurs through the entire altitudinal range of the 
montane forest. A short distance to the north, the lower portions of this 
forest are occupied by P. ponderosa instead. In the Sierra Nevada there 
are numerous sites within the montane forest, often at very low altitudes, 
such as sandbars in rivers or areas of serpentine soil, where P. ponderosa 
occurs infrequently if at all. The less demanding P. jeffreyi, however, is 
frequently encountered on these sites. 

Revealing evidence concerning the distribution limits of P. ponderosa 
and P. jeffreyi has been obtained from east-west transects across the 
mountains as well as by north-south transects along their axes. As an 
example, I shall describe a west to east transect across the Sierra Nevada 
in the vicinity of Yosemite National Park in central California. The west- 
ern and eastern slopes of the central portion of the Sierra Nevada display 
a striking series of contrasts, both topographically and climatologically. 
The western slope is fairly gradual, receives from 30 to 60 inches of pre- 
cipitation annually in the montane forest belt (U.S. Weather Bureau, 
1958), and has a relatively mild climate characterized by temperature 
extremes that are not pronounced. The eastern slope, on the other hand 
is very precipitous, receives only 10 to 20 inches of annual precipitation 
in the montane forest belt because of the rain shadow effect produced by 
the 13,000 foot crest that lies to the west, and has a continental climate 
with great extremes of temperature. 

At the latitude of Yosemite (38° N.), the lowest continuous stands of 
P. ponderosa appear at an elevation of about 3000 feet, where the mean 


1959] HALLER: PONDEROSA AND JEFFREY PINES 69 


A Ponderosa 
12900 ft. Jeffrey [\ 
10,000 ft. a f 
ELLY 7, Mommoth Lokes\ 
8,000 ft. ay Ni A Bey Porcupine Fiat 


6.000 ft. 
\t\ \ \ My Rock Creek \Y 
4,000 ft. [YN Yosemite Valle 
Wy, 
2,000 ft. 
Sea Level 


Fic. 2. Altitudinal distribution of Pinus ponderosa and P. jeffreyi along an ideal- 
ized west-east transect across the central Sierra Nevada. Open triangles represent 
P. ponderosa; solid triangles represent P. jeffreyi. 


annual precipitation is at least 30 inches. At about 6000 feet, P. jeffreyi 
is first found in abundance. The stands of P. jeffreyi growing at this rela- 
tively low altitude are usually not scattered at random over the slopes, 
however, but are concentrated near the margins of meadows or along 
streams, together with Pznus contorta subsp. murrayvana Engelm., which 
occasionally occurs below its usual range in such places. Pinus ponderosa 
remains abundant for several hundred feet above the low places where 
P. jeffrey is first encountered, especially on south-facing slopes, before 
giving way to P. jeffreyi. Above 6500 or 7000 feet, P. jeffreyvi occurs alone 
and continues up to an elevation of approximately 9000 feet (fig. 2). 

If the transect is continued across the 13,000 foot crest to the more 
arid eastern slope of the Sierra Nevada, P. jeffreyi will again be found 
below the 9000 foot elevation. Below 7000 or occasionally 6000 feet, 
where the annual precipitation may be as low as 10 inches, P. jeffreyi 
gives way to more xeric species, such as Pinus monophylla Torr. and 
Artemisia tridentata Nutt. Pinus ponderosa is not at all common on the 
eastern slope, and occurs without exception only along the banks of a few 
perennial streams. It is most abundant along Rock Creek near the Inyo- 
Mono county line at altitudes ranging from 5000 to 6500 feet (fig. 2). 

The distribution pattern of P. ponderosa and P. jeffrevi along this west 
to east Sierran transect appears to be the result of the same limiting fac- 
tors as those suggested by their north-south distribution. Although the 
lowest extensive stands of P. jeffreyi on the western slope of the Sierra 
Nevada are usually near meadows or along streams, the trees are prob- 


70 MADRONO [Vol. 15 


ably not there because they need the moisture, but rather because P. jef- 
frevi is more tolerant of cold than is P. ponderosa. These low pockets 
where P. jeffreyi occurs are colder than the surrounding slopes in winter. 
The occasional presence of the usually subalpine P. contorta subsp. mur- 
rayana in these pockets is further evidence of this fact. Probably P. jef- 
freyi merely tolerates the extra moisture present at its lowest localities, 
because, as already noted, the species occurs in much drier localities in 
other areas. In the harsh climate of the eastern slope of the Sierra Nevada, 
P. ponderosa survives only at low elevations where it is not too cold and 
along streams where it is not too dry. Where P. ponderosa and P. jeffreyi 
are found in the same general area on the eastern slope, the latter, in 
contrast to P. ponderosa often occurs both near the streams and on the 
adjacent slopes. 

Pinus ponderosa is found in abundance on both sides of the Sierra crest 
from Lake Tahoe northward. However, in this region the crest of the 
range is much lower than farther south, and the environmental differences 
between the western and eastern slopes are not nearly as great. On the 
desert-facing slopes of the higher mountains of southern California, P. 
ponderosa is again restricted to unusually moist localities. 


SUMMARY 


Pinus ponderosa and P. jeffreyi are important components of the Cali- 
fornia montane forest. Pinus ponderosa generally occurs on the lower 
coastal slopes, while P. jeffreyi generally occurs on the higher coastal or 
interior slopes. The distribution of P. ponderosa is usually checked by 
lack of moisture at low altitudes and by low temperature at high altitudes. 
Pinus jeffreyt is also limited by low temperature at high altitudes, but at 
its lower margin the distribution of the species is apparently limited pri- 
marily by competition with P. ponderosa. The extensive mixed stands of 
P. ponderosa and P. jeffreyi in southern California where P. ponderosa 
may be less vigorous than in the north, might be the result of decreased 
competition from P. ponderosa. Pinus jeffreyi is more tolerant of extremes 
of low temperature and aridity than is P. ponderosa, and is at least equal- 
ly as tolerant as P. ponderosa of extremes of high temperature and high 
moisture. 


Department of Biological Sciences, 
University of California at 
Santa Barbara 


LITERATURE CITED 


Briiincs, W. D. 1950. Vegetation and plant growth as affected by chemically altered 
rocks in the western Great Basin. Ecology, 31:62-74. 

DAUBENMIRE, R. 1943. Soil temperature versus drought as a factor determining lower 
altitudinal limits of trees in the Rocky Mountains. Bot. Gaz., 105:1-13. 

Hatter, J. R. 1957. Taxonomy, hybridization and evolution in Pinus ponderosa and 
P. jefireyi. Doctoral dissertation, Univ. Calif., Los Angeles Library. 

Hooker, J. D. 1884. Pinus jeffreyi. Gard. Chron., Dec. 27, 1884: p. 814. 


1959] ARNOTT: CORDYLINE AUSTRALIS 71 


Jepson, W.L. 1925. A manual of the flowering plants of California. Berkeley. 

McMinn, H. E. and E. Marno. 1951. An illustrated manual of Pacific Coast trees. 
University of California Press, Berkeley. 

SHAW, G.R. 1914. The genus Pinus. Publ. Arnold Arboretum, No. 3. 

SupworTH, G.B. 1908. Forest trees of the Pacific slope. U.S. Government Printing 
Office, Washington, D.C. 

UniTep STATES WEATHER BurREAU. 1958. Climatological data. 1957 annual summary 
for the state of California. Vol. 61, no. 13. 

WEIDMAN, R. H. 1939. Evidences of racial influence in a 25-year test of Ponderosa 
Pine. Jour. Agr. Res., 59:855-887. 

Wiccrns, I. L. 1940. Yellow pines and other conifers observed in Lower California. 
Jour. N. Y. Bot. Gard., 41:267-269. 


VIVIPARY IN CORDYLINE AUSTRALIS HOOK. 
Howarp J. ARNOTT 


Vivipary is defined by Jackson (1928) as “*.... germinating or sprout- 
ing from seed or bud, while attached to the parent plant.”” Examples of 
vivipary are known in a number of genera of both the monocotyledons 
and dicotyledons. The classical example of this condition occurs in such 
mangroves as Rhizophora mangle Blanco. In this species when the seed 
germinates while still attached to the parent plant, the hypocotyl-radical 
elongates, forming a long sharp structure (Daubenmire, 1947, fig. 10, 
p. 64). When this seedling structure becomes heavy enough, it breaks 
away from the paient plant and drops into the mud below. Because this 
sharp hypocotyl-radical structure penetrates the mud the seedling often 
becomes anchored and is prevented from being washed away from its 
environment, especially in the intertidal zone. 

In the Agavaceae of Hutchinson, of which Cordyline is one member, 
several genera have been reported to show vivipary. Both Agave and 
Furcraea are included in this category. The viviparous condition in Fur- 
craea is a great deal different than that of Rhizophora or Cordyline. In 
Furcraea at certain points along the inflorescence bulbils are formed. 
These bulbils (aerial deciduous buds) consist of a series of papery and 
photosynthetic bud scales surrounding a short axis and a shoot apex. 
These structures are often formed in enormous numbers and literally 
cover the ground when they abscise from the parent inflorescence. Bulbils 
begin to grow immediately when proper conditions prevail; plants pro- 
duced in this manner have a very rapid rate of early growth. 

In a cultivated plant of Cordyline australis Hook. growing in Berkeley, 
California, many cases of vivipary were observed. The bright-green young 
seedlings were easily seen protruding out of the white fruits. A total of 
over fifty separate fruits were found exhibiting this character. 

In most cases the cotyledonary arch and the first leaf were all that 
could be seen of the seedling outside the fruit. Two cases were observed in 


72 MADRONO LVol. 15 


Fic. 1. Fruit of Cordyline australis Hook. showing a viviparous seedling protrud- 
ing through the ruptured fruit wall. The fruit was still attached to the inflorescence. 
The densely stippled areas on the fruit represent the purple pigmented areas which 
normally occur on the white fruit of this species. x 9. 

Fics. 2 and 3. Seedlings dissected from viviparous fruits showing the seed, the 
cotyledon, the first leaf, and the primary root. The primary root is intimately asso- 
ciated with the internal tissues of the fruit (see text). * 4. 


which the seed was pulled free of the fruit by the elongation and straight- 
ening of the cotyledon. Figure 1 shows a fruit in which the “aerial” parts 
of the seedling have protruded through the fruit wall. This protrusion was 
effected by a rupture of the tissues which enclosed the seed. The area sur- 
rounding the rupture seems to be at most only slightly discolored. In some 
cases dissection showed more than one seed to be present inside the fruit, 
but no cases of “multiple-vivipary” were found. 

Figures 2 and 3 show seedlings dissected from ‘‘viviparous”’ fruits. The 
shiny black seed attached to the haustorial cotyledonary tip can be seen 
with the photosynthetic and sheathing parts of the cotyledon extending 
out in an inverted ““U”’. Below the cotyledon is the axis and the primary 
root. 

The most interesting point observed in these cases was the very intimate 
relationship between the internal tissues of the fruit and the primary root 
of the seedlings. This intimacy seems to be caused by the penetration of 
the internal fruit tissues by root hairs. When one tries to dissect these 
seedlings free of the fruit it is almost impossible to separate the primary 


1959] SINGER AND SMITH: SETCHELLIOGASTER Us 


root from the internal tissues. It was not determined whether the root 
hairs grow between cell walls, into intercellular air spaces, or actually pen- 
etrate into the cells. The “fuzzy” appearance of the primary roots in fig- 
ures 2 and 3 is an attempt to show this intimate association of the primary 
roots, root hairs, and internal fruit tissues after dissection. 

One fact which seems to indicate that the root hairs are indeed the 
cause of this intimate association is that the root apex and some short dis- 
tance behind it are completely free from any connection with the fruit 
tissues. This apparently is due to the absence of root hairs on such an 
immature part of the root. 

Viviparous seedlings at a later stage than that shown in figure 1 have 
not been observed im situ. Such seedlings when removed from the fruit 
and placed on moist filter paper in a covered petri dish quickly show the 
production of new roots from the hypocotyl region. Apparently if these 
seedlings were planted they would produce normal plants. 

While a number of cases of vivipary were found in this Cordyline plant, 
the number would be less than 0.1 per cent of the total number of fruits 
on the plant. One wonders what special physiological conditions were 
present in these viviparous fruits which caused or allowed the germination 
of these seeds. Also in the cases where more than one seed was present in 
a viviparous fruit, one wonders why only one seed germinated. One final 
question would be whether this condition occurs in this species in its native 
New Zealand, and what possible adaptive value might be found there for 
this condition, if it does occur. 


Department of Biological Sciences, 
Northwestern University, Evanston, Illinois 


LITERATURE CITED 


DAUBENMIRE, R. F. 1948. Plants and environment. John Wiley & Sons, Inc., New 
York. 
Jacxson, B.D. 1950. A glossary of botanic terms. Hafner Publishing Co., New York. 


STUDIES ON SECOTIACEOUS FUNGI VI. 
SETCHELLIOGASTER POUZAR 


RoLF SINGER AND ALEXANDER H. Sm1TH! 


As a result of studying the types of Secotium tenuipes Setchell, and 
Secotium aurantium Zeller, we believe it is logical to group these two in 
a single genus as designated in our title. They have the following char- 
acters in common: their spores are some shade of rusty ochraceous, have 
an imperfect but often distinctly discontinuous pore-region, are elongate 
in shape, and smooth or ornamented by plugs of material filling canal-like 
passages through the wall. The hyphae bear clamp connections at the 


1 Papers from the University of Michigan Herbarium and the Department of 
Botany, No. 1086, University of Michigan, Ann Arbor, Michigan. 


74 MADRONO [Vol. 15 


cross-walls and the mature gleba does not become pulverulent. The outer 
layer of the peridium is a layer of enlarged to vesiculose cells. The hyme- 
nophoral trama (tramal plates of the gleba) is regular, but in the region 
next to the subhymenium it is composed of characteristically enlarged 
cells often taking the shape of sphaerocysts. 

The subhypogeous habitat, its presence in the forest duff in mild cli- 
mates, the presence of gastroid basidia and spores, and possibly the bright 
color (or the absence of pigment) in the peridium are also features of the 
genus, but these characters do not necessarily separate it too sharply from 
other secotiaceous genera. 

The spore characters and the structure of the outer layer of the peridi- 
um are very distinct from those of Secotium as represented by the type, 
S. gueinzi, and do not allow these species to be placed in any other segre- 
gate of Secotium. The generic name Setchelliogaster was proposed by 
Pouzar for the type species (Secotium tenuipes Setchell) just after we had 
submitted for publication our own account in which a new generic name 
was proposed for the two species treated below; therefore we have had 
to adapt our paper to the use of Pouzar’s name, Setchelliogaster. 


SETCHELLIOGASTER Pouzar, Ceska Mykologie 12:33. 1958. 

Spores varying from light brownish to dark rusty ochraceous, with a 
germ pore which is mostly distinct, more rarely imperfect (or at least 
spore wall at apex of spore partly discontinuous), elongate, smooth or 
with an ornamentation of the type of Metraria insignis (part of spore 
wall conspicuously heterogeneous consisting of a continuous wall through 
which extend minute canals plugged by a resinous substance, these plugs 
appearing as dots or lines or imperfect reticulation on the paler ground 
when spore surface is focussed upon) ; clamp connections present; perid- 
ium covered by an epithelium; hymenophoral trama regular but showing 
some inflated elements and spherocysts near the subhymenium. Peridium 
russet brown or orange or rarely ivory color; gleba brownish; columella 
percurrent; stipe not voluminous. Basidium-spore-configuration of the 
gastromycetoid type. Among thermophilous vegetation, growing subhypo- 
geously in humus under trees. 

Type species of the genus: Secotium tenuipes Setchell. 


SETCHELLIOGASTER TENUIPES (Setchell) Pouzar, Ceska Mykologie 12:34. 
Figs. 1-3. 1958. Secotium tenuipes Setchell, Jour. Mycol. 13:239. 1907. 
Gastrocarp 10-30 mm. tall, and 10-30 mm. broad, subglobose or broad- 

ly ovoid, subumbonate, at the base more or less truncate. 

Peridium membranous in lower (marginal) portion, up to 1 mm. thick 
further up, and rather thick at the point of confluence with the columella, 
glabrous, not viscid, yellow-brown, deep brown (according to Setchell), 
or red-brown (‘Morocco red” R. according to Zeller), usually dehiscent 
from the stipe-columella to expose a narrow ring of gleba. 

Gleba variable in structure, loculate with very regular (in shape and 
position) chambers which become slightly lammellarly extended near the 


1959] SINGER AND SMITH: SETCHELLIOGASTER is 


exposed portion below, in other specimens generally with a gill-like struc- 
ture especially visible in longitudinal sections of the gleba, and then 
consisting of anastomosing plates resembling those of Polyporus alveo- 
larius but not in a regular manner and not permitting the shedding of 
spores, chambers sinuous or equal or completely irregular, gleba decurrent 
on the apex of the stipe-columella but for the most part dehiscent or free 
from the latter, ochraceous brown. 

Stipe variable in length, reaching 20 mm., relatively thin, often bent or 
flattened, more or less concolorous with the pileus, striate, solid, equal 
or slightly attenuate downward, 2—3 mm. thick; columella continuous 
with stipe, percurrent and widened into the upper portion of the peridium, 
sometimes slightly narrowed before reaching the junction, lower portion 
to over half the lower length free from gleba; volva none; veil superior, 
arachnoid, transverse, scanty, evanescent after maturity is reached. Con- 
text fleshy, odorless. 

Spores 14.5-19 & 9.5—12.5 wu, ellipsoid to subovoid, shape in optical 
section slightly more ventricose on the outer line than on the inner one, 
but without a suprahilar applanation or depression, and not so strongly 
asymmetric generally as in agaric spores, deep rusty ochraceous viewed 
in KOH, structure of wall complex at maturity and 1—1.2y. thick. Peri- 
sporium conspicuous and pale ochraceous-tawny. Exosporium heteroge- 
neous, consisting of a continuous wall through which extend minute canals 
plugged by a resinous golden tawny substance, these canals at the surface 
of the perisporium appearing as dots or elongated irregular lines which 
may be fused to form an imperfect inconspicuous reticulation which ap- 
pears tawny on a paler background, under mechanical pressure the com- 
bined exosporium and perisporium tending to separate from the episori- 
um. Episporium tawny and appearing as a thin line. Endosporium interior 
to the episporium, thick and much paler in color than the latter. Spore 
apex complex and peculiar in structure: germ pore generally either poorly 
developed or absent but at times a perforation clearly visible, with a 
tawny-cinnamon, plug-like, thick, heterogeneous (different from wall- 
material) body in the region where the pores should be (abnormal spores 
often with 2—5 such structures), at times showing a slightly mucronate 
apical callus but no distinct perforation (a discontinuity of the wall strata 
may be observed only in the endo- and episporium), not truncate at apex, 
not necessarily germinating through the apex (lateral germination ob- 
served). 

Basidia 30-40 « 7—-10y, (1—) 2— (3—)-spored, hyaline, numerous but 
rarely seen to form a large area of hymenium but rather intermixed with 
very numerous pseudoparaphyses, with a median constriction; sterigmata 
apical, straight or very slightly curved (not typically half-sickle-shaped 
but rather of gastroid type) ; pseudoparaphyses normally vesiculose and 
not projecting beyond the general level of the hymenium, hyaline, some- 
times gigantic (cystidia?), about 32 « 16, generally 20-22 « 13y:; 
true cystidia or pseudocystidia none seen. 


76 MADRONO LVol. 15 


Subhymenium well-developed, subhyaline to hyaline, consisting of 
small irregular elements, some of them subisodiametric, forming a distinct 
if not very thick layer; hymenophoral trama consisting of brownish to 
brownish-hyaline hyphae of very variable diameter, some of them actually 
subisodiametric (e.g. 4-28 u diam.), thin-walled or nearly so, some ele- 
ments incrusted by deep rusty pigment, generally regular but chains of 
broad inflated elements alternating with strands of filamentous narrow 
hyphae, making the trama slightly intermixed, and reminding one of the 
gill trama of Conocybe. Peridial tissue thin, inner layer much like the 
hymenophoral trama, not gelatinous, its hyphae 5—27 ». broad; outer layer 
thick, its elements rusty brown from incrusting pigment, an epithelium or 
pseudoparenchyma consisting of spherocysts in chains or isolated, and 
few filamentous hyphae ending up (as a terminal member) in a sphero- 
cyst (spherocysts 8—28 (50) & 8-22.) hyphae of all tissues with clamp 
connections. 

Subhypogeously on earth and humus under Eucalyptus globulus, Quer- 
cus sp., etc., in California, U.S.A., fruiting during the rainy season (No- 
bember until April), in Oregon also in June. 

Material studied. Catirorn1A. Alameda County: University of California campus, 
Berkeley, Dec. 12, 1904, Gardner 229 (holotype, UC 221827), MICH. Topotypes: 
25 March 1911, Nichols, MICH, NY (distributed as UC Herbarium Exsiccata 399) ; 
March 10 and April 11, 1911, Harper, NY; fall, 1923, Parks, NY; April 10, 1935, 
Copeland (det. E. E. Morse), NY; February and March, 1931, Morse, NY. Santa 
Clara County: Alma, March 2, 1919, Parks Z9, NY; Stanford University campus, 
March 5, 1942, Cooke & Doty 16619, MICH. Also material preserved at FH from 
same regions as above. OREGON: Corvallis, Zeller, NY. 


Illustrations: Setchell (1907), pl. 107, figs. 4-8. Lloyd, Myc. Notes V:788, fig. 1184. 
Heim, Rev. Myc. 25:21. 1950. figs. 1-18. 


Setchell, in his original description, mentions the doubts he felt when 
publishing this species in Secotium: “In appearance and structure, this 
species varies so much from S. Guienzi, the type of the genus, that it may 
well be doubted whether it will ultimately be considered cogeneric with it, 
but, at present, it seems best to refer it to Secotium rather than attempt 
to split up that genus.”’ We, on the other hand, find it rather amazing that 
between 1907 and 1957, in over half a century of mycological progress, 
no such attempt has been made. 


Setchelliogaster aurantium (Zeller) Sing. & Smith, comb. nov. Seco- 
tium aurantium Zeller, Mycologia 39:292. 1947. 

Gastrocarp 14 mm. broad, 15 mm. high, rounded umbonate above; 
peridium thin, not breaking from stipe-columella, bright orange, capucin 
yellow at base, gleba with small cavities, light brown. 

Stipe terete, up to 3 mm. in diameter (about 20 mm. long), white, 
smooth, stuffed; columella also white, thin, percurrent; mycelium form- 
ing white rhizomorphs at base reminding one of those of the phalloids. 
Context white. 

Spores 12-13 & 7.5-8 y, light brownish ochre, smooth, ellipsoid to ellip- 


: SETCHELLIOGASTER 1 


Se 
An Yo 


Fics. 1-3. Setchelliogaster tenuipes: 1, epicutis of peridium, x 450; 2, spores, 
x 1000; 3, basidium, x 450. 


soid-fusoid, with many tiny oil droplets inside, with moderately thickened 
wall composed of at least two layers, the outermost one well-colored, but 
perisporium not noticeable and no ornamentation seen, symmetric, with 
a distinct germ pore in a continuous wall showing lighter different ma- 
terial, but not distinctly showing an opening, and not truncate. 

Basidia 21-22 « 8.2, with slightly oblique, conical, slightly curved 
sterigmata; pseudoparaphyses numerous, 19-20 & 12.5; cystidia none 
seen. 

Hyphae of the hymenophoral trama hyaline, the mediostratum regu- 
larly arranged and consisting of a strand of parallel or subparallel rather 
broad axillarly arranged hyphae, bordered on both sides by layers of 
spherocysts which become gradually smaller as the subhymenium and 
hymenium are approached, hyphae of the mediostratum 4—16.5 y. thick, 
spherocysts about 28 22,4; peridium consisting of hyphous elements 
which are hyaline and 3-16. thick, outermost layer of peridium divided 
from trama proper by a hypodermium-like layer which is not very sharply 
differentiated from both the outermost layer and the internal hyphous 
layer, but consists of spherocysts or broad swollen hyphae forming a 
pseudoparenchymatous tissue which, on its outer side carries a palisade 
of clavate to irregular, more or less erect to isodiametric elements with 
remarkably thick lemon-yellow walls (walls 1.5y thick), the claviform 
ones about 22 & 10y.; all tissues with clamped hyphae. 

On the ground, apparently in the forest, fruiting in summer, Cuba. 


Material studied. Cuba, Santa Clara Prov. July 6, 1941, W.L. White 803 (NY & 
FH), type. 


78 MADRONO [Vol. 15 


This species differs from the other member of the genus in the structure 
of the spore wall which lacks any ornamentation, but the important char- 
acter of the outer layer of the peridium places the species in this genus. 
The microscopic data given in the description are taken from the type, 
but most of the macroscopic data are from the original description. 

There is no other genus of secotiaceous fungi known at present with a 
cellular outer peridial layer such as found in either of these two species. 
Weraroa has a different peridial structure and its spores have a broader 
truncate apex. 

As for the affinities of this genus with representatives of the Agaricales, 
we would say that it is even more closely related to the family Bolbitia- 
ceae than Galeropsis which lacks a cellular outer layer of the peridium. 
While the habit is definitely more reminiscent of Conocybe in Galeropsis 
than in Setchelliogaster, the latter has also some similarity with such 
species as the fleshier 4A grocybes, Conocvbe intrusa, etc. The species with 
ornamented spores may be compared with such forms as are now com- 
bined in the subgenus Ochromarasmius of Conocybe where there is also 
a somewhat protruding germ pore. As a smooth-spored species, Setchel- 
liogaster aurantiacum would be comparable with some of the more thin- 
walled representatives of Conocybe such as have been described in Kuh- 
ner’s monograph. The color of the peridium in this species does not seem 
to be rare in Conocybe. Finally, the relatively voluminous hyphal ele- 
ments making up the hymenophoral trama are distinctive both for this 
species and in Conocybe where they may serve as a generic character 
separating this genus from the neighboring genera of the family Bolbitia- 
ceae. It may be added that careful observation always reveals the exist- 
ence of numerous pseudoparaphyses in the hymenia of Setchelliogaster 
as well as in Conocybe and Bolbitius. The spores of S. tenutpes are some- 
times observed to be forked or otherwise misformed or doubled. Such 
monstrosities are common in the genus Conocybe and also in some other 
bolbitiaceous genera. 

In spite of all the congruence found between Setchelliogaster as a whole 
and the Bolbitiaceae as a whole, it cannot be denied that there are also 
features in this genus which recall similar features of agarics in other 
families such as the particular spore ornamentation of S. tenuipes. This 
remarkable structure was noticed by Singer (1951) and attention was 
drawn to the partly agaric-like characters. Although the species was then 
characterized as agaricoid, no attempt was made to link it up with any 
particular group of agarics, a task which has become easier now that more 
species of Secotium have become known more thoroughly. 

The ornamentation of Setchelliogaster tenuipes is not exactly dupli- 
cated by any known bolbitiaceous species. It is found, with slight varia- 
tions, in Ganoderma among the “polypores”, and in widely separated 
species and groups of species of the Agaricales such as Boletellus, Metra- 
ria, Fayodia (a genus considered as part of Mycena sensu lato by A. H. 
Smith), by a species described as Tubaria thermophila by Singer, and 


1959] QUICK: CEANOTHUS 79 


another described as Kuhneromyces alpinus by Smith.” A similar spore 
type has been observed by us in the type of Secotium eburneum Zeller, 
but at present we do not wish to transfer that species to Setchelliogaster 
because the cellular layer of the peridium is overlaid by a layer of gela- 
tinous filamentous hyphae forming a pellicle which is the outermost layer 
of the peridium. 

LITERATURE CITED 


Pouzar, Z. 1958. Nové rody vyssich hub. II. Ceska Mykologie 12 (1) :31-36. 
SETCHELL, W. A. 1907. Two new hypogaeous Secotiaceae. Jour. Mycology 13:236-241. 
SINGER, R. 1949. The Agaricales (mushrooms) in modern taxonomy. Lilloa 22:1-832, 
pls. 1-29. 

. 1955. Le Genre Melanomphalia Christianson. Rev. Mycologie 20:12-17. 

. 1957. New genera of Fungi X.—Pachylepyrium. Sydowia 11:320-322. 
ZELLER, S.M. 1941. Further notes on Fungi. Mycologia 33:196-214. 

. 1947. More notes on Gasteromycetes. Mycologia 39:282-312. 


CEANOTHUS SEEDS AND SEEDLINGS ON BURNS 
CLARENCE R. Quick! 


Blue Canyon, the large rocky canyon of Big Creek, a tributary of Kings 
River, lies in the Sierra Nevada just southeast of Shaver Lake, Fresno 
County, California. This canyon once supported a magnificent stand of 
timber and in spots still does. Portions of the northerly part of the canyon 
were logged with “steam donkeys” around 1915. The ecologic course of 
reforestation on logged or burned forest areas, especially on high-quality 
forest sites, is often interesting and significant. Causes for variations in 
the reforestation process are not fully understood even yet. 

The areas of Blue Canyon logged in 1915 were clear-cut, but somehow 
the methods used in the logging, the weather cycle after the logging, or 
perhaps other and unrecognized factors caused the cutover area to regen- 
erate timber species promptly, especially sugar pine. For a while the new 
growing forest was somewhat brushy, but the brush slowly gave way to 
the competition of the trees, mostly pine trees, and by 1945 the old cut- 
over area in the northeast corner of Blue Canyon was a beautiful stand of 
pole-sized sugar pine, ponderosa pine, white fir, and incense-cedar. Sugar 
pine predominated in much of the stand. Some decadent brush persisted, 
largely in the forest openings. On a one-acre study plot in the northwest 
corner of section 14, T10S, R25E, were found some 180 sugar pines, 
mostly pole-sized trees 4 to 12 inches in diameter. 


1 Pathologist at California Forest and Range Experiment Station, maintained at 
Berkeley, California, by the Forest Service, U.S. Department of Agriculture, in co- 
operation with the University of California. 

* This latter seems to be congeneric with Melanom phalia nigrescens and M. platen- 
sis. Singer (1955) placed Melanomphalia in the Cortinariaceae, and (1957) pro- 
posed the combination Melanomphalia alpina (Smith) Sing. 


80 MADRONO [Vol. 15 


One hot afternoon in August 1947 a sawmill near the bottom of Blue 
Canyon (at an altitude of about 3,500 feet) caught fire and burned. The 
fire escaped and spread over 4 to 5 square miles of forest in the north- 
eastern part of Blue Canyon. The fire was stopped near the rim of the 
canyon at an altitude of about 6,000 feet. The one-acre study plot was 
completely burned—all conifers and, for that matter, all plant parts above 
the ground surface were killed. Burned forest extended roughly for a mile 
in all directions from the plot. 

In 1948 some small plots were established on the burned one-acre plot 
to see what plants were “coming back” on the burn. Ten separate mil- 
acre plots were distributed over the old one-acre plot. (A milacre—one- 
thousandth of an acre—is a square with sides of 6.6 feet.) Current-season 
seedlings which came up in the spring of 1948, the first spring after the 
burn, and resprouts from subsurface parts of plants which survived the 
burn were counted and recorded. 

From the 10 milacres were removed 281 seedlings of Mariposa man- 
zanita (Arctostaphylos mariposa Dudley), 1,204 seedlings of Sierra Ne- 
vada gooseberry (Rzbes roezli Regel), and 1,994 seedlings of deerbrush 
(Ceanothus integerrimus H. & A.). This is a total of 3,480 seedlings of 
these three brush species on 10 milacres (1/100 acre). 

From a single milacre were removed 83 manzanita seedlings; from an- 
other, 612 gooseberry seedlings; and from still another, 1,814 deerbrush 
seedlings. Subsequent checks of these milacre plots showed that no cur- 
rent-season seedlings of these three species came up in 1949 or in 1950. 
No conifer seedlings appeared on the milacre plots, and almost none with- 
in the boundaries of the old one-acre plot. 

This ability to germinate suddenly after fire is characteristic of the 
seeds of several genera of shrubby plants in California; see, for example, 
Jepson (1925). Species which react in this way are commonly called fire- 
type plants. Seeds of firetype species are mechanically durable, long-lived, 
and come to be relatively ubiquitous in the forest floor of the Sierra Ne- 
vada. Quick (1956) has shown that large numbers of many kinds of seeds 
are “stored” in the duff even under virgin stands of timber. Quick (1935) 
has also shown that treatment in boiling water plus stratification will con- 
dition seeds of many species of Ceanothus L. for immediate germination. 

The longest treatment in boiling water of the experiments reported in 
1935 was 5 minutes. This treatment resulted in very satisfactory germina- 
tion in some species, and it was obvious that the limit of tolerance of some 
collections was not closely approached. So the question arose, just how 
much exposure to boiling water, for example, will Ceanothus seeds endure? 

To seek an answer, samples consisting of 100 airdry seeds of deerbrush 
and of mountain whitethorn (Ceanothus cordulatus) were treated in the 
Berkeley laboratory by tossing them into a screen sieve suspended in 
vigorously boiling tap water. After treatment, the sieve and seeds were 
removed from the boiling water, soused in cold water, planted in auto- 
claved river sand, stratified to obviate embryo dormancy, and germinated 


1959] SNOW: CHROMOSOME NUMBERS 81 


in a greenhouse. Samples of seed collection Q#026 (deerbrush, collected 
on the South Fork of Stanislaus River at about 4,800 feet altitude) were 
treated in boiling water for 1 to 20 minutes. Twelve per cent of these seeds 
germinated after having been boiled for 20 minutes. Samples of seed col- 
lection Q#239 (mountain whitethorn, collected near Strawberry, Stanis- 
laus National Forest, at about 5,400 feet altitude) were treated for 5 to 30 
minutes in boiling water. Twenty-five per cent of these seeds germinated 
after being boiled for 25 minutes, but none germinated after being boiled 
for 30 minutes. A manuscript, now nearing completion, will analyze statis- 
tically the seed germination reactions of several species of Ceanothus with 
respect to seed age, altitude of seed collection, length of exposure to boil- 
ing water, and temperature and length of stratification treatment. 

Many seeds, even Ceanothus seeds, are destroyed in any forest fire, but 
the above preliminary experiments suggest that seeds of many firetype 
plants are very durable, long-lived, and sufficiently resistant to high tem- 
peratures to escape destruction and to completely revegetate a heavy burn. 
Thus starts the first stage in a new cycle of forest development. 


California Forest and Range Experiment Station, 
University of California, Berkeley. 


LITERATURE CITED 


Jepson, Witiis L. 1925. The life zones. Jn A manual of the flowering plants of 
California, pp. 4-8. Berkeley, California. 
Quick, CLARENCE R. 1935. Notes on the germination of Ceanothus seeds. Madrono 
3(3) 2135-140. 
. 1956. Viable seeds from the duff and soil of sugar pine forests. Forest 
Science 2(1) :36—42. 


CHROMOSOME NUMBERS OF CALIFORNIA PLANTS, WITH 
NOTES ON SOME CASES OF CYTOLOGICAL INTEREST 


RICHARD SNOW 


Meiosis has been examined in microsporocytes of various species of 
Californian plants, and the resulting chromosome numbers are listed in 
Table 1. During the course of these observations several meiotic phenom- 
ena have been observed that have cytological interest beyond chromosome 
number. These will be discussed briefly in order to point out some areas 
possibly worthy of further study. 


Mertuops. Buds were fixed in 1:3 acetic-alcohol for one to several days 
and stored in 70 per cent alcohol under refrigeration. After soaking in 
water for a few minutes the material was softened in IN HCl at 60°C. 
for 5-10 minutes, rinsed with water, and the anthers squashed in aceto- 
orcein or aceto-carmine. The Fritillaria material was stained by the Feul- 
gen reaction. Permanent slides were made by Bradley’s (1948) method, 
without removing the coverslip. 


82 MADRONO [Vol. 15 


TABLE 1. DOCUMENTED CHROMOSOME NUMBERS OF CALIFORNIA PLANTS! 


CHROMO- 
SPECIES SOME LOcALITY AND COLLECTOR 
No. (2n) 
Fritillaria biflora Lindl.* 24+0-8ff Near Saddle Peak, Santa Monica Mts., 
Los Angeles County, Lewis in 1958. 
Delphinium polycladon Eastw. 16 Rodger’s Lake, Tuolumne County, 
Snow 284. 
Ranunculus californicus Benth. 28-+f Road from Bridgeport to Highway 
140, Mariposa County, Snow and 
Wedberg in 1953. 
Eschscholtzia mexicana Greene 12 15.3 miles north of Goffs on the road 
to Lanfair, San Bernardino County, 
Snow 18. 
Thely podium lasiophyllum 48 Red Rock Canyon, Kern County, 
(H. & A.) Greene Snow 410. 
Isomeris arborea Nutt.* 40 2.7 miles west of Whitewater, River- 
side County, Raven 11409. 
Point Dume, Los Angeles County, 
Snow (no specimen). 
Red Rock Canyon, Kern County, 
Snow 407. 
Lupinus superbus Hel. var. 48 Matterhorn Canyon, Tuolumne 
elongatus (Greene) County, Snow 292. 
C. P. Smith* 
Astragalus Bolanderi Gray 22 Badger Flats, near Huntington Lake, 
Fresno County, Snow 62. 
Astragalus leucopsis (T. & G.) 22 Point Dume, Los Angeles County, 
Torr. Raven 13857. 
Astragalus oocarpus Gray fey) 4.6 miles north of Santa Ysabel, 
San Diego County, Raven and 
Snow 9557. 
Astragalus pomonensis Jones 22 1.7 miles west of Dripping Springs 
Guard Station, Riverside County, 
Raven and Snow 9543. 
Astragalus crotalariae Benth. 24 Salton Sea State Park, Riverside 
County, Raven 11484. 
Datisca glomerata (Presl) 22 Banning-Idyllwild Road near Grand 
Benth. and Hook. View Guard Station, Riverside 
County, Raven and Snow 11115. 
Epilobium obcordatum Gray 36 Little Slide Canyon, Mono County, 
Snow 275. 

Oenothera californica Wats. 28 11 miles east of Riverside, Riverside 
var. californica* County, Snow and Mosquin 318. 
Gayophytum racemosum 14 Little Slide Canyon, Mono County, 

T.&G. Snow 273. 
Monardella odoratissima 42 Little Slide Canyon, Mono County, 
Benth. Snow 272. 


1 Voucher specimens and prepared slides are in the Herbarium of the University 
of California, Berkeley. 

* See discussion in text. 

f = fragment chromosome. 


1959] SNOW: CHROMOSOME NUMBERS 83 


CHROMO- 
SPECIES SOME LocaLiTy AND COLLECTOR 
No. (2n) 
Datura meteloides DC.* 24 See Table 2. 
Plantago insularis Eastw. 8 Near Red Rock Canyon, Kern 
County, Snow 416. 
Pedicularis groenlandica Retz. 16 Pass between Little Slide Canyon 


and Slide Canyon, Tuolumne 
County, Snow 276. 


Proboscidea Jussieui Keller 30 Putah Creek, west of Davis, 
Yolo County, Snow 605. 
Senecio Fremonti T.&G. 40-++f Little Slide Canyon, Mono County, 
var. occidentalis Gray* Snow 270. 


My thanks are due Dr. Harlan Lewis for the buds of Fritillaria biflora, 
and to Mr. Peter Raven for the material of Astragalus crotalariae and 
for permission to cite his unpublished observations on /someris arborea. 


CYTOLOGICAL NOTES 


FRITILLARIA BIFLORA. The fragments in this species are one-third to 
one-fourth the length of the long arms of chromosomes of the basic com- 
plement at anaphase II. It is not certain how constant their number is 
from plant to plant, since buds from several plants were fixed in the same 
vial. However, since this species has usually only two to four flowers per 
plant it is almost certain that each bud in a meiotic stage was derived 
from a different plant. The number of fragments found in various buds 
from this collection was 2,4, 6, or 8. There was also occasionally a smaller 
variation in number from one PMC to another within the same bud. 

At metaphase I from 1 to 4 “‘fragment-pairs”’ have been seen in various 
buds (fig. 1), and no fragments have been seen at this stage which were 
not “paired.” These dual associations may represent the synapsis of 
homologous fragments or the division of unpaired fragments. The few 
mitotic cells on the slides were not clear enough for a definitive answer. 
The fragments do not congress to the metaphase plate with the other bi- 
valents, but instead usually lie off in the cytoplasm above or below it. 
Their position at metaphase I may account for their frequent inclusion 
in the daughter nuclei. However, lagging of the fragments is not infre- 
quent at anaphase. At anaphase I a fragment-pair may separate, one or 
both halves being included in a nucleus; or the pair may be included, 
without separation, in one daughter nucleus. As a result of lagging and 
irregular separation, an anaphase nucleus may receive from none to as 
many fragments as the PMC contains. Anaphase II distribution is like- 
wise variable. Irregularities of division possibly also occur in mitosis, and 
may account for the variation in the number of fragments observed in 
different PMCs of the same bud. 

Anaphase observations suggest that the fragments probably possess a 
centromere, though one which is perhaps less efficient in division than 
normal. An area which is probably the centromere region appears sub- 


84 MADRONO [Vol. 15 


terminal in mitotic cells. The preparations were not especially good for 
study of this feature, however, but certainly the fragments gave no indi- 
cation of being isobrachial. They do not, therefore, appear to be products 
of mis-division of a centromere followed by the uniting of sister chroma- 
tids to convert a telocentric into an iso-chromosome, as has been found 
in the triploid Fritillaria latifolia major by Darlington (1940; see also 
Darlington, 1939). Whatever their origin it seems likely that all the frag- 
ments observed in the various buds (and hence from several plants) 
are the result of a single primary event of the past, both because their 
similar form and because the formation of nuclei with extra fragments as 
a result of non-disjunction offer cytological evidence of a means for their 
increase. Pollen grains with fragments are no doubt functional at least 
occasionally, if we may extrapolate from the results of Kayano (1956), 
who found almost a 1:1 ratio of 2n:2n+f plants in the cross Lilium cal- 
losum 2n 2 X 2n+f %. In this case the fragment appears to be a telo- 
centric from an originally acrocentric supernumerary. As in F. biflora, 
however, no pairing of the fragment with the basic set occurred at meta- 
phase I, though several fragments could pair among themselves. 

The western Amercan species F. lanceolata, F. pudica, F. recurva and 
F. folcata are reported to have fragment chromosomes (Beetle, 1944; 
La Cour, 1951). In addition, Darlington and Wylie (1955) list several 
Old World species, and also the Californian species F. pluriflora, as hav- 
ing B chromosomes; these may be the same type as those present in F. 
biflora. It is possible that such chromosomes are present throughout the 
genus, in some species frequent, in others rarer. What their role may be 
in the population dynamics and evolution of the species remains to be 
determined. 


ISOMERIS ARBOREA. This plant has enjoyed a measure of fame since 
Billings (1937) reported it to be haploid with a highly anomalous embry- 
ology. Since then several workers have reinvestigated the embryology 
and found it to be essentially normal, although with certain peculiar 
features (Maheswari and Kahn, 1953; Sachar, 1956). The chromosome 
situation, however, has not yet been clarified. Observations of meiosis 
made by Mr. Peter Raven of a plant growing near Whitewater, Riverside 
County, and by the author of one plant from Point Dume, Los Angeles 
County, and of another from Red Rock Canyon, Kern County, have 
shown the species to be a diploid with 20 pairs of chromosomes (fig. 2). 
The chromosomes are small and difficult to stain well, and buds in meiosis 
are found rather infrequently, so that it is not favorable material cyto- 
logically. However, these observations have shown a normal meiotic 
sequence from diakinesis through telophase II. Further, my examination 
of pachytene showed two paired strands in clear cells, and in a few in- 
stances where the synapsed chromosomes were more widely separated 
than usual they could be seen to have a similar chromomere pattern. From 
the figures published by Billings, it seems clear he observed normal meta- 


1959 | SNOW: CHROMOSOME NUMBERS 85 


1 2 
(D ys Domes, 
MEER fveergaotverety = | Tea 
we a= 
WH ao ee 
3 4 


ci rer 


5 6 


Fics. 1-6. Chromosome plates. 1, Fritillaria biflora: 12 bivalents plus 2 “fragment- 
pairs”; 2, Isomeris arborea: 20 bivalents; 3, Lupinus superbus var. elongatus: 24 bi- 
valents, the largest with unequal arms; 4, Lupinus superbus var. elongatus: left, 
five A bivalents at metaphase or early anaphase, right, five A bivalents at anaphase; 
5, Datura meteloides: 10 bivalents plus a chain of 4 chromosomes; 6, Senecio Fre- 
montii var. occidentalis: 20 bivalents plus 1 fragment. 

(Fig. 1, & 358; Fig. 2, * 952; Figs. 3-6, & 1071.) 


phases and anaphases in both of the meiotic divisions, but as he was 
convinced that only univalents were present at diakinesis, he interpreted 
these as equational separations of univalents. 

Isomerts arborea must therefore be returned to the category of rather 
ordinary plants. 


86 MADRONO [Vol. 15 


LUPINUS SUPERBUS var. ELONGATUS. At metaphase 24 bivalents are 
present. One of them is conspicuously larger than the rest, and I desig- 
nate it the A bivalent (fig. 3). In the pollen mother cells from one inflores- 
cence this bivalent was unequal, that is, one chromosome was shorter 
than the other. Judging from metaphase appearances the A chromosomes 
are probably acrocentric, and regularly form a chiasma in the short arm. 
In about ten to twenty per cent of the cells the longer (unequal) arms 
also show evidence of a chiasma. Figure 4, right, shows the appearance of 
5 A bivalents with a chiasma in the short arm; figure 4, left, 5 with chias- 
mata in both arms. It is possible that the A chromosomes usually form 
a chiasma in each arm, and that early terminalization in the longer arms 
releases them from their association. This is suggested by the fact that 
clear one-chiasma bivalents appear further along in anaphase separation 
than those with two chiasmata. 

White (1954) has given a diagram showing the various types of meiotic 
segregations which have been observed or are presumed to occur with 
unequal bivalents of orthopteroid insects. The segregation at the first divi- 
sion will depend upon three factors: the position of the centromere, the 
position of the chiasmata, and the position of the inequalities. His dia- 
gram is based on two assumptions: that the extra segment is terminal 
rather than interstitial, and that only one chiasma is formed in the un- 
equal bivalents. White has based these two assumptions on the evidence 
presently available from grasshoppers and phasmids. In Lupinus superbus 
we have what appears to be an example of an interstitial inequality in 
an unequal bivalent where chiasmata are formed in each arm, a situation 
for which White had no examples. The evidence for the existence of more 
than one chiasma in the A bivalent is unequivocal: two-chiasmata bi- 
valents have been seen in many metaphase cells. The evidence for the 
interstitial position of the inequality is less direct. It depends on the fact 
that only reductional separations of the inequality have been observed 
in both one-chiasma and two-chiasmata A bivalents. Should a chiasma 
be formed in the long arm proximal to a terminal inequality, then an equa- 
tional separation would result, with one chromatid of the anaphase chrom- 
osome longer than the other. Such equational separations were never ob- 
served in about fifty cells analyzed, although it must be admitted that 
equational separations might not be readily detected, since the chromo- 
somes are rather small. Furthermore, only terminal junctions have been 
found in the unequal arm of the A bivalent at metaphase, while a chiasma 
proximal to an inequality would be expected to remain interstitial at this 
stage. It thus appears that reductional separation of the inequality is the 
rule, and hence that it occupies an interstitial position in the chromosome 
arm. Whether it is a duplication, a deficiency, or a heterochromatic region 
(as is so often the case in insects), has not been determined. 

Two other chromosomal types might be expected in the population, 
namely the corresponding homozygotes. The size difference between the 
A chromosomes is great enough so that there should be no difficulty dis- 


1959] SNOW: CHROMOSOME NUMBERS 87 


tinguishing them. First metaphase was studied in another inflorescence 
(and hence probably from another plant). In this case both members of 
the A bivalent appeared about the same size, and were as large as the 
larger member of the heterozygous plant. If the inequality is a deficiency, 
the homozygote for the smaller member may not be viable. A much larger 
sample of the population would be desirable in order to determine the 
prevalence of the three types, and to compare their frequencies with the 
expectation based on the Hardy-Weinberg formula. 


OENOTHERA CALIFORNICA var. CALIFORNICA. The plant examined of 
this collection was a tetraploid (n=14). A varying number of pairs and 
rings of four chromosomes was found at diakinesis and metaphase, the 
maximum seen being six rings of 4 plus 2 pairs. The pairing observed sug- 
gests that this species was derived from a diploid form which formed 7 
bivalents. In Oe. californica var. glabrata, Lewis et al. (1958) reported 
two associations of 8 chromosomes, indicative that the diploid form might 
have been a structural heterozygote. These observations on Oe. califor- 
nica (subgenus Anogra) parallel those of Hagen (1950) on two tetraploid 
species of the subgenus Raimannia, who found only pairs or rings of 4 in 
Oe. tetragona, but long chains (of up to 14 chromosomes) and univalents 
in Oe. speciosa var. Childsi. In this latter species, diploids are known 
which form one or two rings of 4, or 7 bivalents. 

As is usual with multivalent configurations in the Onagraceae, adjacent 
chromosomes were regularly segregated to opposite poles at anaphase, 
thus supporting Garber’s (1954) suggestion that if a diploid species shows 
directed segregation of chromosomes from interchange configurations, the 
autotetraploid will likewise show directed segregations of the quadriva- 
lents. These observations coincide with those of Catcheside (personal 
communication to Garber, l.c.) that “Oenothera tetraploids from struc- 
turally homozygous diploids show almost regularly zigzag orientation of 
the rings of four.” As a consequence of such regular segregation and the 
lack of lagging chromosomes, fertility in this tetraploid should be quite 
high. 


DATURA METELOIDES. Ten wild plants of this species have been exam- 
ined and five have been found to be heterozygous for a reciprocal trans- 
location (table 2). A sixth (from Caliente Creek) suggests by frequent 
univalent formation that it may also be heterozygous for a structural 
change which is unidentified at present. The plants are apparently not all 
heterozygous for the same interchange, because a plant from Isabella 
Reservoir and plants from Putah Creek (Snow 322—5, 324-5) character- 
istically formed rings of four at metaphase, while in the Yucaipa plant 
(11076B) only chains of four were observed. In a third Putah Creek plant 
(Snow 322-1) variable configurations were formed at metaphase. Usually 
the groups of four chromosomes appeared as a branched chain (fig. 5). 
Sometimes one of the pairs of chromosomes of the configuration had a 
chiasma in each arm, so that the association resembled a kite with the 


88 MADRONO 


[Vol. 15 


TABLE 2. LOCALITIES AND METAPHASE ASSOCIATIONS OBSERVED IN WILD INDIVIDUALS 


OF DATURA METELOIDES 


METAPHASE 
LOcALITY AND COLLECTOR ASSOCIATIONS 
Near Ensenada, Baja California, Mexico, Snow (no specimen). 122) 
Near Yucaipa, Riverside County, Raven and Snow 11076A. 12(2) 
Near Yucaipa, Riverside County, Raven and Snow 11076B. 10(2)+ (4c) 
Caliente Creek, Kern County, Snow 248. 1VACD Ee) a 
11,2) £24) 
Isabella Reservoir, Kern County, Snow 253. (coll. 6/18/54). 10(2)+ (4r) 
Isabella Reservoir, Kern County, Snow 253A. (coll. 7/30/57). 12(2) 
Putah Creek, south of Davis, Yolo County, Snow 322-1. 10(2)+ (4c) or 
10(2) + (4r) 
Putah Creek, south of Davis, Yolo County, Snow 322-5. 10(2) + (4r) 


Putah Creek, south of Davis, Yolo County, Snow 322-8. 12(2) 
Putah Creek, west of Davis, Yolo County, Snow 324-5. 10(2)+ (4r) 


(2) = bivalent 
(4c) = chain of four 


(1) = univalent 
(4r) = ring of four 


tail coming off one side. Other configurations observed, much less fre- 
quently, were a “T” with a univalent, a non-disjunctional ring of 4, and 
12 bivalents. 

Staiger (1955) has found interchanges in natural populations of the 
mollusc Purpura lapillus which lead to metaphase configurations similar 
to those which I have found in one plant (Snow 322-1). He has shown 
how they may be accounted for by the interchange of short end pieces 
of metacentric chromosomes plus variations in chiasma formation. The 
same scheme can account for the configurations in this plant of Datura 
meteloides. The translocations in other plants probably represent larger 
exchanges of chromosome end segments. 

Four chromosomal arrangements have been found in this species by 
Satina (1953), and one of them (type I) has been compared to the stand- 
ard race of Datura stramonium. The other three arrangements presum- 
ably differ from type I by reciprocal translocations. The type I arrange- 
ment has been found in twenty-eight races. Unfortunately the origin of 
none of these races is cited in the paper, so that the geographical distri- 
bution of this type is unknown. Furthermore, the existence in nature of 
individuals heterozygous for reciprocal translocations was not reported 
for D. meteloides, and apparently has not been for any other species of 
Datura, even though individuals of the same species may be homozygous 
for different chromosome arrangements while members of different species 
invariably are. 

Study of D. meteloides is being continued with the view of discovering 
the role translocations may play in the populations. 

SENECIO FREMONTII var. OCCIDENTALIS. The extra chromosome in this 


1959] SNOW: CHROMOSOME NUMBERS 89 


material is a small fragment about one-fourth the length of an arm of the 
longer chromosomes of the complement at diakinesis. At this stage it is 
often found lying in the proximity of a bivalent with widely diverging 
arms, one of which appears shorter than the other by about the size of 
the fragment. The fragment may even appear attached to this arm by 
strands of stainable material. With about an equal frequency, however, 
it is found completely free of this readily-recognizable bivalent (fig. 6). 

Stages later than diakinesis were not favorable for study in these prepa- 
rations. In a few metaphase cells which could be analyzed, the fragment 
was attached to a bivalent, probably a continuation of a diakinetic asso- 
ciation which was so often observed. The fragment was included in one 
of the daughter nuclei in all instances save one. In this one instance, out 
of about fifty cells in telophase examined, the fragment had been excluded 
and was dividing. 


Department of Genetics, 
University of California, Davis 


LITERATURE CITED 


BEETLE, D. E. 1944. A monograph of the North American species of Fritillaria. 
Madrono 7:133-159. 

Bitiincs, F. H. 1937. Some new features in the reproductive cytology of Angio- 
sperms, illustrated by Isomeris arborea. New Phytol. 36:301-326. 

BrapLtey, M.V. 1948. A method for making aceto-carmine slides permanent without 
removal of the cover slip. Stain Tech. 23:41-44. 

CLELAND, R. E. (ed.) 1950. Studies in Oenothera cytogenetics and phylogeny. Indiana 
Univ. Publ. Sci. Ser. No. 16. 348 pp. 

Daruincton, C. D. 1939. Misdivision and the genetics of the centromere. Jour. 
Genet. 37:341-364. 

1940. The origin of iso-chromosomes. Jour. Genet. 39:351-361. 
and A. P. Wylie. 1955. Chromosome Atlas of Flowering Plants. George 
Allen and Unwin, Ltd. London. 

GarBER, E. D. 1954. Cytotaxonomic studies in the genus Sorghum. III. The polyploid 
species of the subgenera Parasorghum and Stiposorghum. Bot. Gaz. 115:336-342. 

Hacen, C.W., Jr. 1950. A contribution to the cytogenetics of the genus Oenothera 
with special reference to certain forms from South America, in Cleland, 1950, 
pp. 305-348. 

Kayano, H. 1956. Cytogenetic studies in Lilium callosum. II. Preferential segre- 
gation of a supernumerary chromosome. Mem. Fac. Sci., Kyushu Univ., Ser. E, 
2:53-60. 

La Cour, L.F. 1951. Heterochromatin and the organization of nucleoli in plants. 
Heredity 5:37-50. 

Lewis, H., P. H. RAven, C. S. VENKATESH, and H. L. WeEpBERG. 1958. Observations 
of meiotic chromosomes in the Onagraceae. El Aliso 4:73-86. 

ManesHwart, P. and R. Kann. 1953. Development of the embryo sac, endosperm, 
and embryo in Isomeris arborea—a reinvestigation. Phytomorphology 3:446—-459. 

SacHar, R. C. 1956. The embryology of Isomeris—a reinvestigation. Phytomor- 
phology 6:346-363. 

Satina, S. 1953. Chromosome end arrangements in Datura inoxia, D. meteloides, 
and D. metel. Am. Jour. Bot. 40:638-646. 

STAIGER, H. 1955. Reziproke translokationen in nattrlichen populationen von Pur- 
pura lapillus (Prosobranchia). Chromosoma 7:181-197. 

Waiter, M.J.D. 1954. Animal Cytology and Evolution. 2nd edition. Cambridge 
University Press. 


90 MADRONO LVol. 15 


CHROMOSOME COUNTS IN THE GENUS GAYOPHYTUM? 


Davin G. Drxon 


Gavophytum, an annual member of the family Onagraceae, consists of 
sixteen recognized taxa grouped into eight species. The genus, founded in 
1832 by A. de Jussieu on a South American plant, Gayophytum humile 
Juss., is confined to narrow ecological niches along the cordillera of North 
and South America from Canada to Cape Horn, with an interval in Mex- 
ico and Central America where it has not been reported. It has not been 
found outside the mountain ranges of the western Americas. 

As a genus, Gayophytum is very poorly known owing to the small size 
of the plants, the insignificant flowers (rarely as large as 5 mm. across), 
and the relative difficulty of obtaining specimens. The plants have linear- 
oblanceolate leaves and very slender, diffusely branched stems. The ulti- 
mate branchlets are often filiform and bear numerous tiny four-petaled 
flowers in the upper axils. The bushy habit of the plants often with a great 
number of branching thread-like stems gives them a fuliginous aspect, 
and when in flower, the blossoms appear to be floating near the ground, 
hence the popular common name “Ground Smoke.” 

The flowers are perfect, actinomorphic and tetramerous. The sepals are 
in the orthogonal aid the petais in the diagonal planes. Ovaries are bi- 
locular and inferior, with ovules in one row in each loculus. The fruit is 
dry, splitting loculicidally and septifragally so the inner portion of the 
fertile carpels is left as a seed-bearing column in the center (Saunders 
1940). 

Gavophytum may be distinguished from the very similar Epilobium by 
the lack of coma on the seeds, by the two-celled ovaries and fruits, and 
by having solitary pollen grains. 

Plants of Oenothera of similar size and habit and with equally reflexed 
sepals may be distinguished from Gavophytum by their more elongate 
hypanthium and four-celled ovary. The prevailing flower color of Oeno- 
thera is yellow; the flowers of Gayvophytum are white, turning rose-purple 
with age. 

In the words of Munz (1932): “The genus Gayvophytum offers an un- 
usually interesting series of plants varying in a comparatively small num- 
ber of characters, and these in every conceivable combination. Flowers 
may be small or large; capsules may be sessile or pedicelled, erect or 
spreading-deflexed, short or long, torulose or not torulose; the minute 
hairs may be appressed or spreading; seeds may be glabrous or pubescent; 
branching may be basal or distinctly above the base. The attempt at clas- 
sification . . . may be quite artificial; it has been arrived at only after 


1A thesis submitted in partial fulfillment of the requirements for the degree of 
Master of Science in Botany at the State College of Washington, 1957. The author 
wishes to express his appreciation to Dr. Adolph Hecht who suggested the problem, 
and served as advisor during the course of the research, and to Dr. Marion Ownbey 
who provided many suggestions during the preparation of the manuscript. 


1959] DIXON: GAYOPHYTUM 91 


TABLE 1. CHROMOSOME NUMBER AND SOURCE OF GAYOPHYTUM COLLECTIONS 


STUDIED 
CHROMOSOME 
SPECIES No. 2n SOURCE 
G. Helleri var. 14 Washington, Douglas Co.: 6 mi. N. of With- 
glabrum Munz row, Route 10-B, on Artemisia knolls, 
Dixon 9. 
G. humile Juss. 14 California, Mono Co.: Carnegie Inst. Tim- 


berline Station, Slate Creek Valley, 10,000 
ft. elev. H. Lewis. 


G. lasiospermum 28 Idaho, Valley Co.: 1 mi. S. of Lake Fork, 
Greene Route 15, on coarse gravel of ditches, 
: Dixon 14. 
G. lasiospermum 28 Idaho, Adams Co.: 4.5 mi. S. of New Mead- 
Greene ows, on cut-over Pinus ponderosa flat, W of 
Route 95, Dixon 17. 
G. Nuttallit T.& G. 28 Washington, Okanogan Co.: 5 mi. W. of Re- 


public Route 4, on moist clay and gravel 
road-cut, Dixon 7. 


G. Nuttallit T.& G. 28 Idaho, Valley Co.: within city limits of 
McCall, on sand banks, Dixon 12. 
G. Nuttall T.& G. 28 Idaho, Latah Co.: on roadside sand of Sad- 


dle between Twin Buttes, Moscow Moun- 
tain, Rumely & Dixon. 


G. Nuttall T.& G. 28 California, Tuolumne Co.: 4.4 mi. W. of 
Kennedy Meadows, Highway 108, Deadman 
Creek at 6,000 ft. elevation. Balls & Ever- 


ett 18074. 
G. Nuttalli var. 28 Washington, Stevens Co.: 2 mi. S. of Kettle 
Abramsti Munz Falls, Route 22, fine sand in Pinus flat, 
Dixon 3. 
G. racemosum var. 28 Idaho, Valley Co.: 1 mi. S. of Lake Fork, 
erosulatum Munz Route 15, on coarse gravel road-bank, 
Dixon 15. 


much study. Unfortunately geographical distribution which is so often a 
great aid to the systematist, is not very useful here, and one almost doubts 
the validity of some of the entities maintained because of lack of contin- 
uous or definite distribution. Furthermore, the floral parts are so minute 
that individual specimens can easily be thrown into the wrong group. Yet, 
as I have worked over many hundreds of sheets at various times... , I 
have been forced to conclude that there are several very real and definite 
entities in the genus.” 

The taxonomic treatments of Trelease (1893) and Munz (1932, 1951, 
1952) have proved satisfactory and have been followed in this study. 
Trelease expressed the opinion that Gayvophytum may be of rather recent 


92 MADRONO [Vol. 15 


differentiation from Oenothera, representing an accentuated montane- 
type. 

A possible origin of Epilobium (n=18) by the “addition” of eleven 
chromosomes of Gayophytum ramosissimum T.&G. (n=11) to the seven 
of Boisduvalia (n=7) has been suggested by Johansen (1933). This view 
that Epilobium may be of hybrid origin, however, has not been supported. 

Very little is known about the cytology of Gayophytum. Johansen’s 
(1933) work on G. ramosissimum is the only published report of previous 
cytological study in this genus. It was with the objective of adding to 
the cytological information on this genus that the present study was 
undertaken. 

Plants for this study were collected during the late summer months of 
1955 and 1956, in eastern Washington, central and southwestern Idaho, 
and northeastern Oregon. Additional seeds were supplied by Dr. Harlan 
Lewis and by Dr. P. A. Munz. Dr. Munz kindly identified the plants used. 

Plants of Gavophvtum nuttallii T.&G. were grown to maturity in the 
greenhouse, but no other species survived past the seedling stage in culti- 
vation. Root-tips were the source of the meristematic tissues used in this 
study. 

Field collected root-tips were fixed in Belling’s metaphase modification 
of Navashin’s fixative. Paraffin sections cut at ten microns were prepared 
and then stained by the crystal violet-iodine method (Johansen, 1940). 

Chromosome numbers of 2n=-14 and 2n=28 were found in the plants 
studied. Johansen (1933) lists the number 2n=22 for the North American 
species, Gavophytum ramosissimum. The number 2n=28 was found to 
be constant in collections of Gayophytum lasiospermum Greene, G. Nut- 
tallu, G. Nuttallii var. Abramsii Munz, and G. racemosum var. erosula- 
tum Munz. The number 2n=14 was discovered in Gayophytum Helleri 
var. glabrum Munz and in G. humile. 

On moistened filter-paper in Petri dishes kept at room temperature, 
seeds of the 14-chromosome species had a relatively vigorous germination 
percentage of 11 per cent, whereas those of the 28-chromosome species 
under the same conditions may have as low as one-fourth of 1 per cent 
germination. 

The two 14-chromosome species were collected at elevations of 2,000 
and 10,000 feet, respectively. The 28-chromosome species are from ele- 
vations ranging from 900 feet to over 6,000 feet. The 14-chromosome 
species tend to be smaller in stature and have smaller flowers than those 
with 28 chromosomes, but marked phenotypic differences are not evident 
between the two groups. 


DISCUSSION 


It is suggested that the 14 chromosomes counted in Gavophytum 
Helleri var. glabrum and in G. humile may make up the basic diploid com- 
plement of the genus. If this is the case, it may be assumed that the species 
with 28 chromosomes constitute a number of tetraploid taxa. The chromo- 


1959] DIXON: GAYOPHYTUM 93 


somes do appear in reasonably recognizable sets of four in these species. 

The chromosome numbers reported in this root-tip study are not con- 
sistent with the 22 chromosomes reported for Gayophytum ramosissimum 
(Johansen, 1933) in which embryo-sac meiosis was examined. The species 
previously studied was not available for examination in the current in- 
vestigation. The possibility exists of an aneuploid series of chromosomes 
in Gayophytum. The count of 22 might be an instance of a hyperploid 
(3x + 1) constitution. 

A brief survey of the chromosome numbers (Darlington and Wylie, 
1956) in the family Onagraceae reveals a range of basic numbers occur- 
ring in Clarkia (including Godetia) ranging from x — 5 through 7, 8, 
and 9; Gaura may have x —7 or 9. Other genera reported show x = 7 
constant for Oenothera, x = 8 for Jussieua, x = 11 for Circaea, Lopezita, 
and Fuchsia, with x = 15 for Zauschneria and x = 18 for Epilobium. 

It is apparent that Gayophytum fits most closely into the chromosome 
series of the family near Oenothera. Such a position would agree with 
Trelease’s view (1893) that Gayvophytum may be of rather recent differ- 
entiation from Oenothera. 

SUMMARY 


Mitotic chromosome studies were made on ten collections of six taxa 
representing five species of Gavophytum. Chromosome numbers of 2n= 
14 and 28 were found. The numbers 14 and 28 are in addition to the one 
of 22 previously reported for the genus. 

Chromosome counts for Gavophytum Helleri var. glabrum, G. humile, 
G. lasiospermum, G. Nuttall, G. Nuttallii var. Abramsii, and G. racemo- 
sum var. erosulatum are reported for the first time. Voucher specimens 
are on file in the herbarium of the State College of Washington. 


Department of Floriculture and Ornamental Horticulture, 
University of California, Los Angeles 24, California. 


LITERATURE CITED 


Dar LincTon, C. D. and Wytte, A. P. 1956. Chromosome atlas of flowering plants. 
Macmillan Company. New York. 

JouanseEN, D. A. 1933. Studies on the morphology of the Onagraceae VII. Gayophy- 
tum ramosissimum. Bull. Torrey Club 60:1-8. 

. 1940. Plant microtechnique. McGraw-Hill Book Co. New York and 
London. 

Mownz, P. A. 1932. Studies in Onagraceae VIII. The subgenera Hartmannia and 
Gauropsis of the genus Oenothera. The genus Gayophytum. Am. Jour. Bot. 19: 
755-778. 

. 1951. Gayophytum, in Abrams, L., Illustrated flora of the Pacific states, 
Vol. 3. Stanford University Press, Stanford, Calif. 

———. 1952. Gayophytum, in Davis, R.J., Flora of Idaho, William C. Brown. 
Dubuque, Iowa. 

SAUNDERS, E. R. 1940. Floral Morphology. Chem. Publ. Co. New York. 

TRELEASE, A. 1893. Revision of the North American species of Gayophytum and 
Boisduvalia. Ann. Rep. of the Mo. Bot. Gard. 5:107-122. 


04 MADRONO [Vol. 15 


REVIEWS 


Spring Flowers of the Lower Columbia Valley. By CLARA CHAPMAN Hitt. Illus- 
trated by Mary Comber Miles. pp. xi + 164. University of Washington Press. 1958. 
$3.00. 


Certainly one of the most frustrating and interest-stifling experiences of a novice 
at plant identification is his attempt to name a plant by using the formidable termi- 
nology of the keys and descriptions of technical manuals. Spring Flowers of the Lower 
Columbia Valley is the newest of a small number of “easy” manuals which have been 
designed to accustom the beginner to this terminology and to the technique of using 
the more detailed and comprehensive manuals of the Pacific Northwest flora. 

This small book is not, as its title suggests, a manual of the vernal flora of the 
lower Columbia River valley. Rather it is a florula of what might more appropri- 
ately be called the lower Willamette River valley in the vicinity of Portland, Oregon. 
It describes, in a simplified manner, about 200 of the more conspicuous herbs and 
shrubs of the season in that region. Use of technical words in the keys and descrip- 
tions is held to a minimum; a number of these terms are introduced by means of 
illustrative line drawings. Additional terms can be found in the glossary which, how- 
ever, offers some definitions so oversimplified as to be quite uninformative. Approxi- 
mately one-third of the species treated are adventive. Unfortunately no trees or ferns 
are considered, even though these groups often attract much attention from the be- 
ginners for whom this book was written. Following the introduction is a key to the 
genera; in the text the species are arranged under family headings and in some 
instances there are keys to the species within genera. The text is enlivened by brief 
but interesting notes on the natural history, uses, and lore of various species. Some 
of the families are characterized by short descriptions, a few of which are so trun- 
cated that they do not adequately distinguish one family from another. For example, 
the traits of the Saxifragaceae as stated could apply equally to the Crassulaceae for 
which no diagnosis is given. A problem common to all books which cover only a 
segment of a flora is the chance of misidentification by a beginner of a plant which 
appears to “key out” correctly, but is in fact unlisted in the book. Nevertheless, the 
selection of species considered is good, and few errors should arise providing it is used 
in the season and the region it is intended to cover. 

The format, binding, and price are attractive; the nomenclature is up-to-date and 
the book seems virtually free of typographical errors. For summer and autumn work 
its owners should feel confident and experienced enough to graduate to other more 
technical manuals of broader scope, many of which are listed in a bibliography. 
However, even after they have gone on to other works, they will want to keep this 
book on the shelf for the occasional pleasure of leafing through it to savour the 71 
superb, full-page line drawings by Mary Comber Miles. Perhaps these are its greatest 
recommendation—ROBERT ORNDUFF, Department of Botany, University of Cali- 
fornia, Berkeley. 


Spring Flora of the Dallas-Fort Worth Area [,| Texas. By Ltoyp H. SHINNERS. 
v, 514 pp., 11 plates, 2 maps. Published by the author, Southern Methodist Univer- 
sity, Box 473, Dallas 5, Texas. 1958. $4.75 (by mail, $4.85). 

The author of this unusual book has been working since 1945 in an area for which 
there has never been a complete descriptive flora. It is a particularly interesting area 
because it centers about the Blackland Prairie which supports a large endemic and 
near-endemic flora, and because the tension zones lying east and west of the prairies 
here are complicated and have been little known in detail. The author’s apologia 
explains a little of why this is still not a complete flora; this is worth quoting in 
some detail because it helps to point up one of the most unusual features of the 
book, namely the extent to which it is truly a production of the author’s own work 
and thought: 


1959] REVIEWS 95 


“Spring Floras are a peculiarly American institution. In almost any other 
country one would get a complete flora, not just a sample. I do not know for 
certain all the reasons for this, but most are not scientific. Expediency, lazi- 
ness, indifference, incompetence, and imitativeness are all involved, along with 
distractions, lack of facilities and time, and the difficulties involved in writ- 
ing a complete work in most parts of the country—which itself has never 
had a whole flora completed for it. This book is largely an abstract of my 
Flora of North Central Texas, which after thirteen years’ labor is still unfin- 
ished. A major part of the delay is due to the fact that I have had during the 
same time to create the elaborate facilities needed to produce such a work. 
Some has been due to my conceit in determining at the start that everything 
would be written by myself, and based on live plants and specimens. Nothing 
was to be farmed out to specialists; only the absolutely unavoidable minimum 
would be taken from publications.” 


A little further along Dr. Shinners hints at the real justification for a Spring 
Flora; in his part of Texas the early spring is the only major season of bloom. The 
growing season may begin in January and reach a climax in February or early March; 
before the beginning of May the hot season has set in and everywhere the sun-dried 
skeletons of plants proclaim the onset of summer. 

The book was written to be used. The keys avoid the use of obscure characters, 
and the terms used in them are simple and clear. In the descriptions an attempt has 
been made to combine brevity with emphasis on diagnostic features. A useful feature 
is the inclusion of common spring-flowering cultivated plants likely to be encoun- 
tered in the area. 

The author warns the reader, however, that he has not prostituted science for the 
sake of popularity: 


“There is no magic which will make it child’s play to find out the names 
of so huge a quantity of variable plants. No real familiarity with them can be 
acquired without using technical terms. No worthwhile list of them is possible 
without using scientific names. If you wish something painless and effortless, 
the pursuit of botany is not for you. Nature gives away few secrets to the lazy, 
and none to the incompetent”. 


The total number of species considered in the Flora is about 1650, including three 
that are described for the first time. The largest families, not surprisingly, are the 
Compositae, Gramineae and Leguminosae. The key to families, if it works, will be 
a joy to the earnest student who has so often been frightened away from regional 
floras by keys that are replete with references to details of ovule structure, placenta- 
tion and the like. Dr. Shinners’ key, appropriately enough in a Spring Flora, is based 
entirely upon floral characters and vegetative characters. Some technically minded 
botanists may shudder at having Galium, Orchidaceae, Lonicera japonica and Ari- 
saema associated in the key under the heading of HERBACEOUS DICOTS. I 
found it hard to use this part of the key. In numerous other trials in other parts of 
the key I found family after family readily identifiable. 

The keys to genera represent something of an achievement, especially in the fam- 
ilies like Cruciferae, Leguminosae, Umbelliferae and Compositae where generic dif- 
ferentiations have heretofore been based almost entirely on characters of the fruits. 
Other authors, of course, have attempted, like Shinners, to construct generic keys 
using floral characters only. The late Norman C. Fassett did this with signal success 
in the Spring Flora of Wisconsin. Fassett was dealing with 15 genera of Legumi- 
nosae; Shinners treats 48. Fassett had 13 genera of Compositae; Shinners keys out 
64 genera without mentioning the mature achenes. Specialists may pick flaws in the 
treatments of their favorite groups, but it seems to me this is a praiseworthy attempt 
to focus attention upon floral characters that have usually been overlooked by 
botanists. 

The author’s concepts of families and genera are for the most part broad ones. 


06 MADRONO [Vol. 15 


He makes of Leguminosae a single family, and Rosaceae another. His treatment of 
the Liliaceae-Amaryllidaceae group, although not out of line with modern thought, 
is not conservative. Perhaps his most startling departure from general practice is in 
the recognition of Chamaesyce as distinct from the rest of Euphorbia. His concept 
of the species (based primarily on consideration of morphology and geographic area, 
as he says on page 469) is a rather inclusive one that in general seems appropriate 
in the flora with which he is dealing. In Crataegus, for example, Shinners recognizes 
14 species, relegating to synonymy (sometimes with a question) approximately as 
many additional species described by earlier authors. In Rubus Shinners recognizes 
5 native species, listing in synonymy 12 additional species described by Bailey. 

Throughout the book an appeal is made to the common-sense of the reader. 
Manufactured “common names” are avoided and indeed anathematized. Comments 
on exceptional nomenclatural situations, taxonomic oddities or complexities, and 
extraordinary or unusual features of the plants themselves, are frequently mentioned. 
Nomenclatural synonyms are included sparingly, chiefly for the sake of clarity when 
there have been recent changes in the application of names. 

Following the formal systematic treatment of the flora, Dr. Shinners has included 
a series of plates illustrative of certain plant families, and finally a series of appendices 
explaining technical terms, the use of keys, the justification for a standard system of 
plant-nomenclature, and the preparation of herbarium specimens. There is a glossary 
and a short discussion of ecology, vegetational types and the botanical history of the 
region. 

In general this is a very commendable book that will command a great deal of 
respect from amateur and professional botanists alike. It should be particularly effec- 
tive as an introduction to Botany when used by the student, in or out of school, who 
is initially attracted by a flowering plant and wants to learn more about its identity 
and its characteristics. As a contribution to floristics the book is patently an intro- 
duction to the author’s projected and much larger work. In its present form, hastily 
gotten out with a regrettably large number of typographical errors, it must be never- 
theless regarded as a remarkably good and scholarly flora of an area where such a 
work was urgently needed. The book begins with a quotation from Gerarde’s Herball 
of 1597, and this review may well close with a line from the Book of Daniel, long 
ago quoted by Olof Swartz in expressing his appreciation to those who contribute to 
floristic knowledge: Plurimi pertransibunt et multiplex erit scientia——Rocrers Mc- 
VaucH, University of Michigan, Ann Arbor. 


NOTES AND NEWS 


ALLIARIA OFFICINALIS ANDRZ. IN OREGON.—In the spring of 1959 a cruciferous 
plant which could not be identified in the regional manuals was collected in an un- 
disturbed forested portion of the Reed College campus in southeastern Portland, 
Oregon (Ornduff 5057). This plant has proved to be the European garlic-mustard, 
Alliaria officinalis Andrz., which has not previously been reported from the Pacific 
states. The Reed College colony of this biennial species was composed of about two 
dozen freely seeding plants. It had not been recorded in a thorough, unpublished 
census of the campus flora made in 1938. Subsequently a second colony of about the 
same size was located in a wooded tract in southwestern Portland. This species is well 
established and widespread in eastern North America and appears well on its way to 
becoming a permanent member of the adventive flora of at least the Portland area as 
well.—RosBertT OrNDUFF, Department of Botany, University of California, Berkeley. 


INFORMATION FOR CONTRIBUTORS 


Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree to bear the cost of the ad- 
ditional pages at the rate of $20 per page. Illustrative materials (includ- 
ing “typographically difficult” matter) in excess of 30 per cent for papers 
up to 10 pages and 20 per cent for longer papers are chargeable to the 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
the author assume the complete cost of publication. — 

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

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


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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


VolumeI,1916-1929 . . . . . $5.00 
Volume IT, 1930-1934. . . . . 5.00 


Volume ITI, 1935-1936 . . . . 5.00 
Volume IV, 1937-1938 . . . . 5.00 
Volume V, 1939-1940. . . . . 5.00 


Volume VI, 1941-1942 . . .. . 5.00 
Volume VII, 1943-1944 . . .. . 5.00 
Volume VIII, 1945-1946 . . . . 5.00 
Volume IX, 1947-1948 . . .. . 6.00 
Volume X, 1949-1950. . . . . 7.00 
Volume XI, 1951-1952. . St, 8.00 
Volume XII, 1953-1954 . . . . 8.00 
Volume XITI, 1955-1956 . . . . 8.00 


Volume XIV, 1957-1958 . . . . 8.00 
Single numbers, . 
Volumesland2 .... . 50 
Succeeding numbers, 
(except Volume XI, No.2). . . 1.00 


Volume XI,No.2 . . . . . 1.50 


The subscription price of MapRONo is $4.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual _ 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


Bot. 


aw 
VOLUME 15, NUMBER 4 OCTOBER, 1959 


Contents 


PAGE 


THE GRASS GENERA ORCUTTIA AND NEOSTAPFIA: A 
STUDY IN HABITAT AND MORPHOLOGICAL SPECIAL- 
r/O7 OF ta! 


IZATION, Beecher Crampton Lf 
THE TAXONOMIC RELATIONSHIP BETWEEN PICEA 

GLAUCA (MoENCcH) Voss AND P. ENGELMANNII 

Parry, 7. M.C. Taylor 111 
FIELD STUDIES OF NATURAL HYBRIDIZATION IN THE 

OREGON SPECIES OF IRIS L. SUBSECTION CALIFOR- 

NICAE Diets, Quentin D. Clarkson LS 
VARIATION PATTERNS IN FOUR CLONES OF MERTEN- 

SIA CILIATA, Jeanette S. Pelton 123 
NEw CoMBINATIONS IN ASTER, Roxana S. Ferris 128 
NOTES AND NEWS 128 


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 

$4.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, 
Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


_» HeErBert 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. Davipson, University of Nebraska, Lincoln. 
Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts. 
Mitprep E. Maruzas, University of California, Los Angeles 24. 

Marion Ownsey, State College of Washington, Pullman. 

Ira L. Wiccrns, Stanford University, Stanford, California. 


Secretary, Editorial Board—ANNETTA CARTER 


Department of Botany, University of California, Berkeley. 


Business Manager and Treasurer—WINSLOW R. Briccs. 
Department of Biology, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: James R. Sweeney, San Francisco State College, San Francisco, Cali- 
fornia. First Vice-president: Baki Kasapligil, Mills College, Oakland, California. 
Second Vice-president: Henry J. Thompson, Department of Botany, University of 
California, Los Angeles, California. Recording Secretary: Mary L. Bowerman, De- 
partment of Botany, University of California, Berkeley, California. Corresponding 
Secretary: Francia Chisaki, Department of Botany, University of California, Berke- 
ley, California. Treasurer: Winslow R. Briggs, Department of Biology, Stanford 
University, Stanford, California. | 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 97 


THE GRASS GENERA ORCUTTIA AND NEOSTAPFIA: A STUDY 
IN HABITAT AND MORPHOLOGICAL SPECIALIZATION 


BEECHER CRAMPTON 


Without a doubt, the endemic genera Orcuttia and Neostapfia are the 
most unusual and rarest of the California grasses. Orcuttia comprises five 
species and two varieties, while Neostapfia is monotypic. All are narrowly 
restricted annuals which develop in the summer beds of vernal pools and 
exhibit peculiar morphological features. Their relationship to other grasses 
is not apparent and quite likely they represent a relict group, the ances- 
tors of which are unknown. Very few collections of them had been made 
up to the time Hoover (3) made known the degree of speciation and geo- 
graphical range of the Orcuttia species. A relatively small number of 
these grasses has been collected since. 

In cooperation with the Department of Agronomy, University of Cali- 
fornia, Davis, and its grass research program, the author was privileged 
to spend part of the summer of 1958 observing the nature of the habitat 
and making extensive collections of these fascinating grasses. The results 
were very gratifying, and the success of this specialized exploration has 
prompted the author to present his findings and to provide additional in- 
formation about the habit, habitat, and morphological development of 
these two unique genera in the California grass flora. 

The author is especially grateful for the helpful suggestions and criti- 
cisms by Dr. G. L. Stebbins, Department of Genetics, and Dr. Jack Major, 
Department of Botany, both of the Davis campus, University of Cali- 
fornia. 


I. THE NATURE OF THE HABITAT 


While many of the Californian grasses are relatively unspecialized as 
to habitat, Orcuttia and Neostapfia are restricted to vernal pools. These 
basins, sometimes called “hog wallows,” are best developed on the rolling 
plains surrounding the Great Valley of California, and to a lesser extent 
on the valley floor. Rainwater collects in them and stagnates during the 
winter and spring, and by late spring or early summer it has completely 
evaporated. The pools are quite variable in extent or area, depending 
upon the terrain. Some are small, shallow, or circular to irregular shape, 
and several meters in diameter. Others may be greatly ramified, with 
numerous islands, while some are quite large, perhaps 500 meters or more 
in diameter, being then classed as intermittent lakes. 

Upon evaporation, the recession of the water from the margin initiates 
development of a unique flora on the muddy strand. Some plant species, 
however, depend upon the standing water for seedling development or 
renewal of perennial growth. It is not unusual to find mature annuals on 
the margins of the pools, while very young plants of the same species are 
in active growth at the edge of the receding water. Such annual plants 


Maprono, Vol. 15, No. 4, pp. 97-128. October 30, 1959, 


SMITHSONIAN oe 
INSTITUTION NOV 2 4959 


98 MADRONO [Vol. 15 


as Allocarya, Pogogyne, Downingia, and Navarretia often show this 
pattern and become generally distributed over the beds. The grasses, 
Deschampsia danthonioides and Hordeum hystrix, are restricted to the 
margins. Of the perennials, Eryngium may be generally distributed or 
marginal, while Eleocharis palustris, Marsilea, and sometimes Damaso- 
nium are in the central or deeper portions of the basins. The distribution 
or occurrence of any one plant species in this environment is quite de- 
pendent upon size, depth, and soil type of the pool and the length of time 
of standing water. 

Orcuttia and Neostapfia require a very special type of vernal pool. The 
concept of “hog-wallow” should be amended to ‘“elephant-wallow” to 
satisfy the environmental demands of these grasses. Primarily, the opti- 
mum size of their vernal basins is about 20-100 meters or more in diam- 
eter or length. Secondly, such basins must neither be drained naturally 
nor artificially, for long periods of standing water are a necessity. Dry, 
unfavorable years result in a paucity of the grasses, while years of heavy 
rainfall such as 1958 (Table 1) are responsible for their peak develop- 
ment. The best stands of either Orcuttia or Neostapfia occur mostly in the 
absence of other vegetation (fig. 1E). The adobe muds in the large vernal 
pools, with their barren, dried, cracked, and often well trampled surfaces, 
are ideal sites. The presence of the ubiquitous vernal pool Eryngium 
vaseyi and the sedge, Eleocharis palustris, restricts the density of Orcuttia 
and Neostapfia. Any dense stand of either of these perennials has rela- 
tively few to none of the annual grasses among them. Barren areas or 
clearings in Eleocharis and thin stands or absence of the Eryngium be- 
come excellent sites for Orcuttia development. Marsilea apparently has 
no deterrent effect on the grasses, as it is a frequent associate of them. 
Relatively few annuals offer competition to Orcuttia or Neostapfia, Bois- 
duvalia being probably the commonest, with occasionally Eremocarpus 
and certain Euphorbia species, but in the main most annuals have dried 
before maximum development of the grasses occurs. 

Recognition of the proper habitat of Orcuttia and Neostapfia simplifies 
collection of these unusual grasses. The most numerous, shallow, early 
drying vernal pools can thus be eliminated as sites for their occurrence, 
and only the large-type pool, with some barren portions, need be con- 
sidered. 

In the past evolutionary history, Orcuttia and Neostapfia probably de- 
veloped as shore or strand grasses on the margin of a sea, such as once 
covered the present Great Valley area. The conversion of the sea to a land 
surface involved only minor modification in the character of the habitat 
so that isolated ponds, developing along the shore of the receding sea, 
became an eventual refuge for these grasses. Perhaps the greatest change 
came in elimination of salts, yet Neostapfia has been found growing on 
alkali in Colusa (the type locality) and Solano counties. Orcuttia mucro- 
nata occurs, as far as is known, only on alkali, but other species of the 
genus favor nearly neutral or perhaps slightly acid soils. It also became 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 99 


necessary for the plants to withstand the period of summer dryness as 
developed in the Great Valley and surrounding areas during the geologic 
changes. 
II. THE AREAS OF COLLECTION 

The pioneer work by Hoover (3) established geographical ranges for the 
species of Orcuttia and enabled the present writer to visit several areas 
of known occurrence of these narrow endemics. Aside from studying habi- 
tats and associations, extensive collections were made of all species except 
Orcuttia californica, which occurs only in southern California. 


A. THE SAN JOAQUIN VALLEY 


1. Stanislaus County. The low hill and rolling plain areas bordering 
the Tuolumne River east of Waterford and Hickman, are especially rich 
in Orcuttia and Neostapfia. The topography favors development of the 
larger-type vernal pool. In this region the upland soils are primarily a 
reddish loam of the San Joaquin series, while the pool beds are light to 
dark grey adobe of the Alamo series, well permeated with iron compounds 
which act as cementing materials. Most of the land is grain-farmed, some 
exists as dry rangeland, while other areas have been converted to irrigated 
pasture. 

In the grainfields between the southern part of Modesto Lake and 
State Highway 132, there are seven large vernal pools bordering on Dien- 
stag and Reservoir roads. All but one of these basins had excellent stands 
of Neostapfia, four of them with abundant Orcuttia pilosa, and a single 
one with Orcuttia californica var. inaequalis. Hoover apparently over- 
looked this particular area of Orcuttia and Neostapfia while collecting in 
1936-1938. The type locality for his Orcuttia pilosa lies five miles farther 
east, the region, as far as determinable, now being irrigated pasture. 

Two vernal basins border Dienstag road, the southern one unequally 
divided by the gravel roadbed (fig. 1-A). The larger and western section 
of this latter pool is about 100 meters across at the widest point and 
nearly 125 meters long. Its basin was entirely covered by a magnificent, 
dense stand of Neostapfia (fig. 1-B). A few scattered clumps of Eleo- 
charis palustris and Sida hederacea were found along the fence adjacent 
to the road, otherwise there was pure Neostafia. The green “‘sheet”’ of this 
grass was in marked contrast to the dry barley stubble on the surrounding 
slopes. 

The eastern and smaller portion of this vernal pool is bordered by hilly 
rangeland. At the lowest point, near the road, there was still some stand- 
ing water on July 28, 1958 (fig. 1-A). Surrounding the murky water, ina 
semicircle, was a wide band of barren, dark-grey, well-trampled mud. On 
the periphery of the mud, a pale green band of Neostapfia was developing 
on the cracking, drying, grey adobe. Several resident dairy heifers were 
intermittently, but actively grazing the grass! 

The second vernal pool along Dienstag road is about 100-125 meters 
long and perhaps 75 meters wide. The bed was a solid stand of Neostapfia 


100 MADRONO [Vol. 15 


with an abundance of Orcuttia pilosa along the margin. This unique pat- 
tern invariably occurred whenever the two grasses were associated. 

Three vernal pools along and south of Reservoir Road were replete 
with Neostapfia and Orcuttia pilosa as the sole or major occupants. One 
pool, however, contained mostly Orcuttia californica var. inaequalis, very 
little Orcuttia pilosa, and no Neostapfia. Another pool was of unique 
shape, nearly 150 meters long and about 10 meters wide, its bed a pure, 
solid stand of Neostapfia. 

The barley field operations in this area apparently do not affect the 
successful development of these peculiar grasses in their habitat. Un- 
doubtedly the planting machinery disturbs the vernal pool beds in the 
fall, yet after that they remain undisturbed until harvest, at which time 
Orcuttia and Neostapfia are reaching maximum development. 


A large playa, about 150 meters or so in diameter, is situated in a small 
valley leading out of the hilly country three miles east of Hickman. 
Though the surrounding areas are grain-farmed, the vegetation of this 
bed differs considerably from those of the previous area. Neostapfia is 
restricted to small patches, while Orcuttia californica var. inaequalis is 
more widely dispersed. Active competition is afforded by Centromadia 
fitchu, Boisduvalia, and Eryngium, with an abundance of dried Allo- 
carya, Downingia, and Navarretia leucocephala. The more barren areas 
on the dried and cracked lead-grey adobe support the best stands of both 
Orcuttia and Neostapfia. 

An extensive reservoir or permanent lake! (fig. 1-C) is located on the 
Loren Rouse Ranch six to seven miles east of Hickman, along the road 
to La Grange. Neostapfia and Orcuttia pilosa occur abundantly along the 
strand, which is subject to vernal flooding and summer drying. This lake 
has quite likely been developed from a large vernal playa which originally 
was populated by these two grasses, since the margins of man-made res- 
ervoirs are not ordinarily sites for development of either of these grass 
genera. If planted, they might very well become adapted to reservoir 
strands. 

Summer fallowing of the hilly grainland to the south had closely ap- 
proached, nevertheless had not disturbed the strand. Dry rangeland and 
irrigated pasture surround the lake on the north and east, respectively, 
though neither operation had disturbed the habitat. 


The old Paulsell warehouse is located along the old Sierra railroad 
northeast of Waterford. The land area is now largely devoted to irri- 
gated pastures except for the hilly, dry rangeland to the north. The dis- 
tinctive Orcuttia greenei was found in a large, undisturbed playa at the 
southwestern base of a prominent knob-like hill. The playa basin is per- 
haps 200-250 meters in diameter and composed of black, deeply-cracked 
adobe. Aside from the dense, dried Allocarya cover, Eryngium and Ere- 
mocar pus were dominant, with some Eleocharis palustris and Boisduvalia. 


1 Since the above was written, it has been learned that this is a natural pool. 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 101 


A B 


barley-farmed area south of Modesto Lake, east of Waterford, Stanislaus County. 
Dienstag Road divides the basin. (B) The author in the solid stand of Neostapfa 
present in the far portion of A. (C) Lake on the Rouse Ranch east of Hickman, 
Stanislaus County, the strand of which has an abundance of Neostapfia and Orcuttia 
pilosa. (D) Goose Valley, Shasta County, the type locality of Orcuttia tenuis. The 
author collected the grass in the ditchbed at the right. (E) Neostapfia growing on a 
typical soil of a vernal pool bed. Note the cracks and absence of other vegetation. 
Eremocarpus lines the bed in the background. (F) Habitat of Orcuttia mucronata. 
Frankenia and the prostrate Eryngium aristulatum are the only associates. 


102 MADRONO [Vol. 15 


The Orcuttia was not abundant, but occurred mostly as scattered plants 
in areas containing the least Eryngium and Eremocarpus. 

2. Merced County. In the vicinity of the old Ryer station, located 6.5 
miles south of Montpelier, there are two large-type vernal pools contain- 
ing Orcuttia californica var. inaequalis. The rather uniform rolling hill 
country in this area is all grain-farmed, with rather numerous, smaller- 
type vernal pools. The large playa to the west is basically a lead-grey 
colored adobe, well populated with Centromadia fitchiu, Centromadia 
pungens, Sida hederacea, Boisduvalia and marginal Eryngium as well as 
the dried Allocarya, Downingia and Navarretia leucocephala. The Orcut- 
tia was most abundant in areas of the least competition. 

The eastern vernal pool harbored a dense stand of Orcuttia californica 
var. inaequalis to the near exclusion of other plants. Apparently the 
basin had been completely cultivated during barley planting, for faint 
furrows were evident over the pool bed. The Orcuttia was exceedingly 
robust, some of the plants being 12—15 cm. high and with as many as 60 
culms, a development far in excess of those in the neighboring playa. 
Aside from dried Allocarya, only a few scattered plants of Centromadia 
and Eremocarpus were present. 

North of Legrand, in typical rolling plain rangeland, is a large playa 
perhaps 300-350 meters in diameter. The soil of the basin is a black 
adobe, becoming exceedingly deeply cracked when dry. Some of the cracks 
extended from three-fourths to nearly a meter in depth and from 5—10 
cm. across at their aperture. A dense stand of Evemocarpus covered the 
major portion of the bed, the whole appearing from a distance as a silver- 
grey “lake.”’ Large specimens of Orcuttia greenei were numerous in areas 
of the least Evemocarpus, though scattered individuals occurred through- 
out the playa. Boisduvalia was perhaps the closest and only other 
associate. 


B. THE SACRAMENTO VALLEY AND NORTHERN CALIFORNIA 


1. Solano County. A magnificent alkaline, intermittent lake is situated 
in a rolling plain area twelve miles south of Dixon, Solano County. It is 
shallow, has no drainage, and is about 500 meters in diameter. The pre- 
dominant vegetation in the lake basin is Frankenia grandifolia, Cressa 
truxillensis, Sida hederacea and Eryngium aristulatum, with some patches 
of Eleocharis palustris. The marginal strand is largely Distichlis, Lippia 
and Navarretia bakeri. The dried crust varies from a uniform, glaring- 
white pavement to a tan and cracked surface. On the latter type, Neos- 
tapfia and an undescribed species of Orcuttia grew in association with 
Frankenia and Eryngium (fig. 1-F). Both grasses, at their best, were rare 
occupying an exceedingly small area of about 15-20 meters in diameter. 
The Orcuttia will be described later in this paper as Orcuttia mucronata, 
while the occurrence of Neostapfia here represents a new area for Cali- 
fornia. 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 103 


2. Sacramento County. According to Hoover (3), Orcuttia californica 
var. viscida is not known to occur outside of Sacramento County. The 
Orangevale area now shows a distinct waning of the grass, and in all prob- 
ability the habitat will be eliminated by residential development. 

On the higher, rolling plains north of Sloughhouse, one large vernal pool 
and several smaller, but deep ones, support fairly good stands of this at- 
tractive Orcuttia and represent its type locality. It was conspicuous along 
the margins and in some parts of the bed of the large pool. This greater 
basin, 100 or more meters in length and with some ramification, is largely 
a barren and stony bed. The cracked, lead-grey, adobe soil is well supplied 
with iron concretions, apparently an indication of the type of soil several 
Orcuttia species prefer. Some patches of Eleocharis palustris and Eryn- 
gium were present, but these species were certainly not diffuse in any area 
of the pool. 

3. Butte County. A single vernal pool on the rolling plains about ten 
miles southeast of Chico was well supplied with Orcuttia greenei and 
probably represents the type locality of this species. The basin, 75—100 
meters in diameter, is composed of a grey-black, crumbly, stony, and 
pebbly soil. Over the major portion of the pool the marginal vegetation 
of Eryngium and Eremocarpus gave way to an abundance of the Orcuttia 
and to the prostrate annual Euphorbia hoovert. 

4. Tehama County. Orcuttia tenuis is endemic to northern California, 
preferring soils probably derived from volcanic substrates. Only certain 
areas of Tehama, Shasta, and Lake counties have the proper environ- 
mental conditions, each of them in entirely different settings. 

In Tehama County, on a rocky, volcanic plateau eleven miles northeast 
of Red Bluff, lies an extensive dry lake known as Hog Lake. The basin is 
perhaps 300 meters wide and 1500 meters long, surrounded by mixed, 
open grassland and blueoak woodland. The western shallow portion and 
the marginal strand were largely populated by Eryngium, while the deeper 
portions contained Eleocharis palustris and Damasonium. The best de- 
velopment of Orcuttia tenuis occurred in the barren, stony areas among 
patches of the Eleocharis and to a lesser extent Damasonium. Although 
the grass was rather uniformly distributed among Eryngium, it had dried 
too quickly here for proper seed set. The dried, leached grass stood in 
marked contrast to the grey-green plants in anthesis and fruit matura- 
tion which occurred in the deeper and barren areas of the lake basin. 

5. Shasta County. Orcuttia tenuis occurs in abundance in a series of 
vernal pools around the Redding Municipal airport three to five miles 
north of Anderson, the area being known as Stillwater Plains. The south- 
ern series is largely in open grassland, while those to the west and north 
of the airport are surrounded by oak, digger-pine, and manzanita. The 
typical floristic association pattern of all of the pools can be illustrated 
by that in a single pool, a circular basin about 125 meters in diameter. 
Here Eryngium is marginal, but the majority of the bed contains Eleo- 
charis palustris with intermittent patches of Marsilea. Although some of 


104 MADRONO [Vol. 15 


the Orcuttia grew among the sedge, it was more common in the barren 
situations. None of the grass could be found among the Eryngium. 

Two shallow and smaller pools associated with the open grassland had 
evidently dried too quickly, for the grass was leached and the spikelets 
were without seed. A small but deep ditch along the road and continuous 
with one of these basins contained green and properly maturing Orcuttia. 

The type locality of Orcuttia tenuis is Goose Valley, north of Burney, 
Shasta County. The type collection was made here by Alice Eastwood in 
1912, and since that time no other collections are known to have been 
made. This mountain valley is around 3500 feet in elevation, surrounded 
by mixed conifer forest, and is primarily meadowland with some dry-lake 
habitats in the northern section. Most of the valley is now largely culti- 
vated, either as permanent pasture or cropland. A series of canals which 
effectively drain many areas is consequently disastrous to the survival of 
the Orcuttia. Fortunately, the grass was located in the nearly barren bed 
of an old ditch skirting a northeast portion of the valley (fig. 1-D). Aside 
from small patches of Eleocharis palustris and Damasonium along the 
edges, and some scattered Downingia and Boisduvalia in the bed, Orcut- 
tia tenuis remained the dominant plant of the basin. 

6. Lake County. The occurrence of Orcuttia tenuis in the Coast 
Ranges was first made known by Milo Baker, eminent Santa Rosa bota- 
nist, who collected the grass on the north shore of Bogg’s Lake, Lake 
County. This lake is situated in a yellow pine forest flat on the northwest 
slope of Mount Hannah. The basin is around 1650 meters in diameter, 
contains water the year round, yet is provided with an adequate strand 
for the development of many unique and unusual plants. 

A visit to the lake in August, 1958, proved disappointing because of the 
high water level and resultant flooded strand. Previous to 1958 the author 
had collected the grass on the southwest margin among Eleocharis palus- 
tris and Eryngium though it was exceedingly rare, requiring consider- 
able search to locate perhaps two dozen plants. 


III. YEARLY FLUCTUATIONS IN ABUNDANCE 


The year 1958 was an ideal one for maximum development in the 
stands of Orcuttia and Neostapfia. In all of the areas visited, with one 
exception, these grasses were abundant, the most remarkable being the 
magnificent stand of Neostapfia just south of Modesto Lake. 

What of other years? Even though the grasses must, in the overall 
floristic analysis, be regarded as rare and narrowly restricted endemics, 
why have they not been collected more often? The answer undoubtedly 
lies in their fluctuations in abundance from year to year. Unfavorable, 
dry years would be associated with poor development or scattered stands 
and earlier maturity. A shallow depth of water evaporating rapidly dur- 
ing the spring might prove quite disastrous to the annual stand of the 
grasses, even though they are situated in the required habitat. Such an 
environmental stress would antagonize the genetically-fixed, summer- 


1959 | CRAMPTON: ORCUTTIA AND NEOSTAPFIA 105 


maturation character of Orcuttia and Neostapfia, resulting in heavy seed- 
ling mortality and consequent rarity of plants in the habitat. Further- 
more, supposing good germination in water or on the mud of the vernal 
basins, too rapid a drying might cause premature flowering and poor to 
no seed set. . 

How, then, is perpetuation of the grasses maintained over unfavorable 
periods? Presumably the large seed production effected in such a year as 
1958 might be adequate for the next several years, since there is a sug- 
gestion that the seed remain viable over many years and that a prolonged 
period of dormancy may be necessary before germination occurs. This 
supposition is drawn, but perhaps too hastily, from an attempt to germi- 
nate Orcuttia seed. Orcuttia pilosa grains collected in August, 1957, were 
placed on a blotter in a petri dish on January 31, 1958, kept at room tem- 
perature, and under continuous moisture. By September 1, 1958, no ger- 
mination had occurred, though very few of them had succumbed to mold. 
Aside from the moisture, this artificial medium lacks all of the elements 
of the natural ones. 


TABLE 1. Annual Precipitation (July to June) of four reporting stations in the 
Great Valley of California* 


Season Merced Modesto Sacramento Red Bluff 
1931-1952 (average) 12.35 12.44 16.68 235 
1955-1956 15.42 15.62 25.53 28.53 
1956-1957 7.40 8.63 13.78 12225 
1957-1958 25.63 23.04 28.70 38.03 


* Data obtained from USDA Weather Bureau-Climatological Data, California 
section. 


Table 1 shows some climatological data for most of the areas where 
the writer collected Neostapfia and Orcuttia in 1958. The amount of rain- 
fall during the 1957-1958 season supports the argument that abundance 
of the two grass genera is dependent on the amount of precipitation. In 
all probability there was generally minimum development of the grasses 
during the 1956-1957 season. Dr. G. L. Stebbins, department of Genet- 
ics, and Dr. and Mrs. Louis Mann, department of Vegetable Crops, 
University of California, Davis campus, visited the vernal pool as shown 
in Figure 1-A, in the latter part of June 1957. They could scarcely find 
any Neostapfia, and what few plants were found were thoroughly dried 
and mostly dessicated. The remarkable solid stand of the grass present in 
July, 1958, could not have developed from the 1957 crop. Undoubtedly a 
larger seed crop had been produced in previous and more optimum years. 

Without doubt, judging by observation of the excellent stands of both 
grass genera in 1958, the amount of rainfall and consequent depth of 
standing water in the vernal pools is most critical in their life cycle and 
reflects their yearly abundance. 

The collections by Hoover (3) and others have pretty well established 
the geographical range of the Orcuttia species and Neostapfia. It remains 


106 MADRONO [Vol. 15 


to round out their distribution by detailed collections of the two genera 
from year to year. 

The primary purpose of this paper has been to stimulate interest in 
these most unusual and scientifically interesting grasses and make known 
their specific type of habitat and specific locations. The progress or de- 
cline of the grasses in any one area can be evaluated by the frequency 
of collection. This concern by the author is not without foundation. With 
the steady increase in California’s population and with the resulting modi- 
fied land manipulation that must occur, there is a good possibility of the 
destruction of many habitats, and this may result in the extinction of 
these grass species or their varieties. 


Fic. 2. Orcuttia mucronata sp. nov. Plant from type specimen, approximately 
natural size. 


ITV. AN UNDESCRIBED SPECIES OF ORCUTTIA 


The extensive collections of Orcuttia and Neostapfia made during July 
and August, 1958, uncovered a most unusual and distinctive Orcuttia 
species on the western side of the lower Sacramento Valley. The grass dif- 
fers rather considerably from most other Orcuttia species, and a more 
radical treatment might erect a new genus. However, until all details of 
the plant can be evaluated it seems expedient to assign the grass to 
Orcuttia. 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 107 
ws 


Fic. 3. Orcuttia mucronata sp. nov. (A) spikelet, * 5.5. (B) floret, K 7.3. (C) 
caryopsis, X 7.3. (D) lodicule fused to the palea, x 18. 


Orcuttia mucronata sp. nov. 

Planta annua aestivalis aromatica flavoviridis folia rigida extrinsecus 
curvata 1-4 cm. longa inflorescentia racemiformis basi tantum inclusa 
spiculis 7-19 spiralibus 5-10 floribus nec supra nec infra glumas disar- 
ticulantibus lemmis 5—7 mm. longis apice mucro unico terminatis dentibus 
lateralibus suppressis palea apice trilobata lobis dentatis lodiculis duabus 
hyalinis enervatis paleae adnatis. 

Summer annual; pilose throughout, yellow-green, aromatic. Culms few 
to many, decumbent, 2.5—12 cm. high; leaves eligulate, viscid, 1-4 cm. 
long, somewhat rigid, curved outward tapering to a fine point; inflores- 
cence a raceme, 1.5—6 cm. long, partially included, spikelets 7-19 in 
number, spirally arranged; spikelets 7-13 mm. long, 5—10-flowered, no 
disarticulation between the florets or below spikelet; glumes 4—7 mm. 
long, unequal, nearly approximate, lanceolate, the apex pilose, mostly 
awn-pointed or occasionally with 1 or 2 lateral teeth; lemmas coriaceous, 
5—7 mm. long, the upper portion excurved, sparsely pilose, scabrous, vis- 
cid and light green, the lower portion short-hairy and whitish, the apex 
obtuse with a median mucro, 0.5 to nearly 1 mm. long, the lateral teeth 
suppressed, the margin appearing merely erose; palea shorter than the 
lemma, the apex 3-lobed, the lobes toothed, pubescent towards the margin 
near the apex; lodicules 2, fused to the palea, hyaline, nerveless, about 
0.25 mm. wide and 0.5 mm. long; anthers yellow fading pinkish; caryop- 
sis oblong, flattened, 3 mm. long, the embryo 1.5—2.0 mm. long along one 
side of the grain (figs. 2 and 3). 


108 MADRONO [Vol. 15 


Type. Alkaline lake, 12 miles due south of Dixon, Solano County, Cali- 
fornia, August 1, 1958, Crampton 5057, AHUC. (Isotypes: UC, JEPS, 
DS, US, CAS, K.) Known only from a single dry lake at the type locality. 

Other collections (Crampton 5011, 5059, 5093, and 5113), all collected 
in this single locality, represent a series of developmental stages. Orcuttia 
mucronata is, at the most, rare. Only three patches of the grass, roughly 
3-8 meters in diameter have been found over the large expanse of the 
lake bed. These small populations apparently occur in the deeper portions 
and on a cracked alkali with a brownish film over its surface. The white, 
smooth alkali pavement, characteristic of much of the lake harbored none 
of the grass. The soil type is classified as the Lindsey clay loam series. 
The soil survey of the Suisun area (1) indicates that the surface of this 
soil is a dull, dark or brownish grey material with a large proportion of 
fine to very fine sand. It deflocculates and when dry it becomes hard and 
baked. Organic matter is low, and most areas contain alkali. During the 
rainy season such areas become ponded for weeks or months at a time, 
the soil taking water slowly and having a high water-holding capacity. 
The surface layer extends from 8-10 inches in depth, and certain areas 
contain lime. The subsoil is heavy textured and compact, with some cal- 
careous areas. 

The soil at the area where the type plants of Orcuttia mucronata grow 
shows a pH of 8.0 on saturated paste as determined by the Agronomy 
Soils Laboratory. 


V. MORPHOLOGICAL CHARACTERISTICS 


The classical alliance of Orcuttia and Neostapfia to the tribe Festuceae, 
on the basis of gross spikelet morphology as outlined by Hitchcock and 
Chase (2), is inadequate in establishing relationship of the two genera 
to other grasses. A review of the salient features of both Orcuttia and 
Neostapfia indicates that they are not closely related to any members of 
the Festuceae. 

One of the most conspicuous features of these grasses is the viscid se- 
cretion on all aerial parts of the plant whether young or mature. At first 
the secretion is glistening and watery, but towards maturity it becomes 
a thicker, denser, usually brownish exudate. In Neostapfia, distinctive 
scale-like raised glands on the lemma nerves and leaves contribute to the 
viscidity. In association with the copious secretion, a peculiar odor ema- 
nates from either fresh or dry material in any stage of development. The 
viscidity undoubtediy conserves plant moisture during the warm late 
spring and hot summer temperatures, while the aromatic habit may serve 
to reduce or repel animal depredation. In some of the collection areas 
grasshoppers were in abundance, but the green Orcuttia or Neostapfa 
plants were unaffected by the voracious insects. 

In marked contrast to the Festuceae, Orcuttia and Neostapfia are 
summer-maturing annuals that occupy a highly specific type of habitat. 
Their coloration varies from a pale- to grey-green, and all are hairy. In 


1959] CRAMPTON: ORCUTTIA AND NEOSTAPFIA 109 


Neostapfia, however, hairs on the foliage are sparser, and very minute. 

The nature of the foliage is distinctive, for in both genera there is no 
differentiation into sheath and blade, and consequently a true ligule and 
well-defined collar are absent. In all but Neostapfia and Orcuttia mucro- 
nata there is a definite abscission of the blade portion of the leaf. This 
deciduous character is not apparent until the leaves are dry, although in 
some instances the point of abscission is faintly visible in green tissue and 
might correspond to a ‘‘collar.” The hairiness on the upper surface of the 
Orcuttia blades terminates abruptly at the fracture region, so that the 
resultant line of hairs might be construed as ligular. Otherwise the leaves 
of both genera loosely envelope the culms. 

The culms of Orcuttia and Neostapfia are solid, the internodes being 
filled with pith. In the larger plants of Neostapfia, the decumbent culms 
form a zig-zag pattern, thereby providing additional support of the plant’s 
superstructure. Branching is basal in all of the grasses except Orcuttia 
tenuis, which is literally “top-heavy.” A single, filiform, culm, often with 
adventitious roots from the lower nodes, supports the entire ramification 
above, and when excessively branched the whole plant becomes decum- 
bent. 

The mature inflorescences in both genera are exserted, except for 
Orcuttia mucronata, which is partially included. Neostapfia has a cylin- 
drical, spike-like panicle, the terminal portion differing from the rest of 
the rachis in bearing small, closely appressed, lanceolate bracts. In Orcut- 
tia the inflorescences are spikes or racemes, the shape varying among the 
species. The spikes of Orcuttia pilosa and Orcuttia tenuis are more or 
less elongated, with mostly distichous spikelets. A spiral arrangement of 
spikelets exists in Orcuttia greenei and Orcuttia mucronata, with the in- 
florescence somewhat elongate. In Orcuttia californica var. inaequalis and 
Orcuttia californica var. viscida the spike is sub-capitate, with a secund 
arrangement of spikelets. 

The spikelets in Neostapfia are most unusual. They are without glumes, 
and the florets are so arranged as to suggest a trimerous cluster of spike- 
lets. The florets are secund, the two lower ones divergent, the three or four 
upper ones closely imbricate. Disarticulation normally occurs between the 
florets, but occasionally below the spikelets, the rachis being continuous. 
In Orcuttia the spikelets are solitary, the floret number variable from 
5-10 in Orcuttia mucronata to 10-30-, or even 40 in other species. Except 
for Orcuttia tenuis, there is little or no disarticulation of the florets, the 
whole inflorescence largely non-shattering. The culms of Orcuttia greenei 
are excessively fragile at the base, and the mature seed heads are readily 
deposited upon the surface of, or cracks in, the adobe soil. 

The form of the lemma apex is an excellent diagnostic character for 
both genera. Along with the form of the inflorescence, it is employed con- 
sistently in identification keys and will not be discussed here. The lemma 
texture in Neostapfia is papery except for the tough nerves, while the 
Orcuttia lemmas are coriaceous. 


110 MADRONO [Vol. 15 


Mature grains of both genera were examined from the material col- 
lected during 1958. In common they show: a loose enclosure between the 
lemma and palea, lateral flattening with the embryo extending along one 
side, a large basal hilum, persistent style base at the apex, and compound 
starch grains in the endosperm. The NVeostapfia grains are obovate, com- 
pletely viscid, and dark brown in color. Those of Orcuttia are oblong, not 
or scarcely viscid, and the embryo and hilum are conspicuously brownish 
in contrast to the light-colored endosperm. 

Lodicules are apparently absent in all except Orcuttia mucronata, and 
here their unique appearance (fig. 3-D) suggests that a sectional division 
should be made to accommodate this species in Orcuttia; or possibly the 
lodicules along with other characters suggest that generic rank is merited. 

The similarity of habitats, growth habit, and convergence of some mor- 
phological features indicate a rather close relationship between Orcuttia 
and Neostapfia, while their affinity to other grasses is not readily appar- 
ent. They should be removed from the Festuceae and might well be con- 
sidered as a separate tribe. 


VI. SUMMARY 


1. Orcuttia and Neostapfia are restricted to a special type of vernal pool 
or “hog-wallow” which limits their geographic distribution. The associa- 
tion patterns of both Orcuttia and Neostapfia are truly remarkable and 
would constitute excellent material for detailed ecological studies. In cer- 
tain areas, as revealed by the preceding account, Neostapfia occurs with 
only certain species of Orcuttia or stands alone. Likewise, only rarely are 
two species of Orcuttia associated, but if so, one of them is represented 
by only a few scattered individuals. All, however, demand the relatively 
large vernal pool beds for successful perpetuation. 

2. It is suggested that their varying abundance from year to year is 
dependent upon the amount of rainfall and upon a long period of seed 
viability. 

3. Morphological developments are a natural consequence of the envi- 
ronment with specializations directed towards viscidity, aroma, pubes- 
cence, and non-shattering inflorescences. Both genera are closely related, 
undoubtedly forming a natural grouping among the grasses, though their 
relationship to others is rather obscure. 

4. A new species of Orcuttia from Solano County is described and illus- 


trated. Agronomy Herbarium 


Department of Agronomy 
University of California, Davis 


BIBLIOGRAPHY 


1. CARPENTER, E. J. and S. W. Crossy. 1930. Soil Survey of the Suisun area. U.S. 
Dept. Agric. Bur. Chem. & Soils. Series 1930, Number 18: p. 33. 

2. Hitcucock, A. S. and AcNEs CHAsE. 1950. Manual of the grasses of the United 
States. 2nd Edition. USDA Misc. Publ. 100. 

3. Hoover, Ropert F. 1941. The genus Orcuttia. Bull. Torrey Club 68 (3) :149-156. 


1959] TAYLOR: PICEA 111 


THE TAXONOMIC RELATIONSHIP BETWEEN PICEA 
GLAUCA (MOENCH) VOSS AND P. ENGELMANNII PARRY’ 


T. M. C. TAyLor 


For many years botanists and foresters have been puzzled and frus- 
trated by the spruce complex in British Columbia, particularly the plexus 
centering around Picea glauca (Moench) Voss (White spruce) and P. en- 
gelmannii Parry (Engelmann’s spruce). In the northern parts of the 
province P. glauca appears in its typical form and is a clear-cut entity. At 
higher altitudes in the southern interior of the province P. engelmannit 
may be found in equally characteristic form. Unfortunately, between 
these two easily distinguishable extremes there is a great range of inter- 
mediates. The taxonomic, and hence the nomenclatural, disposition of 
these intermediates is the substance of the present paper. 

Materials studied were collected largely at Banff, in the Upper Colum- 
bia Valley, and in the Cranbrook-Moyie Lake area. This southeastern 
region of the province and adjacent Alberta was selected because here 
the problem raised by the intermediate forms of spruce is particularly 
acute. Collections were made with the primary purpose of gaining a statis- 
tical picture of the variability between individual trees of certain mensural 
characters of leaves and cones. It was felt that only when the extent of 
variation within the individual was established, could one proceed with 
confidence to generalizing from small samples drawn from many trees. 

Materials were collected from about seventy randomly selected trees. 
In case the degree of shading might produce constant differences, cones 
and twigs with needles were taken from both the north and south side of 
trees. Needles were also collected from both vegetative and reproductive 
shoots. Statistical analysis showed no significant difference between sam- 
ples from the north and south side, nor between needles from reproduc- 
tive and vegetative twigs. In the present report therefore this distinction 
is not maintained. 

The difficulty of making accurate measurements of curved leaves was 
overcome by boiling them for five minutes to render them pliable so that 
they could be straightened out. Spreading of the scales in dry cones also 
offered complications for measurement but these complications were also 
overcome by boiling the cones until they sank. By this time the scales had 
contracted and were closely appressed in the cone. It was established that 
prolonged boiling produced no further change in dimensions. 

Mention may be made of the findings on intra-tree variability. Needle 
lengths, with means of the order of 13.5 mm. and standard deviations of 
about 2 mm. showed coefficients of variability ranging from about 11 to 
20. Cone diameters with means of the order of 13.5 mm. have standard 
deviations of slightly over 1 mm. with a range in the coefficients of vari- 


1 This study was supported in considerable measure by financial assistance from 
the National Research Council of Canada to which body grateful acknowledgment 
is made. 


112 MADRONO [Vol. 15 


ability from 6 to 10. Cone lengths were about 42 mm., with standard de- 
viations about 3.5 mm. and variability with coefficients about the same 
as for diameters. 

One can conclude then that needle lengths are very variable on the same 
tree and that a large number would have to be measured in order to obtain 
a Statistically satisfactory mean value. Cones, on the other hand, are much 
more uniform in dimensions and so smaller samples per tree would be 
acceptable. 

The most obvious difference between Engelmann’s and White spruce is 
in the cone scale. Those of the former are thin and somewhat papery, 
wedge-shaped with wavy to erose margins, and commonly erose to trun- 
cate at the apex. The scales of the White spruce on the other hand, are 
obovate-triangular, somewhat stiff with entire margins, the apex being 
rounded or somewhat flattened, not erose. It was possible to interpolate 
three classes of intermediates between these extremes. In the accom- 
panying tables the five cone type classes are numbered in Roman numer- 
als with P. engemannii I and P. glauca V. Table 1 shows the means 
and standard deviations of cone diameters and lengths for the five cone 
type classes. The differences between the means of diameter are not re- 
liable while the differences in mean length of types I and V is reliable at 
the level of .01. The erratic means of length for types in II, III and IV 
are presumably due to inadequacy of the samples. This, however, is a 
matter that requires further investigation. 


TABLE 1. Relationship between cone dimensions and cone type in Picea engel- 
mannii, P. glauca, and intermediates.* 


Cone type Diameter Length 
M SD N n M SD N n 
I 13.7 13 14 521 45.0 Sale 14 518 
II 14.0 1.12 12 349 46.1 6.57 11 328 
III 13.4 1,03 6 207 41.8 Si 6 199 
IV 13.6 1.28 6 368 43.1 3.00 6 377 
V 13.4 127 21 1449 41.4 3.96 vay 1491 


*T represents P. engelmannii, V, “P. glauca,” I-IV, intermediates. M = mean 
(in mm.), SD = standard deviation (in mm.), N = number of trees sampled, n = 
number of cones measured. 


Analysis of needle length showed no significant differences between the 
means for the different cone types. Engelmann spruce needles, however, 
tended to be straight and slender, acuminate and somewhat square in 
cross-section, while those of White spruce were firmer, often curved and 
rounded at the apex, tending to be dorsiventrally flattened or triangular 
in section. Needles with these characteristics in other combinations were 
classed as “intermediates.” Table 2 shows the relationship found between 
leaf type and cone type. It is apparent that foliage characters are not cor- 
related with cone types and are probably due to the independent segrega- 
tion of several genes. 


1959] TAYLOR: PICEA 113 


TABLE 2. Relationship between leaf type and cone type in Picea engelmannii, 
P. glauca, and intermediates.* 


Cone types 
Leaf types I II Ill IV V N 
‘engelmannil’ 9 7 N) ae a 21 
intermediate 12 4 4 7 8 30 
‘glauca’ ie oe ae 1 13 14 
Totals Zi 11 9 8 Da 70 


* T represents P. engelmannii, V, P. glauca, II-IV, intermediates. N = number of 
trees sampled. 


The relationship between the indumentum of the twigs and cone types 
was also studied. Twigs of White spruce are characteristically glabrous, 
while those of Engelmann’s spruce have a short, crisp pubescence. In 
Table 3 it can be seen that all cone types may be borne on trees with 
pubescent twigs, except that cone type V has an equal chance of being 
borne on a tree with glabrous twigs. One can speculate that the develop- 
ment of indumentum is controlled by a single pair of genes and that 
“pubescence” is dominant. 


TaBLeE 3. Relationship between indumentum of twigs and cone in Picea engel- 
mannii, P. glauca, and intermediates.* 


Cone types 
Twigs I II III IV V N 
Pubescent tag 10 9 8 11 59 
Glabrous re 1 = ae 10 i 
Totals ZA 11 9 8 21 70 


*T represents P. engelmannii-type, V, P. glauca-type, II-IV, intermediates. 
N = number of trees sampled. 


It is apparent that White and Engelmann’s spruce are very much alike 
and that even such diagnostic features as mean cone length, and shape and 
character of the scales are merely the extremes of a series of intermediates. 
Both ‘engelmanii’ and ‘glauca’ needle types are found on trees with ‘inter- 
mediate’ type cones and ‘intermediate’ type needles may be associated 
with any type of cone. Glabrous twigs have only been found on trees with 
‘glauca’ cones but, on the other hand, there is an equal chance that the 
twigs on such trees will be pubescent. On several occasions in the past it 
has been commercially important to try to distinguish lumber cut from 
these two spruces. No differentiating histological details have been found 
and Barton and Gardner (1957), using partition chromatography in addi- 
tion to infra-red and ultra-violet spectrographic methods, failed to estab- 
lish any chemical differences between the woods. 

Wright (1955) has examined in considerable detail the question of in- 
terspecific hybrids in Picea. In his paper he attempts “‘to correlate species 


114 MADRONO [Vol. 15 


crossability with geographic distribution, morphology, and phylogeny.” 
He examined thirty-one species with respect to fifty-one characters that 
show differences between some or other of these species. The majority of 
the characters, of course, do not lend themselves to measurement and are 
to quite an extent subjective. It is regrettable that Wright’s observations 
were not analyzed statistically. Had they been, the relationship between 
P. glauca and P. engelmannii would likely have appeared very much closer 
than he concludes. 

Wright (1.c.) has made some investigation of hybridizing between 
White and Engelmann spruce. Using a hybridity index, he shows that very 
considerable introgression is taking place over a wide area extending from 
latitude 51° to 58° N. and between longitude 109° to 124° W. That these 
intermediates are almost certainly of hybrid origin is borne out by his 
report of successful reciprocal crosses between the two species. 

Garman (1957) has made a detailed morphological and distributional 
study of spruce populations in British Columbia. Using a “morphological 
index” on which pure P. glauca rates 6 and pure P. engelmannii rates 18, 
he shows the very extensive geographic area occupied by intermediates 
on this scale. This index has apparently proven useful in distinguishing 
populations, but its value would be greatly enhanced if the mensurable 
characters had been treated statistically. Only average values and ranges 
of measurement are given and, as the latter frequently overlap, their sig- 
nificance cannot be evaluated. 

In the opinion of the present author, the phylogenetic relationship be- 
tween P. glauca and P. engelmannii is best indicated by regarding them 
as subspecies of a single species and so the following changes in status 
are proposed. According to the provisions of Article 57 of the International 
Code of Botanical Nomenclature (Lanjouw 1956) Picea glauca (Moench) 
Voss, which has priority, becomes the legitimate name when the two 
species are combined into one. 


PIcEA GLAUCA (Moench) Voss subsp. glauca 
Abies canadensis Mill. Gard. Dict. Ed. 8, Abies No. 4, 1768; nomen confusum 
Pinus glauca Moench, Verzeichn. Baeume Weissenst. 73. 1785 
Pinus alba Ait. Hort. Kew. 3:371. 1789 
Picea laxa Sargent, Gard. and For. 2:496. 1889 
Picea glauca (Moench) Voss, Mitt. Deutsch. Dendr. Ges. 1907:93. 1907 
Picea glauca var. porsildii Raup, Sargentia 6:102. 1947 


PicEA GLAUCA (Moench) Voss subsp. engelmannii (Parry) stat. nov. 
Abies engelmanni Parry, Trans. Acad. St. Louis 2:122. 1863; nomen nudum 
Picea engelmanni Parry ex Engelmann; Trans. Acad. St. Louis 2:212. 1863 
Picea columbiana Lemmon, Gard. and For. 10:183. 1897 


There is little doubt that Picea glauca var. albertiana (S. Brown) Sar- 
gent (Picea albertiana S. Brown, Torreya 7:126. 1907) is based on an 
individual of the hybrid swarm between subsp. glauca and subsp. engel- 
mannii and in consequence this name should no longer be perpetuated. 
When reference is made to these hybrid intermediates it should be done 


1959] CLARKSON: IRIS 115 


by employing a formula as provided by Article H.2 of the International 
Code with particular attention to the “Note.” 

Goodman (1950), in describing his P. engelmannii var. glabra, almost 
certainly had at hand a biotype of Engelmann’s spruce that showed the 
result of the introgression of genes from the glabrous White spruce. 


Department of Biology and Botany, 
University of British Columbia, 
Vancouver, Canada 
LITERATURE CITED 


Barton, G.M. and J. A. F. Garpner. 1957. Comparison of the heartwood extractives 
of Picea glauca and Picea engelmanni. Forestry Chron. 33 (2) :136-138. 

EKLuUNDH, G. 1943. Artkorsningar inom sl. Picea . . . tillhorande fam. Pinaceae. 
Svensk Papp Tidn. 46:55-61, 101-105, 130-133. (Species crosses within the gen- 
era Picea, etc.) (Sw.) In Forestry Abstr., V. abs. 95. 

GaRMAN, E.H. 1957. The occurrence of spruce in the interior of British Columbia. 
Brit. Col. Forestry Service Tech. Publ. T. 49:1-31. 

GoopMAN, G. J. 1950. A New Variety of Engelmann Spruce. Madrono 10:177. 

Jounson, L.P.V. 1939. A descriptive list of natural and artificial hybrids in North 
American forest-tree genera. Canad. Jour. Res. C. 17:411-444. 

Lanjouw, J., editor. International Code of Botanical Nomenclature adopted by the 
Eighth International Botanical Congress, Paris, July, 1954. Utrecht. 

WricutT, J.W. 1955. Species crossability in spruce in relation to distribution and 
taxonomy. Forest Sci. 1:319-349. 


FIELD STUDIES OF NATURAL HYBRIDIZATION IN THE 
OREGON SPECIES OF IRIS L. SUBSECTION 
CALIFORNICAE DIELS 


QUENTIN D. CLARKSON 


Smith and Clarkson (1956) have discussed the cytological aspects of 
hybridization in /ris, subsection Californicae. They reported that, with the 
exception of /. tenuis Wats., which has been removed to a new subsection 
(Clarkson, 1958), all the members of the subsection studied had a uni- 
form chromosome morphology, and all contained a diploid number of 
forty. Fertile hybrids were produced experimentally without difficulty and 
all were highly fertile except that hybrids involving /. tenuis as a parent 
could not be produced. This paper will discuss some natural hybrids of 
those taxa occurring in western Oregon and will propose certain nomen- 
clatural revisions. Because the subsection was treated taxonomically by 
Foster (1937), conventional citations will not be included except for taxa 
described since that time. 


DISTINGUISHING CHARACTERISTICS. Aside from the usual characteris- 
tics of the genus /ris, the members of the subsection Californicae are dis- 
tinguished by a usually deltoid stigma; D-shaped, cubical or ovoid seeds; 
the absence of foliaceous stem leaves; and tough basal leaves which are 
reddish at base. A number of characteristics have been used for distin- 


116 MADRONO (Vol. 15 


guishing species by various authors, including Foster (1937) and Dykes 
(1912) in the two most complete reviews of the subsection. These have 
been evaluated with herbarium specimens (Clarkson, 1950) and with liv- 
ing material. Those characters which allow ready distinction of taxa and 
which are most easily utilized for study of hybrids are as follows: 

Perianth. The petals are usually narrow and about the same length as 
the sepals which are linear to spathulate in shape. Color of the perianth 
ranges from white, purple, pale yellow, golden yellow, to apricot and 
maroon. 

Perianth tube and pedicel. The perianth tube varies in length from 
0.4 to 12 centimeters with the pedicel varying approximately inversely. 

Bracts of the inflorescence. The bracts, or spathes, are variable in shape 
from ovoid to linear; in length from 2 to 15 centimeters; and in position 
from opposite to alternate. 

FIELD StupiEs. Although a quantitative study of morphological and 
ecological characteristics was desirable, inadequate knowledge of distri- 
bution of the species and the absence of definite knowledge of natural 
hybrids made such work impossible. Field work was therefore conducted 
on a qualitative basis. A limited number of transplants made during cur- 
rent field work and previously by Smith (Smith and Clarkson, 1956) dem- 
onstrate that the characteristics of the taxa are constant under varying 
environmental conditions. Herbarium specimens designed to represent all 
the variation present in a given location were collected. These mass col- 
lections are in the possession of the author. 

Results of field studies are given below together with interpretations 
of their significance. 


HYBRIDS BETWEEN I. TENAX AND I]. CHRYSOPHYLLA 

Iris tenax is characterized by narrow, distant bracts; a perianth tube 
from 4 to 9 millimeters in length and by lavender to purple flowers with 
broad spathulate sepals. The plants are tall and with definite stems. 
Though the species is typically purple-flowered, two yellow-flowered pop- 
ulations have been found. One of these is along Scoggin’s Creek in Wash- 
ington County, Oregon. It has been treated as J. gormanii Piper and more 
recently as /. tenax var. gormanii (Piper) Foster. The second yellow- 
flowered population is on Monument Peak, Linn County, Oregon. This 
local population is apparently not well known and has not been cited in 
the literature of the subsection. Neither of these populations can be dis- 
tinguished from the typical phase of J. tenax by any trait other than color. 

The general range of the species is from the central part of western 
Washington to southern Oregon. Ecologically it is a species of unshaded 
conditions and is abundant on the oak-covered hills of the Willamette and 
Umpqua valleys. It does not extend into coniferous areas unless trees are 
cut, roads built, or conditions otherwise disturbed in such a way that shad- 
ing is reduced. Within the general range there seem to be no soil or mois- 
ture factors limiting its distribution. Factors controlling the northern and 
southern limits of the species range are probably climatic in nature. 


1959 | CLARKSON: IRIS 117 


Iris chrysophylla is characterized by lanceolate, opposite bracts; a 
perianth tube 5 to 9 (occasionally to 12) centimeters in length; a pedicel 
less than 1 centimeter in length; and by pale yellow flowers with narrow 
perianth parts. In southwestern Oregon, where the species is most abun- 
dant, well-marked stems are produced. In the Cascade Mountains of 
northern Oregon, the plants are often nearly stemless. Plants at two pre- 
viously unreported stations of the species, Mill Creek, Polk County, Ore- 
gon, and Prairie Mountain, Benton County, Oregon, are of the Cascade 
type. 

Ecologically 7. chrysophylla is a species of open coniferous forests. It 
grows best on drier soils and will tolerate more shade than /. tenax. It isa 
characteristic species of the ponderosa pine and ponderosa pine-Douglas 
fir communities of southwestern Oregon. 

Distinctive hybrids between J. tenax and I. chrysophylla have been 
found in three locations: (1) 4% miles up Mill Creek from the Dallas— 
Wallace Bridge highway, Polk County, Oregon; (2) 1314 miles southwest 
of Roseburg, Douglas County, Oregon, along Oregon State highway num- 
ber 42; (3) steep hills along the first tributary of the North Santiam River 
west of the Detroit Dam, Marion County, Oregon. In all three locations 
conditions have been disturbed by road construction. 

At all three of these sites individuals have been found which are inter- 
mediate between the parent species and which cannot be assigned to either 
species. Bracts of the hybrids are opposite and are broader than is char- 
acteristic of J. tenax but not so broad as those of J. chrysophylla; flower 
color is maroon to grayish-lavender; and the perianth parts are slender. 
The plants are smaller than typical 7. tenax but have a definite stem. 
Perianth tube length in the natural hybrids averages 2.2 centimeters and 
the pedicel averages 1.4 centimeters. Comparative measurements in the 
known hybrids are perianth tube 2.5 and pedicel 1.8. This indicates that 
the hybrids are possibly of the F, generation. In other characters the 
natural hybrids compare closely with the known hybrids. 

The area of overlapping ranges is, in all three cases, small. At Mill 
Creek and along the North Santiam highway, a transect of less than 100 
yards extends through the area occupied by both species. At the site south- 
west of Roseburg, a similar transect extends less than one-half mile. 

Hybrids between these two species appear to be introgressive toward 
I. tenax. Individuals identifiable as 7. tenax show definite /. chrysophylla 
characteristics for at least ten miles from the area where typical J. chryso- 
phylla occurs. These individuals exhibit a change in color toward a pale 
purple; a reduction in size; and a tendency toward broader and less dis- 
tant bracts. These modifications in /. tenax are what might be expected 
as a result of hybridization with J. chrysophylla. The intermediates pre- 
viously discussed are found in the areas where the two species occur to- 
gether but /. chrysophylla remains constant and no individuals of that 
species have been found which exhibit /. tenax characteristics. 

Introgressive hybridization between these two species is a close parallel 


118 MADRONO [Vol. 15 


of a situation described by Anderson (1949) in which hybridization is 
followed by backcrossing and selection of backcross types. Apparently, 
in this case, only the offspring of the intermediate x J. tenax backcross are 
selected. The factors of the ecology which bring about this selection are 
not known. The introgression, however, has resulted in a considerable in- 
crease in the variability of J. tenax while J. chrysophylla has remained 
constant. 

The occurrence of J. chrysophylla at two locations in the Coast Range 
of Oregon was not unexpected. The presence of individuals of J. tenax 
with lanceolate, opposite bracts in the Coast Range west of Corvallis, 
Oregon, suggested possible hybridization with J. chrysophylla. At the 
Mill Creek location in Polk County, Oregon, the two species are in contact 
at the present time, as was previously mentioned. At Prairie Mountain, 
Benton County, Oregon, there is no present day contact on the north and 
probably not on the south. Iris chrysophylla is abundant on a broad, well- 
drained meadow, along a south-facing slope near the summit of Prairie 
Mountain at an elevation of about 3200 feet. Jvis tenax is not now in con- 
tact on the north, presumably because of the dense growth of Douglas 
fir on that side. The south side of the mountain is more open and the two 
species may come together though no intermediates have been found. J7is 
tenax specimens collected at Horton, Lane County, eight miles to the 
south, have a narrower and more pale perianth which suggests hybridiza- 
tion with J. chrysophylla. 

Thirty miles north of Prairie Mountain are individuals of J. tenax with 
linear-lanceolate, opposite bracts, and narrow, pale purple perianth parts. 
These plants are probably the result of introgression from I. chrysophylla 
to J. tenax. These hybrid forms indicate either that J. chrysophylia is more 
abundant in the Coast Range than is now known or that it was more 
abundant in the past and has survived only on the more favorable sites. 
Either hypothesis may be true in part, but it seems unlikely that these 
forms are the result of recent hybridization. /ris chrysophylla has not been 
found in the Coast Range between Prairie Mountain and Mill Creek, and 
distances are probably too great for plants at those locations to be in- 
volved as parents. The most probable explanation is that J. chrysophylla 
was more abundant in the Coast Range of northern Oregon in the past 
and has remained only at the dry, open sites such as Mill Creek and Prai- 
rie Mountain. The occurrence of a warm, dry period in postglacial times, 
followed by general cooling and increase in moisture (Hansen, 1947) 
makes this a reasonable assumption. Whether J. chrysophylla was distrib- 
uted throughout the general area of northwestern Oregon in the past or 
only on the higher peaks, is a question which cannot be answered. In 
either case, abundant opportunities for hybridization could have existed. 
Introgression, such as occurs today, could have given greater adaptive 
value to J. tenax while I. chrysophylla remained more nearly constant and 
more vulnerable to environment change. 

There is, however, no reason to suppose that the yellow-flowered forms 


1959] CLARKSON: IRIS 119 


of J. tenax on Monument Peak and in Washington County, Oregon, are 
of hybrid origin. All the hybrids between these two species, including the 
artificial hybrids produced for this study, exhibit a tendency toward the 
lanceolate, opposite bracts of J. chrysophylla. The purple flower color is 
reduced to greyish-lavender in the known Fy, hybrids, and none of the 
hybrids found in nature have yellow flowers. An independent origin for 
this color trait is not improbable as evidenced by J. hartwegu, which is 
closely related to 7. tenax and has yellow flowers and by the fact that there 
are yellow-flowered forms in J. macrosiphon which is otherwise purple- 
flowered. 


HYBRIDS BETWEEN I. DOUGLASIANA AND I. INNOMINATA 


Iris douglasiana is a tall species averaging about 60 centimeters in 
height and is characterized by leaves about a centimeter wide; a branch- 
ing stem; lanceolate, opposite bracts; large pale purple to white flowers; 
and a perianth tube 1 to 2 centimeters in length. Two or three flowers per 
branch are produced. Distribution of the species in Oregon is limited 
chiefly to a narrow, open coastal strip from Coquille, Coos County, south- 
ward. The species normally extends inland only along river valleys. It has 
not been seen in dense shade. 

Iris innominata is a low species averaging about 35 centimeters in height 
and is characterized by narrow leaves which are about 4 centimeters wide; 
simple stems; ovate, opposite bracts; small, golden yellow flowers; and a 
perianth tube 1 to 2.5 centimeters in length. Distribution of the species is 
limited to the open meadows and hills of southern Douglas, Coos, and 
Curry counties, Oregon. 

Hybrid colonies of about 100 plants each of the J. douglasiana X I. 
innominata cross have been found along Saunder’s Creek, Curry County, 
Oregon. The two sites are 114 and 2%4 miles from the Rogue River, 3 miles 
upriver from Gold Beach on the south side of the Rogue River. The 
colonies are found in cut-over forest land with most of the plants com- 
pletely exposed to the sun, though a few are found in the shade of Umbel- 
lularia californica. Some of the individuals are similar to the known hy- 
brids grown for cytological study. Perianth size and plant size appear to 
be intermediate between the parent species and the bracts are shorter and 
more ovoid than are those of J. douglasiana but are longer and narrower 
than those of J. innominata. These individuals may be F; hybrids. Other 
plants appear to be the result of backcrossing and segregation. 

Aside from the demonstration of genetic continuity between species, 
the taxonomic significance of the hybrids is considerable. Some individ- 
uals from one of the hybrid colonies compare closely with individuals re- 
ferred to J. thompsonu Foster. There is a strong suggestion of the hybrid 
origin of that taxon. Jris thompsonii has been collected along the Rogue 
River and along United States Highway 101 from Carpenterville to 
Brookings, Curry County, Oregon. In California, it has been collected 
along the Smith River, northeast of Crescent City, Del Norte County. The 


120 MADRONO [Vol. 15 


species can be distinguished from /. innominata by the more lanceolate 
bracts and by the pale purple to lavender flower color. It also seems to be 
taller, averaging about 45 centimeters. These are characteristics which 
could be fixed by backcrossing of the 7. douglasiana X I. innominata hy- 
brids to J. innominata. 

A similar hypothesis can be erected to explain those specimens with 
more ovate bracts and lavender-grey flowers referred by Foster (1937) 
to a new variety, J. douglasiana var. oregonensis. Backcrossing of the in- 
termediates of the J. douglasiana < I. innominata cross to I. douglasiana 
could result in the fixing of these 7. innominata traits in otherwise typical 
I. douglasiana. 

While both assumptions are largely hypothetical, they are lent support 
by the occurrence of these variants in an area where the two species come 
together and hybridize. The only barriers seem to be ecological, though 
exact factors cannot be stated. The removal of the forest trees with the 
accompanying reduction in shade has apparently removed the barrier to 
hybridization in the case discussed here, but elevation and soil factors may 
be important in other locations. However, hybrids probably will be found 
wherever the hills of Coos and Curry counties are near the ocean and 
conditions are disturbed or where /. douglasiana extends inland along river 
valleys. In these areas, at least, the two species can be expected to occur 
together. 


HYBRIDS BETWEEN I. BRACTEATA AND I. THOMPSONII 


Iris bracteata is a tall species averaging about 50 centimeters in height 
and with a thick perianth tube 0.5 to 1.0 centimeter in length, a pedicel 
3 to 6 centimeters long, and golden yellow flowers. Typically the leaves 
are a glossy green on the upper surface and glaucous on the lower. The 
species has been seen only in southwestern Josephine County, Oregon, and 
northeastern Del Norte County, California. Ecologically it is restricted 
to shaded places. The species is found in greatest abundance within pon- 
derosa pine communities, but it is sometimes found on cut-over forest land 
under a cover of bracken fern. 

Iris thom psoni has been discussed as a possible hybrid segregate of the 
I. douglasiana X I. innominata cross. Colonies are best developed along 
the lower Smith River, from 12 to 15 miles northeast of Crescent City, 
California. In this location, the colonies are well established on open rocky 
hillsides, and have not been seen in the shade. 

Proceeding northeastward from Jedediah Smith State Park in Del 
Norte County, over the Siskiyou Mountains into Oregon, the colonies be- 
come more shade tolerant. Color changes gradually from predominantly 
pale purple flowers to predominantly yellow flowers, though a few white- 
flowered plants are present. Bract characteristics appear to be interme- 
diate between the two species, changing gradually from ovoid and 5 centi- 
meters long, to narrower and 7 centimeters long. Height of the plant also 
becomes progressively greater as colonies nearer typical J. bracteata are 


1959] CLARKSON: IRIS al 


examined. Three miles north of the summit of the Siskiyou Mountains 
only typical J. bracteata has been seen. 

No individuals similar to the known F, hybrids have been found in 
nature. The transition between the two species is gradual, probably be- 
cause of ecological requirements which permit more hybridization and 
survival of most of the hybrid offspring. 


HYBRIDS BETWEEN I. BRACTEATA AND |]. CHRYSOPHYLLA 


Distinct hybrids between these two species have not yet been clearly 
demonstrated. Despite the pronounced morphological differences between 
the two, only one individual has been secured which can be considered 
intermediate. The relationship of the perianth tube to the pedicel is re- 
versed in these two species. /vis chrysophylla has a perianth tube 5 to 6 
times longer than the pedicel. In /. bracteata the pedicel is 5 to 6 times 
longer than the perianth tube. In the intermediate specimen cited above, 
the pedicel is twice the length of the perianth tube, clearly not character- 
istic of either J. chrysophylia or I. bracteata. This specimen was collected 
near Bridgeview, Josephine County, Oregon, where the two species occupy 
the same general area. 

Complementing this admittedly limited evidence, field examination re- 
veals hybrid characteristics which are not readily demonstrable with 
pressed specimens. Near Cave Junction, Josephine County, there are indi- 
viduals of J. bracteata with narrow perianth segments characteristic of 
I. chrysophylla. The hybrid origin of these individuals is supported by the 
fact that all the known hybrids involving /. chrysophylla as a parent ex- 
hibit the narrow perianth of that species. There are also individuals, in 
the Cave Junction area, of J. chrysophylla with perianth color similar to 
the golden yellow of J. bracteata. The known J. bracteata < I. chryso- 
phylla hybrid has golden-yellow flowers and all other crosses involving 
golden-yellow and pale yellow-flowered parents show the flower color of 
the golden-yellow parent. 


Taxonomy. Due to the absence of cytological barriers between taxa 
and the presence of natural hybrids, nomenclatural revision which will 
better reflect the biology of the organisms seems necessary. Therefore, the 
following new combinations are proposed. Further field work is needed 
before accurate limits can be set for the California members of the sub- 
section and they will not be considered here. 


1. In1s TENAX Dougl. subsp. TENAX (J. tenax Douglas ex Lindley, Bot. 
Reg. xv. t. 1218, 1829). This subspecies includes the typical form which 
has been discussed previously in this paper as 7. tenax. Also included, 
without nomenclatural distinction, is the yellow-flowered J. tenax var. 
gormanu. The yellow-flowered form from Monument Peak, Linn County, 
Oregon, will be included within J. tenax gormanii and therefore within 
I. tenax tenax without nomenclatural distinction. 


2. IRIS TENAX subsp. CHRYSOPHYLLA (Howell) (J. chrysophylla How- 
ell, Fl. N. W. America 1:633, 1902). No type specimen was designated 


122 MADRONO [Vol. 15 


for Howell’s J. chrysophylla. However, filed with the type specimen col- 
lection at the herbarium of the University of Oregon is a specimen labeled 
“Type specimen” in the distinctive handwriting of Thomas Howell. This 
specimen, collected at Grants Pass, Josephine County, Oregon, in May 
1887, must be considered the type specimen for Howell’s species and there- 
fore for the proposed subspecies. 


3. IRIS TENAX Subsp. BRACTEATA (Watson) (J. bracteata Watson, Proc. 
Amer. Acad. 20:375, 1885). This subspecies includes the species as de- 
scribed by Watson. 


4. IRIS TENAX subsp. DOUGLASIANA (Herbert) (J. douglasiana Herbert, 
Bot. Beechey Voy. 395, 1841). Included in J. tenax douglasiana without 
nomenclatural distinction is J. douglasiana var. oregonensis Foster. So 
long as it is understood that Foster’s variety is probably of hybrid origin, 
there is no need to distinguish a form which obviously belongs to what 
has been described as J. douglasiana. Since this subspecies is found in Cali- 
fornia as well as Oregon, this new combination must necessarily include 
the California plants. This does not mean that the California plants of 
this taxon should be ignored in any further study. There is undoubtedly 
considerable variation present that is not included in the Oregon repre- 
sentatives. 

5. IRIS TENAX subsp. INNOMINATA (Henderson) (J. innominata Hen- 
derson, Rhodora 32:23, 1930). This subspecies has been retained as origi- 
nally described except for the reduction in rank. 


6. IRIS TENAX subsp. THOMPSONII (Foster) (J. thompsoniu Foster, Rho- 
dora 38:199, 1936). The only real difference between this proposed new 
subspecies and /. tenax innominata is in the purple flower color of J. tenax 
thompsonii. Since it has been suggested that J. tenax thompsonu may be 
of hybrid origin with /. tenax innominata as one parent, a close morpho- 
logical relationship is to be expected. The two taxa differ slightly in dis- 
tribution, /. tenax thompsonii is a species of elevations between 400 and 
and 1000 feet while /. tenax innominata is usually at sites over 1500 feet. 
For that reason and because /. tenax thompsonu does form distinct col- 


onies, it is retained. Division of Science: 


Portland State College, 
Portland, Oregon 
LITERATURE CITED 
ANDERSON, E. 1949. Introgressive hybridization. Wiley and Sons. New York. 
CrarkKson, Q. D. 1950. Population studies in Iris. M.S. thesis. University of Oregon. 
Eugene. 
. 1958. Iris section Apogon subsection Oregonae. Subsect. Nov. Madrono 
14:246-247. 
Dykes, W.R. 1913. The genus Iris. Cambridge University Press. London. 
Foster, R.C. 1937. A cyto-taxonomic survey of the North American species of Iris. 
Contr. Gray Herb. 119:1-82. 
Hansen, H.P. 1947. Postglacial forest succession, climate and chronology in the 
Pacific northwest. Trans. Am. Philos. Soc., N.S. 37, part 1:1-130. 
SmitH, F.H. and Q. D. Crarxson. 1956. Cytological studies of interspecific hybrid- 
ization in Iris, subsection Californicae. Am. Jour. Bot. 43:582-588. 


1959] PELTON: MERTENSIA 123 


VARIATION PATTERNS IN FOUR CLONES OF 
MERTENSIA CILIATA! 


JEANETTE S. PELTON 


Mertensia ciliata (James) G. Don is well delineated from other species 
of the genus in the monograph by Williams (1937). Field observations 
in the area of the present study, Gunnison County, Colorado, bear out 
this distinctness of M. ciliata from other sympatric species, although con- 
siderable intraspecific variation is easily detected. In spite of the fact that 
certain cther species of the group grow sympatrically in the study area 
with M. ciliata, five years of observation by the author have not uncovered 
a single likely instance of hybridization between M. ciliata and any of the 
other species. Nor did artificial cross pollination produce any fertile seed 
between MV. ciliata and M. fusiformis Greene. In the observed popula- 
tions, therefore, /. ciliata seems an excellent species in which to study 
quantitatively intraspecific variation patterns which are probably uncom- 
plicated by any present inflow of genes from another species. With this 
objective in mind, three floral characteristics within and among four 
clones of M. ciliata were chosen for study. The clones were selected such 
that each was separated from the other by a distance of one-third to two 
miles. Such a distance probably assures that each clump is an individual 
clone with a different genetic origin. Hence the four clumps will be re- 
ferred to hereafter as clones A, B, C, and D. Since each clone was grow- 
ing in a different combination of environmental factors, at least four of 
the many micro-habitats to which individuals of M. ciliata are adapted 
are represented. By selecting clones in the above manner, it was presumed 
that a measure of somatic variability could be obtained, since the pheno- 
typic measurements would be made upon single genotypes each produced 
in a slightly different environment. In addition, differences in gene ex- 
pression among the clones imply possible genetic variation patterns. 


METHODS AND RESULTS 


Clones A, B, C, and D were collected in the summer of 1953 in or near 
the Rocky Mountain Biological Laboratory, Gunnison County, Colorado. 
No clone was nearer than approximately one-third mile from another, 
and all clones were located on different drainage channels. The altitude 
is approximately 9,500 feet for three of the clones and 10,000 feet for 
Clone B. Voucher specimens of the collections are in the personal herba- 
rium of the author. 

Length of calyx, corolla-tube, and corolla-limb were measured from 
these herbarium voucher specimens. In this study the corolla-limb, as 
defined by Williams (1937), will include that portion of the corolla above 
the fornices. Individual mature flowers were measured for the three floral 
characteristics to 0.5 mm. using low power of a binocular microscope to 


1 The author is grateful for the use of facilities of the Rocky Mountain Biological 
Laboratory and to Dr. John F. Pelton for criticisms and additions to the manuscript. 


124 MADRONO [Vol. 15 


TABLE 1. Arithmetic mean, standard deviation, and range of variation for length 
of calyx, corolla-tube, and corolla-limb in four clones of Mertensia ciliata 


Calyx Length Corolla Tube Length Corolla Limb Length 
Mean Stand- Range Mean Stand- Range Mean Stand- Range 
Clone No. of ar of Vari- ard of Vari- ard of Vari- 
Indi- Devi- ation Devi- ation Devi- ation 
viduals ation ation ation 


150 25 024 20-35 54 0.52 3.5-65 5.9 0.70 4.5-8.0 
94 18 O30 15-25 62 151 5075 73 1.04 5.0-9.5 
132 18 O33 15-25 68 0.50 55-80 61 088 4.0-8.0 
40 21 O30 15-25 5.9 036 50-65 7.1 0.79 5.0-9.0 


wo @ Woo Ee 


increase accuracy. Arithmetic mean, standard deviation, and range of va- 
riation were determined for each characteristic measured in the four 
clones. These results are presented in Table 1. In Figure 1 variation 
among the four clones is diagrammed using mean length of calyx, corolla- 
tube and corolla-limb. Range of variation within the clones is diagrammed 
in Figure 2. 

Descriptions of the clones are as follows: 


CionE A. Collected July 15 from a moist roadside site one-fourth mile 
north of the laboratory. The clump was growing on the edge of a dense 
willow thicket in full sun. There was a total of 14 individual stems in the 
clone, averaging 10.7 mature flowers per stem. Total number of mature 
flowers measured was 150. 

CLONE b. Growing in a very wet location in the partial shade of a spruce- 
fir forest on the edge of a beaver pond about two miles northeast of the 
laboratory at 10,000 feet and collected on July 19. The average number 
of mature flowers was 3.1 on a total of 30 individual stems; 94 mature 
flowers were measured. 


CLONE C. Chosen from a population in an aspen forest on laboratory 
property. The clone was collected on July 23 in a partially shaded rocky 
stream bed. This clone had 32 stems, the largest number of individual 
stems of the four clones. Average number of mature flowers per stem was 
4.1; 132 mature flowers were measured from this clone. 


CLONE D. Growing in a willow thicket near Copper Creek adjacent to 
the laboratory. The soil was wet and rocky, the clone growing in partial 
shade. Collection was on July 7. The three individual stems of this clone, 
all flowering, averaged 13.3 mature flowers per stem and totaled 40 ma- 
ture flowers suitable for measurement. 


DISCUSSION 


The variation pattern for each of the four studied clones of Mertensia 
ciliata is striking enough that a given clump can be identified on the basis 
of a distinctive combination of average length of calyx, corolla-tube and 
corolla-limb (fig. 1). On the other hand, individual measurements within 


1959] PELTON: MERTENSIA 125 


| 2 3. 4 5 6 7 8 9 10 I l2 13 14 
LENGTH IN MILLIMETERS 


Fic. 1. Idiograms showing average lengths in millimeters of calyx, corolla tube, 
and corolla limb in clones A, B, C, and D of Mertensia ciliata. 


each clone vary considerably for these three characteristics (fig. 2). How- 
ever, the range of variation of a character in a clone usually does not over- 
lap completely with that of the same character in other clones. These 
observed variations within and among the clones could be the result of 
three factors mentioned by Stebbins (1950): environmental modifica- 
tion, gene recombination, and mutation of genes or chromosomes. Con- 
siderations of the role of these factors as possible explanations for the 
variability observed is discussed in the following. 


ENVIRONMENTAL MopiFicaTions. Variability within a given clump 
should be a measure of environmental influences except for rare bud 
mutations or the unlikely possibility that one clump was derived from 
two or more seedlings. Floral characteristics were chosen for study be- 
cause they are known to be frequently less easily influenced by environ- 
mental factors than are many vegetative characteristics (Clausen, Keck, 
and Hiesey, 1940; Anderson, 1929; Brainerd and Peitersen, 1920). All 
three characteristics vary considerably in range of measurements within 
each clone, such as the 1.5 mm. variation in Clone A calyx length which 
averages only 2.5 mm., the 3 mm. variation in corolla tube length in Clone 


126 MADRONO [Vol. 15 


(@) 


FLOWERS 


OF 


NUMBER 
@ 
° 


ie) rae 35 ro) 45 S59 6.5 fis) 85 45 hohe] 65 (as) 65 95 
CALYX LENGTH COROLLA TUBE LENGTH COROLLA LIMB LENGTH 
Fic. 2. Histograms showing distribution of individual measurements in millimeters 


of length of calyx, corolla tube, and corolla limb in clones A, B, C, and D of Mertensia 
ciliata. 


A which averages 5.4 mm. in length, or the 4.5 mm. variation in corolla 
limb length for Clone B while average length is only 7.3 mm. This indi- 
cates that gene action, even in the fairly uniform environment of a single 
clone, differs in the final expression of length of calyx and corolla. To 
determine the various environmental factors that control the diverse 
action of these genes is difficult even in the imagination. External condi- 
tions such as soil, temperature, light, humidity, and biotic interactions 
would usually be expected to vary comparatively slightly during the de- 
velopment of the flower primordia of a single clump. Internal conditions 
such as amount and distance of the vascular supply, internal temperature, 
chemical environment, and the differing interaction of other genes in 
ontogeny would perhaps be more important than external environment 
since a slight variation of internal environment during the delicate inter- 
actions between gene initiation and the end result of expression could 
alter the phenotype. Whichever of the external or internal conditions 
may be important, their effect on the genes controlling calyx and corolla 
length accounts for a large proportion of the variation observed in this 
study, probably all of the intraclonal variability. This would support the 
idea that corolla and calyx length in this case are quantitative character- 
istics, dependent on multiple genes, since quantitative characteristics are 
usually subject to considerable modification by environment (Srb and 
Owen, 1952). 


GENE RECOMBINATION AND MUTATION. While the somatic variation 
discussed above cannot result in permanent changes in the species, gene 


1959] PELTON: MERTENSIA Ld 


recombination and/or gene and chromosomal mutation are thought to 
contribute to variation that can foster evolutional change in the species 
(Stebbins, 1950). Whether gene recombination and mutations could ac- 
count for the differences among these four clones cannot be determined 
from the results of this study. Probably some of the observed differences 
among clones would be attributed to dissimilarities in external or internal 
environments of the four clones. It must be noted again, however, that 
floral characteristics are probably less subject to environmental modifica- 
tion than are other features of external morphology. Floral differences 
based on the pattern of average length of calyx, corolla-tube, and corolla- 
limb illustrate a distinctive combination in each clone (fig. 1). On close 
examination of the individuals compounded in these means it is found 
that only a few flowers approach the extremes of the large range of varia- 
tion, and that standard deviations, given in Table I, are not large. Also, 
the histograms for each clone do not closely coincide with those for the 
other clones, although considerable overlapping does occur (fig. 2). These 
patterns of difference among the clones are probably the result of gene 
recombination, mutation because of its rarer occurrence being a less like- 
ly source. If Mertensia ciliata has a large number of genes active in regu- 
lating corolla length, such as the estimated twelve or more controlling 
corolla size in Nicotiana (Smith, 1937), it would be plausible to assume 
such recombination of the many genes in different individual plants or 
clones. Close linkage between the polygenes that determine quantitative 
characteristics, however, is often assumed to restrict the range of recom- 
bination of characteristics (Smith, 1944). Nevertheless, while somatic 
variation is doubtless the main factor in accounting for the variation 
within each clump, the characteristic variation patterns presented here 
for calyx and corolla length would imply some genetic differences among 
the clones, probably, as a result of gene recombination. 


SUMMARY 


Length of calyx, corolla-tube, and corolla-limb were measured for four 
widely separated clumps, presumably clones, of Mertensia ciliata that 
were collected from four differing and widely separated sites in Gunnison 
County, Colorado. Comparisons of calyx and corolla lengths were made 
within and among the four clones. The considerable variation of the three 
characteristics found within each of the clones is attributed to external 
and internal environmental factors, internal conditions probably being 
more important. Variation patterns among the clones differ enough to give 
each clone a distinctive combination of average lengths for the three 
characteristics. In most cases, the range of variation in calyx and corolla 
length within each clone does not completely coincide with that of the 
other clones. These differences in variation patterns imply some genetic 
differences among the clones, probably as a result of gene recombination. 


Department of Botany, 
Butler University, Indianapolis, Indiana 


128 MADRONO [Vol. 15 


LITERATURE CITED 


ANDERSON, E. 1929. Variation in Aster anomalus. Ann. Missouri Bot. Gard. 16: 
129-144. 

BRAINERD, E., and E. K. PEITERSEN. 1920. Blackberries of New England—their clas- 
sification. Vermont Agr. Exp. Sta. Bull. No. 217. 84 pp. 

CiausEN, J., D. D. Keck and W. M. Hirsey. 1940. Experimental studies on the 
nature of species. I. The effect of varied environments on western North Ameri- 
can plants. Carnegie Inst. Publ. No. 520. 452 pp. 

SmitH, H. H. 1937. Inheritance of corolla color in the cross Nicotiana Langsdorffii 
by N. Sanderae. The relation between genes affecting size and color in certain 
species of Nicotiana. Genetics 22:347-375. 

. 1944. Recent studies on inheritance of quantitative characters in plants. 
Bot. Rev. 10:349-382. 

Srp, A. M., and R. D. Owen. 1952. General Genetics. W. H. Freeman and Co. 
San Francisco, Calif. 561 pp. 

Stepsins, G. L., Jk. 1950. Variation and evolution in plants. Columbia Univ. Press. 
643 pp. 

WitiiaMms, L.O. 1937. A monograph of the genus Mertensia in North America. 
Ann. Missouri Bot. Gard. 24:17-159. 


NEW COMBINATIONS IN ASTER 
ROXANA S. FERRIS 


Through an inadvertence the following new combinations were not 
legally made in the recent “Flora of the Marshes of California” by Her- 
bert L. Mason. 


ASTER OCCIDENTALIS var. parishii (Gray) Ferris, comb. nov. A. fre- 
montu var. parishi Gray, Syn. Fl. N. Amer. 1 (2): 192. 1884. 


ASTER OCCIDENTALIS var. delectabilis (H. M. Hall) Ferris, comb. nov. 
A. delectabilis H. M. Hall, Univ. Calif. Publ. Bot. 3:82. 1907. 


Both of these varieties occur in California in the Sierra Nevada and in 
the mountains of southern California, and they reoccur in the San Pedro 


Martir of northern Baja California, Mexico. 
Dudley Herbarium, Stanford University, 
Stanford, California. 


NOTES AND NEWS 


Some publications of interest follow: 

Under the auspices of the Gobierno del Estado de México, Direccion de Recursos 
Naturales (Toluca) publications on the Flora del Estado de México have continued 
to appear. During 1958 Professor Maximino Martinez completed the Flora Medicinal 
as well as the treatment of the Cactaceae and some forty smaller families; Professor 
EKizi Matuda treated the Gramineae, Umbelliferae and the Compositae. 

Drawings of British Plants, by Stella Ross-Craig. Part XI. Droseraceae—Ficoi- 
daceae. 39 pls. 1958, 9s. 6d. G. Bell and Sons, Ltd. London. Part XII. Umbelliferae (1). 
36 pls. 1958. 9s. 6d. 1958. 


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. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume I, 1916-1929 . . . . = . $5.00 
Volume IJ, 1930-1934. . . .) . 5.00 
Volume ITT, 1935-1936 . . . . 5.00 
Volume IV, 1937-1938 . . . . 5.00 
Volume V, 1939-1940. . . . ., 5.00 
Volume VI, 1941-1942 . . .. . 5.00 
Volume VIT, 1943-1944 . . .. 5.00 
Volume VIII, 1945-1946 . . .. . 5.00 
Volume IX, 1947-1948 . eo ee ii 6.00 
Volume X, 1949-1950. . . .. . 7.00 
Volume XJ, 1951-1952. ww, 8.00 


Volume XIT, 1953-1954 . . .. . 8.00 
Volume XIII, 1955-1956 . . . 8.00 


Volume XIV, 1957-1958 . . . . 8.00 
Single numbers, 

Volumes I and 2) .)).3) 2. ee .50 
Succeeding numbers, 

(except Volume XI, No.2). .. 1.00 

Volume XI; No. 20530 a toe 1.50 


The subscription price of MADRONo is $4.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


g07/V'1 


ew 
VOLUME 15, NUMBER 5 JANUARY, 1960 


Contents 


PAGE 
THE DISTRIBUTION OF DwarF MISTLETOES, ARCEU- 
THOBIUM, IN CALIFORNIA, Job Kuzjt 129 


NUCLEAR CYTOLOGY OF THE CALIFORNIA MousE-TAILs 
(Mvyosurus), Donald E. Stone 139 


VARIATION IN SECTION TRIGONOPHYLLAE OF NICO- 
TIANA, Philip V. Wells 148 


STUDIES ON SECOTIACEOUS FuNGI VII. SECOTIUM AND 
NEosEcoTIUM, Rolf Singer and Alexander H. Smith 152 


Review: Adriance S. Foster and Ernest M. Gifford, Jr., 
Comparative Morphology of Vascular Plants 
(Sanford Tepfer) 158 


NOTES AND NEWS 160 


A WEST AMERICAN JOURNAL OF BOTANY > > Pe 


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 Madrono, 
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. Davipson, University of Nebraska, Lincoln. 

Ivan M. JounstTon, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MitprepD E. Maruras, University of California, Los Angeles 24. 
Marion OwNBEY, State College of Washington, Pullman. 

IrA L. Wiccrns, Stanford University, Stanford, California. 


Secretary, Editorial Board—ANNETTA CARTER 


Department of Botany, University of California, Berkeley. 


Business Manager and Treasurer—JOHN THOMAS, 


Dudley Herbarium, Stanford University, Stanford, California 


CALIFORNIA BOTANICAL SOCIETY, INC. 


President: Herbert G. Baker, Department of Botany, University of California, 
Berkeley, California. First Vice-president: Malcolm A. Nobs, Carnegie Institution of 
Washington, Stanford, California. Second Vice-president: Herbert F. Copeland, Sac- 
ramento City College, Sacramento, California. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley, California. Corre- 
sponding Secretary: Francia Chisaki, Department of Botany, University of California, 
Berkeley, California. Treasurer: John Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1960] KUIJT: ARCEUTHOBIUM 129 


THE DISTRIBUTION OF DWARF MISTLETOES, 
ARCEUTHOBIUM, IN CALIFORNIA 


Jos Kuljt 


There has long been need of a more careful geographical study of the 
western dwarf mistletoes than has been available thus far. This paper 
attempts, first of all, to correct certain misconceptions which have been 
carried along in the relevant literature. Secondly, it represents an effort 
to bring up to date what is reliably known about the distribution of dwarf 
mistletoes in California. Finally, it tries to discover whether there is geo- 
graphical evidence for the existence of host forms in Arceuthobium cam- 
pylopodum Engelm. 

The California distribution of dwarf mistletoes is indeed of crucial im- 
portance in geographical considerations in the genus as a whole. For it is 
in this state that the Pinaceae, a family embracing all known hosts of 
North American dwarf mistletoes, reaches a high level of diversity. The 
geographic peculiarities of the genus in California may thus contribute 
to the understanding of similar features elsewhere. 

The lists and maps contain all Californian collections of A. america- 
num Nutt. ex Engelm. and A. douglasii Engelm. which I have been able 
to find in the herbaria of the University of California at Berkeley and Los 
Angeles (UC and LA, respectively), the California Academy of Sciences 
at San Francisco (CAS), the Dudley Herbarium of Stanford University 
(DS), the two herbaria at Claremont (POM and RSA), the Santa Bar- 
bara Museum of Natural History (SBM), and the United States Depart- 
ment of Agriculture Forest Pathology herbaria at Albuquerque (FPA) 
and Berkeley (FPB). Such an approach is at this time not feasible for A. 
campvlopodum, as hundreds of collections of this species have been made 
in California. I have chosen instead to select a number of collections of 
this species to be recorded here. This selection has been carried out with 
two objectives in mind: firstly, to give as extensive a picture of the dis- 
tribution of A. campylopodum in the state as possible; and secondly, to 
compare the geographic ranges of the supposed host forms of this species. 
Such a procedure does not give a reliable idea of frequency of occurrence. 
This would have been equally true, however, had all collections been taken 
into account, because some recreational areas are represented in herbaria 
by disproportionally large numbers of collections. This is true particularly 
for the Monterey Peninsula. In a similar way, the mistletoe on digger pine 
is much more accessible and much more in evidence than, for example, the 
same species on sugar pine, and therefore the former is present in herbaria 
in numbers exaggerating its relative frequency. I believe therefore that 
selection of specimens, when based on some acquaintance with the species 
as they occur in nature, in this way can give a somewhat more accurate 
idea of geographic distribution. I am also refraining from citing specimens 


Maprono, Vol. 15, No. 5, January 29, 1960. 


130 MADRONO [Vol. 15 


A) 5A Gougias 


| 
| 
| 
E | e 6A. americanum 
| 
| 
| 


Fic. 1. Distribution of Arceuthobium americanum and A. douglasii in California. 


which I have not personally seen. Especially in Gill (1935) some addi- 
tional citations may be found which do not, however, modify the distri- 
butional patterns as here described, except where this author has repeated 
erroneous records from previous publications. 


ARCEUTHOBIUM AMERICANUM Nutt. ex Engelm. 


The lodgepole pine mistletoe is known from the boreal forests of the 
four western Canadian provinces and from all states west of the Rocky 
Mountains. It has been collected from a number of pines as well as from 
Picea glauca (Moench) Voss (Kuijt, 1955), but in California it is known 
from only Pinus contorta, P. ponderosa, and P. jeffreyi (see map, fig. 1). 


1960] KUIJT: ARCEUTHOBIUM 131 


On Pinus contorta Dougl. ex Loud. 

TuLare County. North fork of Kern River, 2150 m., Coville & Funston 1596 
(DS); Bakeoven Meadows, 8100 ft., Howell 27038 (CAS, DS, UC); ridge between 
Monache Meadows and Bakeoven Meadows, 8200 ft., Munz 15220 (RSA); Funston 
Camp, Kern River Canyon, 6700 ft., Peirson 1720 (RSA) ; Junction Meadows, Kern 
River, 8000 ft., Raven 8349 (CAS, UC). Fresno County. Huntington Lake, 7000 ft., 
Solbrig 2420 (UC). Maprera County. Red’s Meadows to Rainbow Falls, 7100 ft., 
Raven 3678 (CAS, RSA). Mariposa County. Merced Lake trail, Yosemite National 
Park, Schreiber 1948 (UC); Little Yosemite Valley, Bolander 5095 (UC), Rodin 877 
(UC). TuoLUMNE County. Glen Aulin Camp, Baldauf (CAS); Morrison Creek, 
Mount Adams, Braekett (LA); YMCA camp, Pinecrest, 5500 ft., Gill (FPA). Mono 
County. 4.3 mi. west of Highway 120-395 junction, Kwzjt 1390 (UC). ALPINE 
County. Camp Wolfeboro, Kuwijt 1411 (UC); 0.5 mi. east of Grade Summit, Kwuzjt 
1412 (UC). Amapor County. East of Lower Bear River Reservoir, Quick 54-88 
(CAS). Ex Dorapo County. Near Lily Lake, Glen Alpine Canyon, Abrams 12753 
(DS, RSA, UC) ; 16 mi. south of Tahoe City, Kuzjt 1335 (UC) ; Highway 50 at Pyra- 
mid Creek, Kwuijt 1513 (UC). PLAcER County. 8.5 mi. west of Soda Springs, Kwzjt 
1330 (UC); 10 mi. north of Tahoe City on Highway 89, Kwuzjt 1331 (UC); 3.5 mi. 
north of Tahoe City on Highway 89, Kuwuzjt 1332 (UC); Tahoe Meadow, Schreiber 
776 (UC). Nevapa County. Near Donner Lake, Dudley (DS); Mayen’s Meadow, 
near Truckee, Sonne (UC); Trout Creek near Truckee, Thomson (CAS) ; Truckee, 
6300 ft. (FPA); Hobart Mills (FPA). Srerra County. 9.5 mi. north of Truckee, 
Kuijt 1344 (UC); Salmon Lake, Sutliffe (CAS); Gold Lake (FPA). PLumMas 
County. 18 mi. north of Greenville on Highway 89, Kuzjt 1350 (UC); Gold Lake, 
Mason 1079 (UC). TEHAMA County. 7 mi. west of Highway 36-89 junction at Lake 
Almanor, Kuwuzjt 1352 (UC); 13 mi. west of Highway 36-89 junction at Lake Alma- 
nor, Kuijt 1354 (UC) ; near Ranger Station, Mineral Campgrounds, Kuwuzjt 1358 (UC) ; 
along Deer Creek, 1.5 mi. south of Highway 32-36 junction, Kuzjt 1501 (UC) ; Spring 
camp near Childs Meadow, Quick 53-129 (CAS). SHAsta County. Thousand Lake 
Basin, 6400 ft., Pezrson 10136 (RSA). Siskiyou County. Military Pass, Mount 
Shasta, Cooke 16034 (DS, UC); north fork, Sacramento River, 6600 ft., Raven 
10456 (CAS). 


On Pinus jeffreyi Grev. & Balf. 
PLACER County. 9.2 mi. north of Tahoe City on Highway 89, Kuizjt 1343 (UC). 


On Pinus ponderosa Dougl. ex Laws. 
TEHAMA County. 13 mi. west of Highway 36-89 junction at Lake Almanor, 
KRuit 1359 (UC): 


Discussion. California represents one of the two southernmost exten- 
sions of the lodgepole pine mistletoe, the other being in the Rocky Moun- 
tain area of Colorado. From a comparison of Figure 1 with the known 
distribution of Pinus contorta (Critchfield, 1957, fig. 26), it is evident 
that at least in California the geographic range of the lodgepole pine mis- 
tletoe corresponds rather closely to that of its most common host. The 
exceptions to this rule are seen in the isolated and apparently healthy 
populations of lodgepole pine in the San Bernardino and San Jacinto 
mountains, California, and in Baja California, Mexico, and of P. contorta 
subsp. bolanderi (Parl.) Critchf. on the Mendocino coast of California. 
It is also notable that the subsp. contorta, although in direct contact with 
subsp. murrayana (Balf.) Critchf. in northern California, has not yet 
been reported as host for Arceuthobium americanum. Indeed this appears 


ey MADRONO [Vol. 15 


to be true outside the state, as I have seen no record of A. americanum 
anywhere in the range of Pinus contorta subsp. contorta (Kuijt, 1956). 
This is more likely to be a case of ecological or spatial isolation than re- 
sistance on the part of the host, as various other species of pine and even 
a spruce may be parasitized by Arceuthobium americanum. Furthermore, 
all subspecies of Pinus contorta are susceptible to Arceuthobium campy- 
lopodum in one or more localities. 

It isa remarkable fact that A. americanum appears to be unable to per- 
petuate itself for long periods of time in stands of Pinus jeffreyi and P. 
ponderosa. Wherever infected individuals of these hosts are found, in- 
fected lodgepole pine is nearby and almost surely is the source of the 
former infections. This is the more remarkable because the brooming 
induced on Jeffrey and ponderosa pine is almost identical to that on 
lodgepole pine (Kuijt, 1958). There are factors quite apart from symp- 
tomatology, therefore, which distinguish the spread of this species of 
dwarf mistletoe in stands of various pine species. Whether these factors 
are climatological, or whether in Jeffrey and ponderosa pine there is a 
greater percentage of resistant trees, is impossible to say at present. 


ARCEUTHOBIUM CAMPYLOPODUM Engelm. 


This species is the most abundant one in California (see map, fig. 2). 
Outside the state it is found from Alaska and British Columbia, largely 
west of the Rocky Mountains, to the Mexican border (and across into 
Baja California). It parasitizes species of Abies, Larix, Pinus, Picea, and 
Tsuga. All but Larix have been reported as hosts for Arceuthobium cam- 
pylopodum in California, but in greatly different frequencies. 

I have attempted, both in the listing and mapping of the specimens 
selected for my purpose, to group herbarium specimens according to the 
affinities of their hosts. In this way I have used the following host cate- 
gories as criteria for division: Abies; Picea; Tsuga; and the three main 
subdivisions of Pinus, namely, the yellow, white, and pinyon pines. I want 
to make clear that such a subdivision is not based on my acceptance of the 
corresponding host forms as recognized by Gill (1935), but rather is an 
effort to test their validity. 


On Abies. 


TULARE County. Cone Peak Camp, Kaweah and Kings River, Dudley (DS) ; 
between Junction Meadows and the hot springs, 7500 ft., Raven 8381 (CAS). FRESNO 
County. Charlotte Creek, Bubbs Creek, Howell 15674 (CAS) ; 2 mi. south of summit 
of Shaver Lake-Dinkey Creek road, Quick 53-27 (CAS) ; Huntington Lake, 7000 ft., 
Wall 67 (CAS). TuoLuMNE County. Southeast of Strawberry Lake, 6500-7200 ft., 
Quick 1734 (CAS) ; southeast of Pinecrest, Quick 50-40 (CAS) ; Tuolumne Canyon, 
Clemens (CAS). AmMapor County. East of Lower Bear Reservoir, Quick 54-87 
(CAS). ALPINE County. Silver Creek, east side of Ebbetts Pass, 6800 ft., Howitt 
(CAS). Et Dorapo County. Near north end of Echo Lake, Howell 22902 (CAS) ; 
near Camino, Kuijt 1272 (UC); Tehoma, Lake Tahoe, Kuijt 1340 (UC). PLACER 
County. Trail to Mount Ellis above Homewood, Schreiber 891 (UC). NEVADA 
County. Baltic Trail, Dudley (DS). PLtumMas County. Taylorsville, Clements (CAS) ; 
8 mi. north of Greenville, Kuzjt 1349 (UC). Butte County. Jonesville, 1600 m., 


1960] KUIJT: ARCEUTHOBIUM 1G) 


hee | A. campylopodum on: 
A : * 
po OQ, ee | 
re es e Sigiiag-tea | e . ° 
has ; i Pinus sect. Diploxylon 
x z hi: ; 

a ae eee | © Pinus sect. Haploxylon 
: i. ee (excl. pinyon ) 
dee pe tees ee Boke e : 

4 ‘ POs : Beate é ; 
Kile. Pa or | + Pinus monophyllia 
© ; as 
} AC ; 
a 4 Abies 
vy Picea 
T Tsuga 
NS 
~ 
s 
+ 
oa 
“= a Cee 


Fic. 2. Distribution of Arceuthobium campylopodum in California. 


Copeland (CAS). TEHAMA County. 11 mi. west of Highway 36-89 junction at Lake 
Almanor, Kwijt 1353 (UC) ; 5 mi. west of Mineral, Kuzjt 1359 (UC). SHasta County. 
Highway 89 at Pondosa junction, Auzjt 1368 (UC); 1.5 mi. east of Viola, Kuzjt 1364 
(UC). Lassen County. 1 mi. below Drakesbad, Mount Lassen, 5400 ft., Cain 114 
(DS) ; Crater Mountain, ca. 15 mi. west of Eagle Lake, 7300 ft., Whitney 1703 (UC) ; 
Mopoc County. Campgrounds at Cedar Pass, Warner Mountains, Alava (UC). 
Siskiyou County. East of Bartle, at border of Modoc National Forest, Newcomb 
156 (UC) ; Sisson southern trail, Mount Shasta, Cooke 13574 (DS) ; summit of Cay- 
enne Ridge, near Marble Mountain, Ownbey 2212 (CAS, DS). HumBorpt County. 
Lasseck’s Peaks, between Goat Camp and Signal Peak, Kildale 2634 (DS); ridges 
east of Corral Prairie, Trinity Summit, Tracy 10571 (UC) ; Eureka, Tracy 3984 (UC). 
Trinity County. South Fork Mountain, Parks & Tracy (UC). MENDOocINO CouNTY. 
Van Damme State Park, Kuijt 1216 (UC). GLENN County. Plaskett Meadows, 6000 
ft., Howell 19283 (CAS). Laxkrt County. Mackie (UC): see the discussion under 
A. douglasii. 


134 MADRONO [Vol. 15 


On Picea breweriana Wats. 


Siskiyou County. Ridge above Applegate Creek, Dry Lake Lookout, 6500 ft., 
Meinecke (FPB). 


On Tsuga mertensiana (Bong.) Carr. 

PLACER County. Emigrant Gap, Jones (POM). TEHAMA County. Mineral, 6300 
ft., Long (FPB). Siskrvou County. Along head of Applegate Creek, north side of 
pass to Fort Gough, 6000 ft., Meinecke (FPB). 


On Yellow Pines. (Pinus sect. Diploxylon: P. attenuata Lemmon, P. con- 
torta, P. jeffres, P. ponderosa, P. coulteri Don, P. sabiniana Doug. ex 
Don, P. radiata Don, P. muricata Don.) 

BAJA CALIFORNIA, MEXICO:! Low hills northwest of La Encantada, Sierra 
San Pedro Martir, 7300-7400 ft., Wiggins & Demaree 5018 (DS, UC, LA); “San 
Pedro Martir”, Brandegee (UC). CALIFORNIA: San Dirco County. Pine Hills, 
near Julian, ca. 4250 ft., Brown (RSA); base of Stonewall Peak, Wiggins 2725 (DS). 
RIvERSIDE County. Idyllwild-Banning, Clokey & Anderson 6574 (RSA, UC); south 
side of San Jacinto Mountains, 5400 ft., Hall 2616 (UC). SAN BERNARDINO COUNTY. 
Mill Creek, Smith 15A (UC); Lake Arrowhead, MacFadden 14737 (CAS). Los An- 
GELES County. Elizabeth Lake Canyon, Liebre Mountains, Dudley & Lamb 4411 
(DS) ; Pine Flats, 20 mi. north of Sierra Madre, Angeles N.F., 6000 ft., Sloan (FPB). 
VENTURA County. Mount Pinos, Hall 6642 (UC). SANTA BaBRARA County. Figueroa 
Mountain, Pollard (CAS). Kern County. Near Havilah, 900 m., Coville & Funston 
1073 (DS); 4 mi. west of Kernville, 4000 ft., Gould 1010 (DS). San Luts Opispo 
County. Cambria, Hoover 6448 (CAS); Santa Margarita, Mason 525 (UC); near 
Paso Robles, Summers 926 (UC). TuLareE County. Near Mineral King, 2700 m., 
Coville & Funston 1460 (DS); Peppermint Valley, Dudley (DS). Fresno County. 
Vermillion Valley, 7700 ft., Raven 5825 (CAS). San BEniTO County. On ridge above 
New Idria reservoir, Kuzjt 1310 (UC) ; on road to New Idria, 4.5 mi. south of Bitter- 
water junction, Kuijt 1300 (UC). MonTEREY County. Pacific Grove, Coleman (DS) ; 
Millers Canyon, Santa Lucia Mountains, 4300 ft., Ferris 12158 (DS). SANTA CLARA 
County. Loma Prieta, Dudley (DS) ; slopes of Mount Umunhun, Ferris 2083 (DS) ; 
Arboretum, Stanford University, Long (DS).2. Contra Costa County. Mount 
Diablo, Abrams 4356 (DS). MAperA County. Ca. 9 mi. west of Oakhurst, Kuzjt 1254 
(UC). Martposa County. Ca. 3 mi. west of Mount Bullion, Kuzjt 1253 (UC); on 
road to Wawona, Jussell 13 (UC). TUOLUMNE County. Cottonwood Meadows, east 
of Mather, Clausen 1777 (DS); near Pinecrest, Quick 55-46 (CAS). CALAVERAS 
County. West of Stanislaus River on road from Vallecito to Columbia, Quick 53-141 
(CAS); 1.5 mi. east of Copperopolis on Highway 4, Kuijt 1410 (UC). Amapor 
County. On road to Buena Vista, ca. 1 mi. south of Ione, 400 ft., Newcomb (UC). 
Mono County. 3 mi. east of Sonora Pass, Kuijt 1432 (UC). ALPINE County. Silver 
Creek Public Camp, Munz 21347 (RSA). Ex Dorapo County. Lily Lake, Glen Alpine 
Canyon, Abrams 12752 (DS) ; near Camino, Kuijt 1273 (UC). PLAcER County. 1 mi. 
north of Tahoe City on Highway 89, Kwijt 1333 (UC). Nevapa County. Spence- 
ville, Eastwood 3420 (CAS). SterrA County. Gold Lake, Barker 254 (DS). BUTTE 
County. Hills near Big Chico Creek, east of Chico, Heller 11144 (DS); Bangor, 
Rose (CAS). PLuMas County. 21.5 mi. north of Sierraville, Kuzjt 1346 (UC). SHASTA 
County. Near Morleys, Baker (UC). Siskryou County. Sisson southern trail, Mount 
Shasta, Cooke 11593 (CAS, DS, UC) ; west of Craggy Mountain, northwest of Yreka, 


1 As far as I can ascertain, these are the only Mexican records of A. campylopo- 
dum. The species A. vaginatum, which is not known from California or Baja Cali- 
fornia, does occur in areas of non-peninsular Mexico. 

2 Introduced into the Arboretum at an early date, since the mistletoe was already 
there at the turn of the century (Peirce, 1905). 


1960] KUIJT: ARCEUTHOBIUM 135 


Ownbey & Brown 2425 (UC). Det Norte County. Gasquet Mountain, Eastwood 
12138 (CAS); Elk Camp Ridge, Parks 24063 (UC). Trin1ty County. Near Scott 
Ranch, Cantelow 1659 (RSA); near Trinity Center, Howell 12790 (CAS). TEHAMA 
County. 15 mi. west of Highway 36-89 junction at Lake Almanor, Kwzjt 1357 (UC) ; 
Manton, Kuijt 1362 (UC). MEnpocrno County. Pygmy forest above Van Damme 
State Park, Kuijt 1215 (UC); Point Arena, Mason 7168 (UC). GLENN CoUNTY. 
5 mi. above Long Point Fire Lookout Station, 3750 ft., Newcomb 148 (UC) ; 2.1 mi. 
north of Stonyford, Kuijt 1506 (UC). LAKE County. Kelseyville, Jussell (CAS) ; 
near Lucerne, Sutliffe (CAS). Cotusa County. Stonyford—Upper Lake road, below 
Old Mill Campgrounds, 3700 ft., Newcomb 146 (UC). Napa County. Mount St. 
Helena, Howell 2204 (CAS); 3 mi. from Aetna Springs on Butts Canyon Road to 
Middletown, Howell 5618 (CAS). Sonoma County. Fort Ross, Mason 4285 (UC). 
Marin County. Inverness Ridge, Howell 19686 (CAS). 


On Pinus monophylla Torr. & Frem. 

San BERNARDINO County. East slope of Providence Mountains, Munz, Johnston 
& Harwood 4272 (POM) ; north slope of San Bernardino Mountains, Parish & Parish 
1442 (DS, UC); Arrastre Creek, 3 mi. southeast of Baldwin Lake, 6700 ft., Jaeger 
(POM); 5 mi. southeast of Ivanpah, Gill & Wright (DS). Ventura County. Sey- 
mour Creek, Mount Pinos, 6000 ft., Pezrson 3251 (POM, RSA). Inyo County. Ca. 3 
mi. west of Lone Pine on Mount Whitney road, Kuzjt 1389 (UC). Mono County. 
6 mi. south of Coleville, Kuzjt 1413 (UC). 


On white pines, excluding pinyon (Pinus sect. Haploxylon, excl. pinyon: 
P. albicaulis Engelm., P. lambertiana Dougl., P. flexilis James, and P. 
monticola Doug]. ex Don). 

SAN BERNARDINO County. West slope, Job’s Peak, 5000 ft., on P. lambertiana, 
Ewan 3564 (POM, DS, UC). Los ANncELEs County. Between Wrightwood and 
Kratka Ridge, Angeles N.F., on P. lambertiana, Embree (UC). Mariposa County. 
Fish Camp Creek, on P. lambertiana, Hedgecock and Meinecke (UC). Mono County. 
On saddle above Convict Lake, 8800 ft., on P. flexilis, Kuijt 1415 (UC) ; 2 mi. south- 
east of Lundy Lake, on P. (?) flexilis, Hendrix 616 (UC). PLAcER County. Near 
Summit, Tahoe N.F., 7500 ft., on P. monticola, Wagener (FPB). SrerrA COUNTY. 
Between Downieville and Forest, 5000 ft., on P. lambertiana, Boyce (FPB). BuTTE 
County. Big Bar Mountain ridge east of Pulga, on P. lambertiana, Quick 53-32. 
PLrumas County. Southeast of Meadow Valley, on P. lambertiana, Quick 53-120 
(CAS) ; Meadow Valley, on P. lambertiana, Weatherby 1667 (UC). TEHAMA CounTY. 
2 mi. north of Hole-in-Ground Campgrounds, on P. lambertiana, Kuijt 1502 (UC). 
SHASTA County. Highway 89 at Pondosa junction, on P. lambertiana, Kuijt 1369 
(UC). Siskryou County. North slope of Shastina, on P. albicaulis, Cooke 11576 
(DS) ; west fork of Molly Creek, on P. albicaulis, Butler 272 (UC); head of Apple- 
gate Creek, 5800 ft., Wagener (FPB). DEL Norte County. Gordon Mountain, 4100 
ft.. on P. monticola, Newcomb 165 (UC). 

Discussion. All major areas of yellow pine in California are infected 
by A. campylopodum. This conclusion cannot be avoided when the speci- 
mens cited above are considered. There are, however, differences in the 
frequency of A. campvlopodum which are not evident from this enumera- 
tion. Such differences are difficult to measure, but are nevertheless recog- 
nizable in the field. 

The only California yellow pine which appears to be free of this mistle- 
toe is P. torreyana Parry ex Carr. I have searched for the parasite in the 
Del Mar area without success, and I have never seen a herbarium speci- 
men with this pine as host. The spatial isolation of this pine from its fellow 


136 MADRONO [Vol. 15 


species would lead one to conclude that its health is not a question of im- 
munity, but rather that isolation from other pines has, at the same time, 
kept Arceuthobium from reaching the Torrey pine.* I am currently germi- 
nating A. campylopodum on some seedlings of this pine, but cannot as yet 
report on it.? 

Infected white pines have been collected a great deal less frequently 
than yellow pines. This is of course partly due to their comparative rarity 
in the state, especially P. albicaulis, P. flexilis, and P. monticola. Pinus 
balfouriana Grev. & Balf. has also been reported as host from Black Butte, 
Siskiyou County (Gill, 1935). The latter author also makes reference to a 
supposed host record of California P. aristata Engelm. by Garrett (1921) 
which, however, is erroneous, as Garrett reports this host only from Bryce 
Canyon, Utah. 

The sugar pine I believe is more frequently infected than the record 
indicates, and this discrepancy is probably due to the inaccessibility of 
infections on this tree. It is nevertheless true, as Gill (1935) pointed out, 
that infected trees of this pine are fairly infrequent and are usually asso- 
ated with other infected members of the Pinaceae. 

The pinyon pines have an even more spotty collection record. This situ- 
ation indeed is evident in the field since infected trees are rare and appear 
to occur in small groups. The interesting fact is that such a small number 
of collections (these are all the pinyon records I have found from Cali- 
fornia) should be so widely spaced. I have heard it said that the pinyon 
pine mistletoe is equally spotty in its occurrence outside California. 

As far as Abies is concerned, it seems to be fairly commonly infected in 
northern California. The incomplete record for some counties in the Sierra 
Nevada I suspect to be due to an infrequency of collecting rather than to 
a rarity of occurrence. This can be checked only by further collections, 
however. The notable fact in this host genus is that the geographically 
most isolated fir, Adies bracteata (Don) Poit., is free of dwarf mistletoe. 
Do we here have a parallel to the situation seen in Pinus torreyana? Is 
the lack of infection of Abies bracteata due to immunity, or to isolation 
from dwarf mistletoe? Cross-inoculations may well supply the answer to 
this question. It is an interesting fact that within a few miles of the A dies 
bracteata populations in the Santa Lucia Mountains the Coulter pine is 
heavily infected with Arceuthobium campylopodum. 

There remain to be considered, finally, the only California records 
known of A. campylopodum on Picea and Tsuga. The small number of 


3 Dr. H. L. Mason reports that the pines of Santa Cruz Island are apparently free 
of mistletoe. 

4 Since writing the above, the inoculations have been inspected and found success- 
ful. The source of the mistletoe seed was Mount Diablo where the dwarf mistletoe 
grows indiscriminately on both Pinus coulteri and P. sabiniana. Seeds were placed 
individually in axils of leaves and fascicles of seedlings of P. torreyana (from Del Mar) 
less than a year of age, on November 6, 1957. At present (January, 1960) a large 
number of mistletoe shoots are present and one pine has died, perhaps as a result 
of heavy infection. 


1960] KUIJT: ARCEUTHOBIUM 137, 


collections of these conifers as hosts, and the distances between their 
localities of origin, would certainly militate against basing host forms 
upon these genera. The three isolated collections on Mountain Hemlock 
are significant in this respect, but the infected Picea breweriana in Siski- 
you County is an even better case in point. According to Gill (1935), the 
nearest known infected spruces are from Idaho and Arizona! 

How then does the geographical evidence bear upon the status of the 
host forms of Arceuthobium campylopodum? It is quite clear that it does 
not, in itself, support the notion. The host forms considered by Gill as 
“minor” forms [f. cyanocarpum (Nelson) Gill, f. blumeri (Nelson) Gill, 
and f. microcarpum (Englm.) Gill] are found within the areas occupied 
by even a single “major” form, f. campylopodum. In fact, f. campvlopo- 
dum is known from practically every county where any of the other forms 
have been collected. Gill admits that his three ‘“‘minor” forms are found 
only in association with other infected species. In California, at least, the 
host forms do not have geographic independence. These facts alone, of 
course, do not preclude racial differentiation as to hosts. 

It is a common field experience to find a heavily infected species of one 
tree together with another species, apparently healthy here, but infected 
elsewhere. This puzzling situation is frequently observed in mixed fir and 
pine stands of the Sierra Nevada and elsewhere. It seems to me that such 
situations more than any other considerations have led to the supposition 
of host forms. In my opinion the taxonomic recognition of such host forms 
ignores two important possibilities. First of all, it fails to take into account 
a possible variation in susceptibility even within a host species. Secondly, 
it largely ignores those isolated but significant instances where, for exam- 
ple, a lodgepole pine has become infected obviously from the heavily in- 
fected fir towering above it. When geographic data are thus considered 
together with the results of the past cross-inoculations (Weir, 1918) 
and natural apparent transfers between Pinus, Picea, Abies, Larix, and 
Tsuga (Kuijt, 1955) it becomes evident that the host forms are not natu- 
ral groups and are, indeed, misleading. The species A. campylopodum 
cannot, in my opinion, be subdivided into natural groups until more is 
known about the resistance differences (if any) both between and within 
host species. 

ARCEUTHOBIUM DOUGLASII Engelm. 

The Douglas fir mistletoe is the rarest of California dwarf mistletoes 
(see map, fig. 1). Its range outside the state shows similarities to that of 
A. americanum. It is found from southern British Columbia to California 
and Arizona and New Mexico. It is not known from the coastal areas of 
the Pacific Northwest, even where the Douglas fir reaches its maximum 
development. It rarely grows on anything but Douglas fir (Pseudotsuga 
menziesi), the only known host from California. 


On Pseudotsuga menziesii (Mirb.) Franco. 
SHASTA County. Highway 89 at Pondosa junction, 4000 ft., Kwijt 1367 (UC). 
Siskiyou County. Southwest slopes of Mount Shasta, 5000 ft., Cooke 13920 (LA) ; 


138 MADRONO [Vol. 15 


north side of Cascade Gulch, Mount Shasta, 5000 ft., Cooke 17729 (CAS); near 
Upton, Mount Shasta, 4000 ft., Hall & Babcock 4078 (UC); 1 mi. east of Highway 
89-99 junction, Kuijt 1371 (UC) ; road to Gumboot Lake, south fork of Sacramento 
River, west of Shasta, 4500 ft., Smith & Bacigalupi (UC) ; west fork of Cottonwood 
Creek, Siskiyou Mountains, Wheeler 2783 (CAS, POM, LA); 2 mi. below Dry Lake 
Lookout, Oak Knoll Ranger Station, 5000 ft., Gill (FPA). 


DiscussIon. So much confusion existes as to this species in California 
that it is necessary first to make a few corrections. 

To begin with, there is the question of Jepson’s (1923) reference to 
Arceuthobium douglasu on Pseudotsuga macrocarpa (Vasey) Mayr in 
southern California. As far as I can discover there is no voucher for this 
suggestion in the Jepson Herbarium or in the University of California 
Herbarium at Berkeley, or elsewhere. In Jepson’s field notebooks there is 
no mention of Arceuthobium douglasii. Whatever the statement was origi- 
nally based on, at present the record is unacceptable. 

A second source of confusion has been Jepson’s (1914) misquotation of 
Engelmann (1880). In this work, Engelmann refers to A. douglasi var. 
abietinum Engelm. as occurring on Abies concolor (Gord. & Glend.) 
Lindl. ex Hildebr. in Sierra Valley, Sierra County. Jepson, however, sim- 
ply repeats this locality for Arceuthobium douglas, and this error is per- 
petuated by Gill (1935). The var. abietinum is undoubtedly referable to 
A. campylopodum, as are the early A. douglasi var. laricis and var. 
tsugense. There is no reliable record of A. douglasii (in the modern sense) 
from Sierra County. 

A third error was first introduced by Jepson (1923) and again repeated 
by Gill (1935). It concerns a specimen collected by Mackie, “Lake Co., 
Aug. 1902, on Pseudotsuga taxifolia.”’ Both Jepson and Gill refer to Ar- 
ceuthobium douglasi in Lake County. Gill’s reference is based on the 
Mackie collection, and Jepson’s probably also. The specimen in question 
(UC 54672) includes some fragmented material, a couple of infected 
branches of Adzes sp. [probably 4. grandis (Dougl. ex Don) Lindl.]|, and 
a cone of Pseudotsuga menziesiu. I have no doubt, both because of the 
flowering condition of the plants and because of their large size, that the 
collection is Arceuthobium campylopodum, and that the Douglas fir cone 
was included by mistake. 

As it stands, then, A. douglasi is known only from the northern part of 
the state. In fact, the only known collection outside Siskiyou County was 
made within half a mile of the county line. There is in this species a most 
striking discrepancy between the geographic ranges of host and parasite. 
The common Douglas fir may be found in the Coast Ranges as far south 
as the Santa Lucia Mountains, and in the Sierra Nevada as far south as 
Big Creek (San Joaquin River), Fresno County. The dwarf mistletoe, al- 
most exclusively restricted to this tree, somehow has not been able to in- 
vade large portions of its host’s range. Whether these extensive areas of 
Douglas tir have remained healthy because of spatial isolation, resistance, 
or climatic barriers, or whether differences in forest composition here play 


1960] STONE: MYOSURUS 139 


a decisive role in limiting the spread of A. douglasti, are questions which 


remain to be clarified. 
Department of Biology and Botany, 
University of British Columbia, 
Vancouver, Canada 


LITERATURE CITED 


CRITCHFIELD, W. B. 1957. Geographic variation in Pinus contorta. Maria Moors 
Cabot Found. Publ. No. 3. 
ENGELMANN, G. 1880. In Watson, S. Botany of California, 2:106-107. 
GarreETT, A. O. 1921. Forest tree diseases. Trans. Utah Acad. Sci. 2:182-189. 
Girt, L.S. 1935. Arceuthobium in the United States. Trans. Conn. Acad. Arts & Sci. 
32:111-245. 
Jepson, W.L. 1914. A flora of California. San Francisco. 
. 1923. A manual of the flowering plants of California. Berkeley. 
Kurjt, Jos. 1955. Dwarf mistletoes. Bot. Rev. 21:569-628. 
. 1956. A new record of dwarf mistletoe on lodgepole and western white 
pine. Madrono 13:170-172. 
. 1958. Morphological aspects of parasitism in the dwarf mistletoes (Arceu- 
thobium). Dissertation, Univ. Calif., Berkeley. 
Peirce, G.J. 1905. The dissemination and germination of Arceuthobium occiden- 
tale Eng. Ann. Bot. 19:99-113. 
WErr, J. R. 1918. Experimental investigation on the genus Razoumofskya. Bot. Gaz. 
66:1-31. 


NUCLEAR CYTOLOGY OF THE CALIFORNIA MOUSE-TAILS 
(MYOSURUS)! 


DONALD E. STONE 


INTRODUCTION 


Published accounts of the chromosome numbers in the genus M yosurus 
are limited to three brief reports concerned exclusively with European 
representatives. In the 1945 edition of the “Chromosome Atlas,” a single 
citation (Gregory, 1941) noted the chromosome number of M. minimus as 
n=—8. A check of Gregory’s paper, however, reveals that Myosurus was 
one of the few genera in the family for which he had no first hand 
information. Instead, his citation is based upon the work of Mann (1892) 
and Hocquette (1922), who found n=8 and 2n=—16 respectively. The 
haploid number was published by Mann as a footnote to his figure 5: 
‘“Monaster stage of archesporium, with 8 chromatin segments.” Hoc- 
quette’s account was likewise lacking in details, as his study was part of a 
general survey of the Ranunculaceae. 

The third reference to original work is in the 1955 edition of the 
“Chromosome Atlas.” It is of interest to note that here the earlier cita- 
tions of Mann and Hocquette are dropped in favor of a more recent 


1 Part of a dissertation submitted to the University of California at Berkeley as 
partial fulfillment of the requirements for the degree of Doctor of Philosophy. 


140 MADRONO [Vol. 15 


count by Ehrenberg (1945). Working on Swedish material, Ehrenberg 
found about 28 chromosomes in the somatic cells. His counts of nine cells 
showed variations of from 27 to 30 chromosomes, with the best three slides 
having 28, 28 and 29. He suggests that the Swedish material is tetraploid, 
being derived from a diploid race with a base number of 7. Hocquette’s 
report of a haploid number of 8 is considered to offer little difficulty as the 


TABLE 1. Myosurus SPECIMENS CYTOLOGICALLY EXAMINED AND DOCUMENTED.1 


Collection Data N 2N Fig. 


M. sessilis Watson 
Stone 1(14): 5 April 1953, 3 miles east 


of Maxwell on the Maxwell Road, Colusa County.* 16 1 

Stone 1(10): same data as above. 8 4&5 
M. sessilis subsp. alopecuroides (Greene) Stone 

Stone 7(22): 10 April 1953, same locality 16 — 

as above. 8 6&7 


M. minimus subsp. apus (Greene) Campbell 
Stone 5(1): 10 April 1953, Manning Flat, 
514 miles west of Lower Lake on the road to 
Kelseyville, Lake County 8 8 
Stone 5(2): same data as above. 8 9 
H.L. Mason 14275(2): 26 April 1952, 5 miles 
northeast of Crows Landing on the Crows Landing 
Road, Stanislaus County. 16 — 
M. minimus L. 
J. Lid, Stone 73(1): 21 June 1955, Hud Island 
Vestfold County, Norway. 16 11 
Stone 3(5): 4 April 1953, Manning Flat, 5% miles 
west of Lower Lake on the road to Kelseyville, 


Lake County. 8 10 
H. L. Mason 14501(4): 2 April 1953, 3 miles east 
of Hanford on the road to Visalia, Kings County. 8 — 


M. minimus var. filiformis Greene 
Stone 7(13): 10 April 1953, 3 miles east of Maxwell 


on the Maxwell Road, Colusa County. 16 2 

Stone 9(2): Spring, 1953, Ajax Field in Willows, 

Glenn County. 8 12-14 
M. aristatus subsp. montanus (Campbell) Stone 

R. Bacigalupi 4238, Stone 15(5): April, 1953, 16 — 

Big Bear Lake, San Bernardino County. 8 15 
M. cupulatus Watson. 

T. Robbins 3480, Stone 17 (1): 29 April 1952, 8 16 

Providence Mountains, San Bernardino County. 16 3 


1 Specimens documenting the chromosome counts have been deposited in the 
Herbarium of the University of California at Berkeley. 
* All localities are in California unless otherwise noted. 


closely related genus of Ranunculus, which like Myosurus has large- 
Ranunculus-type chromosomes (Langlet, 1932), ranges from n=7, n=8 
to n=64 (Darlington and Ammal, 1955). 

When the problem of the existence of sympatric biotypes was first sug- 
gested (Stone, 1959), it was hoped that cytology might provide some 


1960] STONE: MYOSURUS 141 


M, sessilis 
* o — |! e2e3 4 5 6 7 8 
6 Lu sm st sm sm sm_ st sm_ st 


M. m. filiformis 


M. cupulatus 


Whitt 


Fics. 1-3. Somatic metaphase chromosomes and idiograms of three Myosurus 
species: 1, M. sessilis, shoot-apex squash, x 2000; 2, M. minimus var. filiformis, root- 
tip squash, &K 2200; 3, M. cupulatus, root-tip squash, x 1360. 


142 MADRONO [Vol. 15 


ee ee ee eo ieee ee oem Se ae 


; 


6 - 7 


Fics. 4-7. Meiotic configurations of M. sessilis and subspecies: 4, M. sessilis, diaki- 
nesis, X 1850; 5, M. sessilis, metaphase I, * 1850; 6 and 7, M. sessilis subsp. alope- 
curoides, diakinesis, & 1850. 


1960] STONE: MYOSURUS 143 


clue to the mechanisms involved in isolation. This hope, unfortunately, 
was not realized. A survey of chromosome numbers in nine heterogeneous 
California valley populations (Mason, 1957; Stone, 1957), representa- 
tives of two high-mountain species, and a collection from Norway, how- 
ever, showed only diploid plants with n = 8 and 2n = 16. Although no 
deviations from the basic number of 8 were found, it is possible that 
specialized biotypes found in disjunct pools throughout California might 
prove to be exceptions. Additional information was sought in a karyotype 
study of three of the most extreme morphological types (figs. 1, 2 and 3). 
Here again, no differences could be established. 


MATERIAL AND METHODS 


All of the material examined cytologically was grown in the Botany 
Department greenhouse, University of California, Berkeley. 

Mitotic stages were most readily obtained from the root tips of young 
seedlings or from the embryonic tissues of leaf bases and shoot apices. 
Fixation with acetic-alcohol (1:3) and staining with iron aceto-carmine 
proved satisfactory in root-tip squashes of young seedlings. Root tips of 
more mature individuals, however, were extremely difficult to squash, 
hence special techniques were found necessary. The following four-step 
process worked well on material examined immediately after squashing: 
(1) fixation of root tips in acetic-alcohol for 24 hours; (2) pre-staining 
of the material in aceto-carmine at 60° C. for 2 hours: (3) hvdrolvzing 
in IN HCl at 60° C. for 1 hour; and (4) washing in distilled water for 
15 minutes, after which the material was stored in 70 per cent ethanol. 
Processed root tips were then squashed using additional iron aceto-car- 
mine stain. Cells hydrolyzed in such a manner have light-stained nucleoli 
and dark stained chromosomes, and thus are quite favorable for observa- 
tion of chromosome-nucleolar associations in mitotic prophase. Appar- 
ently there is a differential reduction of acidity in the pre-stained nucleus 
during the hydrolysis (Rattenbury, 1952). Due to the obvious difficulty 
of chromosome distortion in squashes, paraffin-section methods were tried. 
However, the minute size of the secondary roots (0.1—0.2 mm. in diam- 
eter) and the restricted meristematic region made sectioning efforts fruit- 
less. 

Stages of microsporogenesis were used in the study of the meiotic 
chromosome behavior. When obtained, active pollen mother cells were 
extremely useful in determining chromosome number, size, and pairing 
relationships. Three features affecting satisfactory results should be 
noted: (1) the period of active microsporogenesis; (2) the position of the 
bud; and (3) the size of the bud and stamens. 

The period of microsporogenesis was found to be of extremely short 
duration. Out of a total of four or five young buds on a plant it was 
common to find that most had already matured, while the remainder were 
premeiotic. Possibly, poor greenhouse conditions were responsible for the 
shortened meiotic period, but judging from the luxuriant specimens, this 


144 MADRONO [Vol. 15 


: lO lI 


Ee 


Fics. 8-11. Meiotic and mitotic configurations of M. minimus and subspecies: 
8, M. minimus subsp. apus, metaphase I, &* 1850; 9, M. minimus subsp. apus, diaki- 
nesis, X 1850; 10, M. minimus, diakinesis, * 1850; 11, M. minimus, mitotic meta- 
phase, 2800. 


1960] STONE: MYOSURUS 145 


I2 13 
mon __ : | 
( re ” 
¢ 
; ~ eel 
Nie 


14 


Fics. 12-14. Meiosis in M. minimus var. filiformis: 12-13, diakinesis, x 1850; 
14, anaphase II, * 1500. 


146 MADRONO [Vol. 15 


16 


Fics. 15-16. Meiosis in two montane species of Myosurus: 15, M. aristatus subsp. 
montanus, diakinesis showing attachment of chromosome No. 1 (large satellited one) 
with nucleolus, & 2000; 16, M. cupulatus, metaphase I, * 2000. 


does not seem likely to be the case. Perhaps a condition such as this, where 
meiosis occurs at a very early stage and over a short period of time, has a 
selective advantage in plants that survive in ephemeral environments 
such as vernal pools. 

In all biotypes of Myosurus examined, meiosis occurs before peduncle 
elongation takes place. The meiotic buds are found buried deep in the 
basal rosette of leaves and peduncles of the more mature flowers. Meiotic 
buds are usually less than 1.5 mm. in length and hence are extremely 
difficult to find and remove. Killing and fixing of the entire plant, however, 
was found practical. Buds were removed under a dissecting microscope at 
30, and flower dissection was completed at 60%, with one stamen 
(anthers 0.3—0.5 mm. long) at a time being removed for squashing. Plants 
used for meiotic studies were fixed either in acetic-alcohol or in Linnert’s 
fixative. 

The photomicrographs in the accompanying figures were made using a 
Bausch and Lomb compound microscope having either a 90 (N.A. 1.30) 
or a 60 (N.A. 1.40) apochromatic objective and a 15 compensating 
eyepiece. The magnification of each figure is noted in the legend of the 
figure. 

RESULTS 
Mitotic Chromosome Number and Morphology 


Detailed cytological studies were limited to the Manning Flat, Maxwell, 
and Willows populations (Table 1), but a survey of additional biotypes 
from the other California Valley populations established a single chromo- 
some number of 2n = 16. The photographs of figures 1-3, and 11, are rep- 
resentative of the mitotic squashes that were observed in this study. The 


1960] STONE: MYOSURUS 147 


generic karyotype consists of 5 submedian and 3 subterminal chromo- 
somes. The largest of the set bears a conspicuous satellite. The idiograms 
are based on the average of the homologous pairs of chromosomes, as 
measured in the corresponding photographs. The arbitrary classification 
of the centromere is based on the relative length of the two arms (Good- 
speed, 1945); median (m), arm ratio 1:1; submedian (sm), arm ratio 
greater than 1:1 but less than 3:1; subterminal (st), arm ratio 3:1 or 
greater. It is quite apparent that although the satellited chromosome fits 
in the submedian class, it is very close to the median class, and for all 
practical purposes it can be considered as such. The conspicuous uniform- 
ity in the size gradient from the large satellited chromosome to the small- 
est subterminal chromosome is common to all three taxa. The most 
notable difference between the idiograms is the absolute size of the 
chromosomes. For example, the satellited chromosome is about 6 microns 
in figure 1, 6 microns in figure 2, and 7 microns in figure 3. As cell size 
and chromosome size are more or less interdependent and seem to fluctu- 
ate considerably in the same plant, no significance was attached to the 
slight differences in length. 

Examination of Norwegian material (fig. 11) has proven the chromo- 
some number to be identical to that of California specimens. Ehrenberg’s 
polyploid counts still remain to be verified. 

Meiotic Chromosome Number and Pairing Relationships 

Diakinesis (figs. 4, 6, 7, 9, 10, 12, 13, and 15) was by far the most 
common meiotic stage encountered. In part, this occurrence might be 
attributed to the selected time of fixation: it was found that best results 
were obtained if fixation was limited to the time between 12 noon and 2 
p.m. Infrequently, metaphase I stages (figs. 5, 8, and 16) were found. 
Later stages in the meiotic sequence were so rare that only two pollen 
mother cells were observed in the anaphase II stage (fig. 14). The second 
meiotic anaphase is frequently useful in denoting karyotype differences 
(Chambers, 1955) and in the case of figure 14 it is possible to verify 
centromere positions established in mitotic preparations. All of the figures 
show 8 pairs of chromosomes with no indication of pairing difficulties. 
It is of interest to note the association of the large satellited chromosome 
(No. 1) with the nucleolus in the diakinesis figures. 


SUMMARY 


Mitotic and meiotic chromosome counts have been made for each of 
seven taxa of Myosurus, on six of which no counts have previously been 
reported. All examined specimens of the genus Myosurus displayed a 
diploid number of 16, and a haploid number of 8 chromosomes, with no 
meiotic irregularities. 

Tulane University, 
New Orleans, La. 


LITERATURE CITED 


CHAMBERS, K. 1955. A biosystematic study of the annual species of Microseris. 
Contr. Dudley Herb. 4:207-312. 


148 MADRONO [Vol. 15 


DariincTon, C. D., and E. K. JANAKI AMMaAL. 1945. Chromosome atlas of cultivated 
plants. Allen and Unwin, London. 397 pp. 

DarLincToNn, C. D., and A. P. Wy ir. 1955. Chromosome atlas of flowering plants. 
Allen and Unwin, London. 519 pp. 

EHRENBERG, L. 1945. Kromosomtalen hos nagra karlvaxter. Bot. Not. 4:430-437. 

GoopsPEED, T. H. 1945. Chromosome number and morphology in Nicotiana. VII. 
Karyotypes of fifty-five species in relation to a taxonomic revision of the genus. 
Univ. Calif. Publ. Bot. 18:345-367. 

Grecory, W. C. 1941. Phylogenetic and cytological studies in the Ranunculaceae. 
Amer. Phil. Soc. Trans., New Series, 31:443-521. 

HocQuETTe, M. 1922. Observations sur le nombre des chromosomes chez quelques 
Renonculacées. C. R. Soc. Biol. Paris, 87:1301—1303. 

LANGLET, O. 1932. Uber Chromosomenverhiltnisse und Systematik der Ranuncula- 
ceae. Svensk. Bot. Tidskr. 26:381-400. 

Mann, G. 1892. The embryo sac of Myosurus minimus L. Trans. and Proc. Bot. Soc. 
Edinburgh, 19:351-428. 

Mason, H. L. 1957. Flora of the marshes of California. Univ. of Calif. Press, Berke- 
ley. 878 pp. 

RATTENBURY, J. A. 1952. Specific staining of nucleolar substance with aceto-carmine. 
Stain Tech. 27:113-120. 

STONE, D.E. 1957. Studies in population differentiation and variation in Myosurus of 
the Ranunculaceae. Ph.D. Thesis. Univ. of Calif., Berkeley. 

. 1959. A unique balanced breeding system in the vernal pool mouse-tails. 

Evolution 13:151-174. 


VARIATION IN SECTION TRIGONOPHYLLAE OF NICOTIANA 
Puitiep V. WELLS 


Section Trigonophyllae of the genus Nicotiana is peculiar to the warm 
deserts of southwestern North America, and ranges from California to 
Texas and southward locally as far as Oaxaca. The section, as defined by 
Goodspeed (1954), includes two species: N. trigonophylla Dunal, the 
range of which coincides with that of the section, and NV. Palmeri Gray, 
which is apparently found only in southwestern Utah and western Arizona. 

During the course of an ecological investigation of NV. trigonophylla 
throughout its range in the United States, the writer encountered facts 
which cast doubt on the validity of the specific rank of the taxon JN. 
Palmeri. 

The two members of the section Trigonophyllae are segregated as fol- 
lows by Goodspeed (1954) in his key and text: 

Calyx 8-11 mm. long; corolla 12-23 mm. long, limb 3-4 mm. wide, erect in 

bud; seed ca. 0.5 mm. long; cauline leaves obtuse to acuminate....V. trigonophylla 


Calyx 15-17 mm. long; corolla 23-32 mm. long, limb 5-6 mm. wide, oblique in 
bud; seed ca. 0.7 mm. long; cauline leaves acute to acuminate........ N. Palmeri 


Both taxa have the same chromosome number (12 pairs) and Kostoff 
(1943) reported that Fi hybrids between the two show twelve homolo- 
gous pairs of chromosomes at meiosis. 

The writer visited several of the major herbaria! of the United States 
and examined the collections of Nicotiana, section Trigonophyllae. Only 


1960] WELLS: NICOTIANA 149 


nine collection numbers were encountered which were labelled or anno- 
tated as NV. Palmeri. Of these, one was an intermediate mentioned by 
Goodspeed (1954), and of the remaining eight specimens only two pos- 
sessed corollas appreciable larger than those of NV. trigonophylla: 1) Keck 
4255 (UC) Gillespie Dam, Maricopa County, Arizona, March 22, 1936. 
“Tn lava rock at cliff base.” 2) Gould 1611 (NY) St. George, Washington 
County, Utah, April 20, 1942. “Southern slope of Black Hill... on rock 
ledges and among volcanic boulders.” Both of these specimens have corol- 
las more than 30 mm. in length. This is larger than the type collection 
(Palmer 433), which is intermediate between these extremes and JN. 
trigonophylla. 

In the herbarium at Dixie College at St. George, Utah, there were 
seven collections of section Trigonophyllae from the basalt-capped mesa 
just west of St. George (the site of Gould 1611). Of these, only one was 
labelled NV. Palmeri; the other six (including one determined by I. Tide- 
strom) were labelled N. trigonophylla. None were more than interme- 
diate between the two taxa. On the other hand, the writer has grown 
plants from seed collected from this same site which produced corollas 
27 mm. in length, which is within the size range for the flowers of JV. 
Palmeri. 

Specimens and seed were collected over a large part of the range of 
section Trigonophyllae in the United States. Measurements of various 
taxonomic characters were made both on collected plants and on plants 
grown from seed in the greenhouse. The results are presented in Table 1, 
where a number of size classes are established for each character investi- 
gated. The number of measurements falling within each size class is given, 
thus illustrating the modal class and the range of variation. For the 
greenhouse-grown plants each figure represents the number of plants 
having that particular mean character size, while for the collected speci- 
mens, each figure indicates the number of measurements falling in a size 
class. 

The populations investigated show a trend toward N. Palmeri char- 
acters as one approaches the Washington County, Utah area. Whether 
one regards NV. Palmeri as a distinct species or prefers to sink it to the 
subspecific or varietal level, it is apparent that the two taxa are not clearly 
delineated, but intergrade with respect to all characters measured. This 
morphological intergradation is probably best interpreted in a topoclinal 
sense, since no ecological gradients appear to be involved. In the range of 
N. Palmeri, section Trigonophyllae occupies its three usual ecological 
niches: 1) bedrock outcrops and talus; 2) dry washes; 3) ruderal sites 
(roadsides, etc.), but it is most prevalent in the first mentioned (Wells, 
1959). 


1 The following institutions were visited: Bureau of Plant Industry (Beltsville, 
Md., Missouri Botanical Garden, St. Louis, New York Botanical Garden, Rancho 
Santa Ana Botanical Garden, Claremont, California, University of California, Berke- 
ley, and the United States National Herbarium, Washington, D.C. 


N 
WwW 
(S9) 
s+ 
ne) 
mH 
— 
N 
N 
= 
N 


yey ‘uoysuryse 
qeip) “uoysuryse A 
I yey ‘uoysuryse 

if ie yey ‘uosuryse 

T I I c I yey ‘uoysuryse A 
T BpeaAIN ‘YARD 

YyeiQ ‘uo urIyse A 

I BpeaAdN “YyAe[O 
I Sexo], ‘1oJsMoIg 

I BUuOZIIY ‘asIyIod 
euoZzly ‘eUulld 

Sexa], ‘Io}SMOIg 

sexo], ‘a[qury 

euoziiy ‘[eulg 

euozly ‘eWIg 

c OoTxeT, MON ‘APPA 
I T sexay, ‘uosIeq{ng 
RIUIOJI[LD ‘IPISIOANY 
I T PIUIOJI[ED ‘OUIPIVUJEg UBS 


[Vel. 15 


bol 
lee) 

al al 

= Ss OA MN 
iS) 


ise) 
~onnw~rtntawtwnst 
So 
+ 
Set NN 
Saal 
Sm NNN NY 


mn~ OM Net M CO MON 
Soe | 
™N 
— 
N 
MAAN Te ANN 


~ 


MADRONO 


a a RS 


COs OMe O 905 0 St tt ¢f crt It OT 6 8 LO’ 9C Se FC $e €¢ -1e “Oc GI Sk ZE 9ST 
“wu “paas Jo yysuaT ‘mu ‘xATRD JO YBUIT “uu “(QUIT SUIPN[Ox9) YWeLeg JO yyWuUeT 


SNAWIOUdS GHLOAXTTIO;) 


i> 6) 


N 
[3 ) 
- 


I [ae ee: ye} ‘uo uUrIYyse 


N 
+ 
N 


BPRAIN ‘YIPTO 
b I PIUIOFe) ‘OAUT 
Z ¢ Z BUOZLIY ‘RWI 
Z (iar me | lo VEare Sexay “a[qury 


_— 
=a NN ™X © 
Mmm ON 
~— 
+ 
man TN 
cn oe oe 
a 
NN OG 
_ 
= es OQ 
Sal ~*~ 
~N 
> 
_— 
Sn ions Iie iio] 
al 
eal 
el 
N 


ov ea OFS ei Sel Or “6 Sh tie Cie cl Sh Ol Gn 8 2670 Ce hee SZ cece cs 0G) Ol. Sh Er OT 


She ae “WU “UPI, QUIT [RIO], ‘wu ‘xATBD JO YSuUeT “ul (QUIT, SUIPN]OXd) P[[OIOD Jo YyIBuaT (Ayunog Aq) sant[eoTy 
; ASNOHNATH NI NMOUD) SNAWIOTdS 


150 


DONVY SLT 40 SLUVG INAXAAAIC, NI DUD210I1N7 AO AVTTAHdONOODINT NOTLOUS NI SYALOVUVH.) OIWONOXVY, NI NOILVINVA ‘T FTAV EE 


1960] WELLS: NICOTIANA Pot 


The tendency of plants of section Trigonophyllae to occur in small, 
isolated populations, and the localization of plants showing NV. Palmeri 
characters in certain northwestern portions of the range of the section, 
suggest the operation of the Sewall Wright effect in bringing about dif- 
ferentiation. A related possibility is the selection of larger flowers by some 
local pollinator in the V. Palmeri range. 

The taxon NV. trigonophylla Dun. is acknowledged by Goodspeed 
(1954) to be a variable one. With respect to the validity of N. Palmert, 
Goodspeed (1945) wrote: “...N. Palmeri is morphologically so closely 
related to NV. trigonophyila as doubtfully to deserve specific recognition 

..”. In his 1954 monograph, he wrote: “This species (N. Palmeri) is 
obviously close to NV. trigonophylla. It is distinguishable even from large 
flowered races of the latter by the greater coarseness throughout, by 
longer corolla, and by broader, whiter, more horizontal limb with lobes 
at times slightly concave. Maguire and Blood 1456, 15 miles SW of Leeds 
(Washington Co.), Utah (UC) is an example of an intermediate between 
the two.” 

In the herbarium of the New York Botanical Garden, there is a series 
of collections of section Trigonophyllae from Sonora and Baja Cali- 
fornia. On one of the herbarium sheets (MacDougal 41), there are some 
remarks by a reviewer of these collections (unsigned). In summary, he 
finds the material “not uniform but contains 2 forms with very different 
pubescence. One is densely glandular villous-tomentose and is so oily as to 
heavily stain collecting paper. The two forms deserve nomenclatural rec- 
ognition, but at present it seems impossible to determine which is typical 
form. NV. Palmeri Gray of Arizona seems intermediate in its characters, 
but nearest to eglandulose form.” The writer also found wide variation 
in several characters (including flower size) in the Mexican collections of 
section Trigonophyllae. It seems likely that an intensive study of the NV. 
trigonophylla complex in Mexico might uncover variants at least as di- 
vergent as the currently accepted NV. Palmert. 

Considering the variability of N. trigonophylla and the continuous 
intergradation between it and NV. Palmeri, and also the very meager rep- 
resentation of the latter taxon in herbaria, it does not seem too conserva- 
tive to relegate NV. Palmeri to subspecific or varietal status. This, in fact, 
has already been done by Marcus E. Jones (1908) who reduced NV. 
Palmert Gray to N. trigonophylla Dun. var. Palmeri (Gray) Jones. 

Department of Biological Sciences, 


California State Polytechnic College, 
San Luis Obispo, California. 


LITERATURE CITED 
GoopsPEED, T. H. 1945. Cytotaxonomy of Nicotiana. Bot. Rev. 11:533. 
1954. The genus Nicotiana. Chronica Botanica, Waltham, Mass. 
Jones, M. E. 1908. Contr. West. Bot. 12:52. 
Kostorr, D. 1943. Cytogenetics of the genus Nicotiana. State Printing House, Sophia. 
WELLs, P. V. 1959. An ecological investigation of two desert tobaccos. Ecology 40: 
626-644. 


152 MADRONO [Vol. 15 


STUDIES ON SECOTIACEOUS FUNGI VII. 
SECOTIUM AND NEOSECOTIUM. 


RoutF SINGER AND ALEXANDER H. Sm1TH! 


We have had occasion to mention the genus Secotium in the narrower 
sense as based on Secotium gueinzii Kunze from South Africa. These 
references to S. gueinzi in the preceding parts of our studies on secotia- 
ceous fungi were motivated by the necessity of comparing the characters 
of the type species of the genus with the characters of the other genera 
of the family as proposed in our previous papers. 

We have now arrived at the question: what is the position of Secotium 
gueinzi, and which other species of Secotium are close to it? 

In the first place we wish to redescribe the species to facilitate the 
comparison. 


SECOTIUM Kunze, Flora 23:321. 1840. 
SECOTIUM GUEINzII Kunze, l.c. p. 322. 


Gastrocarp convex, truncate below or not, always deeply sinuate under- 
neath along the apex of the stipe (like Endoptychum depressum), semi- 
globose to campanulate, 30-60 mm. broad, about 32-42 mm. tall. 

Peridium (inner) tessellate (with canal-like depressions), not gelatin- 
ized, white-buff (dried fuscidulous-yellowish), sometimes covering the 
gleba completely, sometimes pulled back (down) to expose some part of 
gleba; gleba loculate, with small chambers which do not show lamellar 
arrangement in any form but are winding, irregular, and unequal, not 
pulverulent, the walls white, thin, ochraceous buff to light brownish and 
said to have been pale olive, the exposed surface of gleba (if any) ver- 
tically surrounding the apex of the stipe. 

Stipe up to 70 mm. long and 5—10 mm. broad at apex, up to 22 mm. 
broad at base, buffish colored, apparently smooth and glabrous, dry, 
stuffed; columella continuous with the stipe and either percurrent or 
not, if not, then sending out tramal plates with thick branches which 
merge with the normal thin tramal plates making up the loculi, at times 
changing direction in relation to the stipe and becoming oblique rather 
than vertical, broadened into the upper portion of the peridium if per- 
current, white; volva said to be present, whitish, eventually disappearing 
except at the base of the stipe where it appears cothurnate, a distinct 
annular veil such as seen in Endoptychum depressum not described, not 
seen in the fragments available, and not clearly shown in illustrations. 
Context white, dry-fleshy, probably when fresh somewhat like Endop- 
tychum agaricoides. 

Spores (8.5—) 11-14 & (6.3—) 8.2—9.7y, short ellipsoid but ovate in 
frontal view, somewhat inequilateral in profile (asymmetric), with 
oblique eccentric sterigmal appendage, pale olive-melleous to melleous- 


1 Papers from the University of Michigan Herbarium and the Department of 
Botany, No. 1087, University of Michigan, Ann Arbor, Michigan. 


1960] SINGER & SMITH: NEOSECOTIUM 153 


hyaline, the wall thick and complex (at least four wall-layers discernible ) , 
smooth, some with an apical germ pore, or some with an apical truncation 
but without a demonstrable discontinuity in the wall (or else pore incom- 
plete and spore not truncate), slightly metachromatic in cresyl blue but 
absorption of the cresyl blue very variable, if weakly stained, a lilac line 
along the endosporium visible and inner two layers sometimes remaining 
incolorous, when strongly stained the whole wall and interior deep blue as 
in spores of Chlorophyllum molybdites, correspondingly, with Melzer’s 
reagent distinctly pseudoamyloid but some remaining inamyloid and a 
few discoloring only partly; not forming a pulverulent mass in the gleba. 

Basidia about 28 « 9.7u., 4-spored, sterigmata variable, some thin and 
straight but oblique, some thin and slightly curved (somewhat interme- 
diate between “‘agaricoid” and “gastroid”’) ; cystidia not seen, but yellow 
“Pollinarien” described and illustrated by Corda. 

Trama hyphous throughout, in places very slightly gelatinized, hyaline, 
in peridium extremely irregular but more radially arranged in outermost 
layer of endoperidium, the hyphae of all layers 2—6 y. in diam., inamyloid 
and with clamp connections. 

Terrestrial on the sand steppes of the Cape of Good Hope, South Africa, 
Uitenhage, fruiting in summer (December). Leg. Queinzius, comm. M. 
C. Cooke (NY, part of type). 

Another part of the type is at Kew (Singer has seen but not studied it, 
but he is certain that it is part of the same collection, which is corrobo- 
rated by the fact that what little Heim communicates about the “‘Berke- 
ley-type”’ coincides well with our findings). It is possible that part of this 
was also in Corda’s Herbarium which is in Prague. 

This species has the same essential spore-wall characters as Endop- 
tyvchum agaricoides and FE. arizonicum, namely the pseudoamyloid reac- 
tion, the thick, many-layered wall, and the relatively light color (varying 
to hyaline). In spite of these similarities, there are important differences 
such as the presence of a volva, non-pulverulent gleba, abundant clamp 
connections, and large ellipsoid spores with a germ pore. In view of these 
differences it appears illogical to us to place S. gueinzii in the same genus 
with Endoptyvchum. This was apparently also Zeller’s point of view. 

However, two other species, intermediate in their characters, need to 
be considered here. One is Secotium macrosporum Lloyd. It is interme- 
diate in such basic characters as the pulverulent gleba and degree to 
which clamps are present, but is strikingly distinct because of the comp'ex 
ornamentation of the spores. With some modifications the spore orna- 
mentation is the type that is found in some tropical Lepiotas (Leucoagari- 
cus) and/or (this is significant), in the Lycoperdaceae. We shall discuss 
the affinities of this interesting species later, but considering its differ- 
ences from both Endoptyvchum and Secotium, we cannot convince our- 
selves that according to any generic concept short of re-establishing 
Secotium sensu lato, can S. macrosporum be considered congeneric with 
either Secotium or Endoptychum. 


154 MADRONO [Vol. 15 


This establishes Secotium as a monotypic genus, and necessitates estab- 
lishing a new genus for S. macrosporum. We propose for it the new generic 
name NVeosecotium, this being a New World Secotium and a species only 
now critically analyzed (Neo—new; secotium—a loculate system). 


Neosecotium gen nov. 

Carpophoris haud volvatis, pallidis, stipitatis, columella percurrente; 
gleba demum paulum vel manifeste pulveracea; sporis hyalinis leviter 
ochrascentibus, pseudoamyloideis, poro germinativo instructis, globosis, 
membrana admodum crassa reticulatim fracta ornamentatis; fibulis 
praesentibus sed sparsis. 

Typus generis: Secotium macrosporum Lloyd. 


Neosecotium macrosporum (Lloyd) Sing. & Smith, comb. nov. 
Secotium macrosporum Lloyd, Mycol. Writ. 1:139. 1903. 

Gastrocary 1-3 cm. high and 1-2 cm. thick, subelliptic to nearly 
globose, the margin not separating from the stipe-columella. 

Peridium smooth, avellaneous or paler, the lower portion whitish-pallid 
at times. 

Gleba chambered at first, but somewhat pulverulent and at maturity 
little if any structure visible, in immature stages showing chambers ori- 
ented in an obscure lamellar orientation, about wood brown (R) near 
maturity or finally becoming more cinnamon, not separated from 
columella. 

Stipe-columella percurrent, pallid throughout as dried, 2—3.5 mm. diam. 
in widest place (as dried), very little (2-3 mm.) projecting below the 
gastrocarp as a Stipe. 

Spores globose and 13.5-18 ». or 14-18 & 12-15 yu. and subglobose to 
slightly ovate, ochraceous to tawny in KOH (depending on degree of 
maturity), dark red-brown in Melzer’s solution (pseudoamyloid), with 
a short to rather long sterigmal appendage as in many Lycoperdaceae, 
the pedicel hyaline and thin-walled except for the area where the thick- 
ening of the spore wall projects down into it slightly, centrally attached 
or rarely slightly eccentric; spore wall complex, at maturity with a thin 
hyaline perisporium which adapts itself to the configuration of the wall 
beneath (exosporium and possibly endosporium combined) ; exosporium 
and endosporium together 3—5 y. thick, rusty brown in KOH at maturity 
but nearly hyaline earlier, deep red-brown in Melzer’s solution, smooth at 
first but soon becoming cracked into an areolate pattern and the fissures 
gradually deepening to produce a warty to almost echinulate effect and at 
this time the spore surface appearing distinctly roughened, but peris- 
porial membrane still visible over warts and depressed into the crevices, 
in young stages where the inner thick wall is still hyaline an apical germ 
pore can be observed in some spores, and in abnormal spores a lateral 
beak furnished with a distinct pore is clearly evident, the pore obscured 
in old spores by the cracking up of the thick inner layer of the wall. 


1960] SINGER & SMITH: NEOSECOTIUM 155 


Fics. 1-5. Neosecotium macrosporium: 1, upper part of basidium showing tubu- 
lar sterigmata and young spores, * 450; 2, mature spores in optical section, * 1000; 
3, surface view of nearly mature spores, * 1000; 4, optical section of immature 
spores, X 1000; 5, young basidium, x 450. 


Basidia large, 25-37 & 14-17.5, clavate to subelliptic-pedicellate, 
thin-walled and hyaline or the wall slightly thickened and brownish— 
hence the cell more persistent than usual; sterigmata typically 4 and 
tubular, not often tapering appreciably and the young spore acropetally 
attached or very rarely appearing slightly eccentric. Cystidia, none 
observed. 

Subhymenium of broad intricately interwoven hyphae, hence in section 
appearing somewhat cellular from cut hyphal ends; hyphae of the 
peridium filamentous, many hyphal cells somewhat to markedly inflated 
(4-12-18 ». in diam.), the outer layer more or less radially arranged and 
melleous to dingy ochraceous in KOH, gradually paler toward gleba, not 
at all gelatinous or toward gleba only sub- gelatinous (slightly translucent 
in KOH); clamp connections absent to rarely present. 

The type was collected near Dallas, Texas, by E. P. Ely. The best mate- 
rial we have seen, however, is a collection by R. Sprague, June 13, 1941, 
in grass plots on sandy soil at Mandan, North Dakota (NY). 

This is a most interesting species in many respects: the long, tubular 
sterigmata which often break off leaving the upper half attached to the 
spore as a pedicel or appendage, the tendency of the gleba to become 
powdery at maturity, and the type of spore ornamentation in mature 
spores are all strongly reminiscent of the Lycoperdaceae, so much so, in 
fact, that we are inclined to believe that N. macrosporum actually does 
represent a true link connecting the Secotiaceae to that group. The outer 
surface of the dried peridium is almost Calvatia-like in texture, but this, 
of course, may not have any phylogenetic significance beyond that indi- 
cated by the type and arrangement of the hyphae of the outer zone of the 
peridium. 

The pallid to avellaneous tone of the mature gastrocarp and its texture 
are also reminiscent of Lepiota naucina. Actually, aside from the shape 


156 MADRONO [Vol. 15 


of the spore and the peculiar way in which the inner wall layers break up, 
the spores themselves show resemblances to those of Macrolepiota and 
Leucoagaricus by the presence of a germ pore (though it is obscured at 
maturity) and the strong pseudoamyloid reaction of the thick inner wall. 
Also the spores are metachromatic in Cresyl blue—at least the pale col- 
ored spores are. These characters appear to us to connect Neosecotium 
macrosporum to the Agaricales, family Agaricaceae sensu Singer, and 
very likely in the vicinity of Chlorophyllum and Macrole piota. Hence we 
have here a connecting link, as we see it, between the Lycoperdaceae on 
the one hand and a family of agarics on the other. 

As far as we are aware, this is a heretofore unsuspected connection 
between the two groups, and when viewed in this way it is cause for much 
interesting speculation on the course which evolution has followed. Since 
in this series of papers we are not discussing the direction of evolution, 
we shall limit ourselves to considerations which we believe to be based on 
facts as follows: 

The lycoperdaceous fungi show a wide range of spore color just as does 
the family Agaricaceae, and, though the spores in the Lycoperdaceae are 
small, many show a sufficiently similar type of ornamentation to make it 
imperative that spore structure in that order now be studied by the tech- 
niques in use for the study of spores in the Agaricales. 

The problem of the powdery gleba in the Lycoperdaceae is now no 
problem at all as far as its being an obstacle to ascertaining connections 
to the Agaricales. In a number of species of Agaricus the gills become 
very soft and almost collapse after maturity, and in carpophores which 
did not open but which dried out im situ it is a simple matter to understand 
how these structures could break down to a powdery consistency. The 
presence of a highly developed capillitium is certainly to be regarded as 
an advanced character in the Lycoperdaceae, but this, no matter from 
which source one derives the Lycoperdaceae—the agarics or lower Gas- 
tromycetes—does not offer any serious hurdle to establishing relation- 
ships in either direction. Any hymenophoral trama with thick-walled 
hyphae could easily give rise to “capillitium” if the remaining trama con- 
sisted of thin-walled perishable hyphae. There is no reason why thick- 
walled hyphae should not appear “de nova” in the glebal trama of Gas- 
tromycetes in more than one evolutionary series, since wall-thickenings 
of hyphae are one of the commonest types of hyphal adaptation in the 
fungi as a whole. 

The second species of Neosecotium was found among the collections of 
Arcangelliella in the Zeller Herbarium. A redescription of it follows: 


Neosecotium africanum (Lloyd) comb. nov. Octaviania africana 
Lloyd, Myc. Writings 7:1142. 1922. Octaviania africana Verwoerd, S. 
Afr. Journ. Sci. 22:164. 1925. Arcangeliella africana (Lloyd) Zeller & 
Dodge, Ann. Mo. Bot. 22:365. 1935. 

Fructifications spherical, 10-15 mm. thick, drying cinnamon-brown to 


1960] SINGER & SMITH: NEOSECOTIUM 157 


TABLE 1. COMPARATIVE FEATURES INDICATING INTERMEDIATE POSITION OF 
NEOSECOTIUM MACROSPORUM BETWEEN ENDOPTYCHUM ARIZONICUM AND 
SECOTIUM GUEINZII. 


Endo ptychum Neosecotium macrosporum 


arizonicum (Secotium macrosporum ) Secotium gueinzit 
SEPTA without clamp some with, some with clamp 
connections without clamps connections 
VOLVA none none present 
PERIDIAL rough but not smooth tesselate 
SURFACE tessellate 
GLEBA pulverulent pulverulent non-pulverulent 
(strongly) (moderately ) 
SPORES 
ORNAMENT smooth ornamented smooth 
SIZE small large large 
SHAPE subglobose globose ellipsoid 
PORE none present present 


Dresden brown; peridium hard, duplex, the outer layer 140-160 y. thick, 
composed of closely woven slender, hyaline hyphae 1.5—2 y. in diam., the 
inner layer 375-400 vy. thick, composed of hyaline, more loosely woven 
septate hyphae 3—4y in diam., “with lactiferous ducts”.... Zeller & 
Dodge, separable; gleba drying from ferruginous to snuff brown; tramal 
plates 15-30 yu thick; basidia clavate, 23-30 « 7-8 u, sterigmata 10— 
15 long and filiform. 

Spores (giant spores) 17-20 « 14-16y, “normal” spores 13-15 
10-13 y, subglobose to broadly ellipsoid; sterigmal appendage pedicellate; 
dingy yellowish in KOH, dark red-brown in Melzer’s reagent (pseudo- 
amyloid) ; ornamented and thick-walled, inner wall about 2 y. thick, outer 
wall broken up into a pattern of broad obtuse to flattened warts due to 
the cracking of the wall; no germ pore found. 

The description of the spores was taken from the part of the type in 
the Zeller Collections of the New York Botanical Garden. The hymenium 
and tissues of the fruiting body failed to revive sufficiently for critical 
study. It is more than evident to us that because of the pseudoamyloid 
spores with their characteristic ornamentation the species belongs in 
Neosecotium even though in the material available we failed to establish 
the presence of a germ pore. The hard, brown peridium should amply 
distinguish NV. africanum from N. macrosporum. From what we were able 
to ascertain from the limited material available, it appears to us that V. 
africanum is more gastroid than N. macrosporum, in fact may represent 
a, distinct genus at the level of Martellia. Because it represents a different 
level of evolution, it is not included in the chart with the other distinctly 
secotiaceous species. 

Naturally, the genera Endoptychum, Neosecotium, and Secotium form 
a definitely circumscribed and sharply outlined group—a tribus or sub- 


158 MADRONO [Vol. 15 


family—which may also contain such genera as Polyplocium Berk., Gyro- 
phrangmium Mont., and Longula Zeller. 

Since we do not wish to enter such intricate questions of purely gastro- 
mycete taxonomy as the possibility of maintaining all three last-named 
genera (which seem to us extremely close to each other), and since our 
experience with them is relatively limited, we prefer to omit these genera 
for the time being. However, their close relationship to Secotium sensu 
stricto as well as Endoptychum cannot be overlooked. 


REVIEW 


Comparative Morphology of Vascular Plants. By ADRIANCE S. FOSTER, and ERNEST 
M. Grirrorp, JR. 555 pp., 213 figs. W. H. Freeman, San Francisco. 1959. $9.00. 

The literature of vascular plant morphology has been greatly enriched by this 
new textbook by two prominent teachers and researchers at the University of Cali- 
fornia at Berkeley and Davis. In contrast with other morphology texts that have 
appeared in recent years, this is a product of men who have devoted their entire 
careers to the higher plants. As a result, the book is organized in a manner that em- 
phasizes morphological problems of current interest in this area, with subdued treat- 
ment of the burning questions of morphology of the early years of this century that 
are currently only of historical interest. This book is likely to enjoy a long active life 
as a textbook and reference work, therefore a detailed review seems justified. 

A unique feature that sets apart “Comparative Morphology” from earlier text- 
books is the organization of material into two sections. In the first part, consisting 
of six chapters, the principal characteristics of the vascular plants are surveyed in a 
comparative fashion; in the second part individual chapters are devoted to treat- 
ments of the plant groups in systematic sequence. The classification system of Tippo 
is followed throughout. Extinct groups are treated in an integrated manner alongside 
their living relatives, but the emphasis is on modern plant types. Detailed descriptive 
material is not presented for its own sake, but rather as evidence for morphological 
or phylogenetic conclusions. The detail might be described as interpretative and illus- 
trative rather than as encyclopedic. 

The opening chapter tells the beginning student what morphology is all about. 
There is a discussion of the concept of homology, and of the kinds of morphological 
evidence that have proved most useful in reconstructing concepts of phylogeny, such 
as ontogeny, adult form, and the fossil record. The frontiers of modern experimental 
morphology and morphogenesis are described briefly and some pertinent unanswered 
questions are posed. The following chapter deals with the overall characteristics of 
the phylum of vascular plants, giving an outline of a typical life cycle involving an 
alternation of generations. The existence of apospory and apogamy and the signifi- 
cance of these phenomena on the classical theories regarding the origin of alternate 
generations is discussed. The phylum is then divided into the usual four subphyla of 
Eames and Tippo. 

Four chapters dealing with the principal areas of morphological investigation are 
devoted to the vegetative sporophyte, the sporangia, the gametangia, and to embry- 
ogeny. Under the heading of vegetative sporophyte are included discussions of the 
general structure of shoot and root, types of branching, types of leaves, microphylls 
versus megaphylls, and the phylogenetic origin of leaves according to Bower. The 
Telome Theory is presented briefly. The area of plant anatomy is entered with a dis- 
cussion of the problems of classification of tissues and tissue systems. The system of 
Sachs is presented, and the structure and development of the principal tissues are 


12960] REVIEW 159 


described. The chapter closes with a résumé of the historically important Stelar 
Theory. In this section the uses of the terms dictyostele and eustele are clarified. 

The chapter on sporangia describes their function, position, and the organization 
of sporophylls into strobili in some groups. The structure and development of the 
two types of sporangia, the eusporangium and the leptosporangium are described, with 
an excellent series of comparative developmental drawings. The phylogenetic sig- 
nificance of the presence of two sporangium types is discussed. The following chapter 
on gametangia contrasts antheridia with archegonia in development, structure, and 
position. The concluding chapter in the first section of the book deals with embry- 
ogeny; the parts of embryos, polarity, and the development of the embryo from the 
zygote. The phylogenetic aspects of the study of embryo development are discussed. 

The second section of the book opens with three chapters dealing with the sub- 
phyla usually known as the lower vascular plants. The Psilopsida are introduced 
with an historical treatment of their discovery, followed by synopsis of their classi- 
fication into two orders and three families. Rhynia, Horneophyton, and Asteroxylon 
are described. Treatment of these fossils is limited to their general organography and 
anatomy. Psilotum and Tmesipteris are covered in much greater detail. Included in 
the description of the sporophyte structure is a discussion of the interpretation of the 
nature of the stem appendages, and of the multilocular sporangia found in these gen- 
era. The gametophyte generation is discussed in greater detail than is usual in recent 
texts, incorporating the results of contemporary workers like Bierhorst. The brief 
section on embryo development is followed by a concluding summary for the group. 
In the presentation of the details of structure and development of the sporangia, 
gametangia, and embryo, the earlier introductory chapters on these organs serve as a 
basis for comparison. In the usual treatment which lacks such introductory chapters, 
the organs of Psilotum must be studied by the beginning student, detached from the 
reality of the plant world that he knows. 

The Lycopsida are treated in similar fashion, but here the living genera Lycopo- 
dium and Phylloglossum are described first, followed by the extinct Protolepidoden- 
dron and Baragwanathia. Then follow Selaginella, the Lepidodendrales, /soetes, and 
the Pleuromeiales. Throughout this chapter much recent research is presented, along 
with the necessary details of structure and development that are part of the usual 
subject matter. The chapter on the Sphenopsida follows, using the same pattern of 
presentation. Equisetum is described first in detail, followed by a brief statement on 
Hyenia. Sphenophyllum and Calamites conclude the subphylum. 

The Pteropsida include the vast majority of living vascular plants and are de- 
scribed under a series of ten chapter headings. The first is a brief introductory de- 
scription of the group, followed by another chapter which introduces the Filicinae. 
This chapter includes a very brief discussion of fossil fern foliage in general, and of 
the Coenopteridales in particular. The taxonomic summary for the ferns is included 
here, followed by a list of critical areas of morphological study compiled by Bower. 
The Eusporangiate Ferns and Leptosporangiate Ferns are treated next under separate 
headings. The latter group, which includes most living fern genera, is treated as a 
whole in great detail. Excellent series of drawings illustrate the degrees of compound- 
ing of fern fronds, variations in venation patterns, variation in sorus structure and 
position, the development of the sporangium, sporangium structure and various types 
of annuli, types of sorus in regard to order of maturation of the sporangia. The vast 
array of stele types found in the ferns is illustrated by an excellent series of photo- 
micrographs. The gametophytes and embryos are described in the same manner as in 
the lower groups. A section dealing with special problems in fern morphology dis- 
cusses “phyletic slide’ and the relationship between sorus position and phylogeny. 
Recent work in experimental morphology of the ferns by Wardlaw and others is care- 
fully reviewed. A brief résumé of the problems of fern systematics is included, illus- 
trating the relationship between morphology and phylogeny, between phylogeny and 
classification. In the course of the systematic treatment, the principal families are 
briefly described, with fuller treatment of the Marsileaceae. 

The Gymnosperms are covered by four chapters. The first is an introductory 


160 MADRONO [Vol. 15 


conspectus of the group which includes brief treatments of the extinct orders Cycado- 
filicales, Cordaitales, and Bennettitales. The development and structure of the seed 
are covered here, including the details of ovule ontogeny, megasporogenesis, the for- 
mation of the megagametophyte, pollination, and fertilization. This discussion is fol- 
lowed by a brief statement on embryogeny and seed maturation. The second chapter 
in this series is devoted to the living cycads and Ginkgo. Megasporophyll evolution is 
illustrated by drawings of various cycads; the cycad life cycle by another series. The 
details of ovule development and of embryogeny not found in the introductory chap- 
ter are included here. The Coniferales occupy the third chapter and are introduced 
by a systematic treatment of the principal families. This is followed by the usual 
section on organography and anatomy. Florin’s work on Paleozoic and Mesozoic 
conifers is reported in connection with leaf and strobilus evolution and structure. The 
life.cycle of modern conifers is illustrated by Pinus. Included here are the details of 
fertilization, embryogeny and seed development. Then other conifers are compared 
with Pinus. The final chapter on the Gymnosperms deals with the Gnetales. The 
structure and life cycle of Ephedra are presented in detail, followed by a brief 
statement of the differences between Ephedra and the other genera, Gnuetum and 
Welwitschia. 

The final section of the book consists of two chapters on the Angiosperms. The 
first of these chapters treats the general structure and evolution of the group, while 
the second is devoted to the reproductive cycle. Under general structure, leaf mor- 
phology is described in detail, with series of illustrations of venation patterns. Stem 
and root structure are covered more briefly, but a concise statement of modern views 
on nodal anatomy and its phylogenetic significance is included, as is a brief statement 
on wood anatomy. The major part of the chapter is devoted to the problems of floral 
morphology, including theories of the nature of the flower, and the impact of evidence 
from floral vascular anatomy and from floral ontogeny on these theories. The vast 
body of work on primitive woody Ranales by Bailey and his associates during the 
past twenty years is drawn upon for evidence on phylogeny of stamens and carpels. 
The last chapter on angiosperm reproduction describes microsporangium development 
and microsporogenesis, the development of the male gametophyte, the ovule, mega- 
sporocyte, megasporogenesis, and embryo sac, with detailed discussion of the im- 
portant types of the latter. The events of fertilization, endosperm development, and 
embryogeny follow, with a final discussion of seeds and seedlings. 

In summary, Foster and Gifford’s ‘““Comparative Morphology of Vascular Plants” 
is an excellent work featuring clear discussions and illustrations, with an organization 
that should prove a boon to morphology teaching—SAnrorp S. TEPFER, University 
of Oregon, Eugene. 


NOTES AND NEWS 


From June through late December, 1959, with the aid of a National Science 
Foundation grant, Dr. Fritz EHRENDORFER intensified his field and laboratory studies, 
started several years ago, on the genus Galium in the western United States. He re- 
turned to Vienna to take up his new duties as Assistant Curator of the Naturhis- 
torisches Museum. 


PROFESSOR HERBERT L. MAson, who recently was the recipient of a Fulbright 
award, will be taking a sabbatical year from the University of California commenc- 
ing February 1. He will be in residence at the University of Auckland, New Zealand, 
devoting his time mainly to studies of floristic relations in the Southern Hemisphere. 


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 lost 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. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume I, 1916-1929... 3)... 2) = es 55:00 
Volume IT, 1930-1934 . . . .. . 5.00 
Volume ITI, 1935-1936 . . . . . . 5.00 
Volume IV; 1937=1938 15. 2 ae Seno 
Volume V, 1939-1940), a7) ee eo 00 
Volume VI, 1941-1942 . .... . 5.00 
Volume VII, 1943-1944 . . . . . . 5.00 
Volume VIII, 1945-1946 .... . 5.00 
Volume IX, 1947-1948 . . .. . . 6.00 
Volume X, 1949-1950 i>. 2 8 2a) 7 7200 
Volume XI, 1951-1952 . . . .. . 8.00 
Volume XII, 1953-1954. . . . . . 8.00 
Volume X1IT, 1955-1956. Ae S06 
Volume XIV, 1957-1958 ... . . 8.00 
Single numbers, 

Volumessl. and)2> 27 ).) 3) aie .50 
Succeeding numbers, 

(except Volume 2a Non 2) = 094 i ae O0 

Volume XI, Nov2°> 1.40. ete 1 esO) 


The subscription price of MaproNo is $6.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


Bot. 


MADRONO 


VOLUME 15, NUMBER 6 APRIL, 1960 


Contents 


PAGE 
THE REPRODUCTIVE STRUCTURES OF FRAXINUS 
VELUTINA (OLEACEAE), Herbert F. Copeland 161 


A NEw SILENE FROM NORTHWESTERN CALIFORNIA, 
A. R. Kruckeberg 172 


FREED HoFrrMaNn, John L. Morrison 178 


CLEARED CARDIOCARPON LATE-ALATUM LESQ., 
CORDAITEAN SEEDS FROM Micuican, J. F. Davidson 180 


THE BAsic CHROMOSOME NUMBER OF THE GENUS 
NEPTUNIA (LEGUMINOSAE-MIMOSOIDEAE), 


B. L. Turner and O. S. Fearing 184 
HYBRIDIZATION AND INSTABILITY OF YUCCA, 

John Milton Webber 187 
Review: The Physiology of Forest Trees, edited by 

Kenneth V. Thimann, et al. (N. T. Mirov) 192 
NoTEs AND NEws 192 


A WEST AMERICAN JOURNAL OF BOTANY 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription 
price $6.00 per year. Published quarterly and issued from the office of Madrofio, 
Herbarium, Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


HERBERT L. Mason, University of California, Berkeley, Chairman 
EpcAR ANDERSON, Missouri Botanical Garden, St. Louis 
LymMAN BENSON, Pomona College, Claremont,. California. 
HERBERT F. COPELAND, Sacramento College, Sacramento, California. 
Joun F. Davipson, University of Nebraska, Lincoln. 

IvAN M. JoHNsToNn, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MitprepD E. MATuias, University of California, Los Angeles 24. 
MARION OWNBEY, State College of Washington, Pullman. 

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: Herbert G. Baker, Department of Botany, University of California, 
Berkeley, California. First Vice-president: Malcolm A. Nobs, Carnegie Institution of 
Washington, Stanford, California. Second Vice-president: Herbert F. Copeland, Sac- 
ramento City College, Sacramento, California. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley, California. Corre- 
sponding Secretary: Francia Chisaki, Department of Botany, University of California, 
Berkeley, California. Treasurer: John Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1960] COPELAND: FRAXINUS VELUTINA 161 


THE REPRODUCTIVE STRUCTURES OF FRAXINUS 
VELUTINA (OLEACEAE) 


HERBERT F. COPELAND 


The observations on floral structure and embryogeny in Fraxinus velu- 
tina Torrey, called the Arizona ash, which are here presented, were under- 
taken because an abundance of material was available; and because the 
facts as to Oleaceae assembled in Schnarf’s (1931) Vergleichende Embry- 
ologie der Angiospermen were fragmentary, including no reasonably com- 
plete account of any single species. 

Material was collected from trees cultivated on the grounds of Sacra- 
mento City College during the years 1957 to 1959. It was treated by 
routine microtechnical methods: fixed in Bouin’s fluid and stained with 
Delafield’s haematoxylin, Heidenhain’s haematoxylin, or safranine and 
light green. 

In the library, consulted when a certain acquaintance with the plant had 
been attained, I found out that the microscopic reproductive features of 
the Oleaceae are no longer poorly known; also, that there has been dis- 
agreement as to the proper place of Oleaceae in the taxonomic system. 
These matters are discussed in later sections of this paper. 


THE TREE 


Fraxinus velutina occurs near springs and along streams in western 
Texas, New Mexico, Arizona, and southern California. Pratt (1922?) 
wrote of it as apparently new in cultivation and recommended it as resis- 
tant to alkali and drought. It is widely planted in northern California, 
where it is seen to survive with little or no irrigation, but to flourish in 
watered lawns. 

Munz and Laudermilk (1949) refer all plants of this species which are 
native in California to var. coriacea (Watson) Rehder, and I have been 
uncertain of the identity of the cultivated material. Taylor (1945) found 
the species diploid (2n = 46) and the variety tetraploid (2n =92). The 
cultivated trees are diploid (n = 23), and are to be referred to the species. 

As a typical ash, this is a deciduous tree bearing opposite pinnate leaves 
and producing samaras. It belongs to the group of ashes in which the 
flowers are apetalous and dioecious. Flower clusters appear in the axils 
of the proximal fallen leaves of the previous year during the month of 
February. Pollination is evidently by wind. In March, the staminate 
flowers fall, and the leaves begin to unfold. The samaras grow to their full 
size by the end of May, but the seeds are not mature until autumn. The 
samaras are shed, along with the leaves, in autumn storms. 

The flower clusters are dense glomerules. After anthesis, the axes of the 
pistillate clusters become elongate, and it is seen that they are freely 
branched in a decussate pattern. All of the flowers or fruits of a particular 


Maprono 15, No. 6, pp. 161-192, April 18, 1960. 


162 MADRONO [ Vol. 15 


cluster are at a particular time in nearly the same stage of development; 
most axes bear terminal flowers; hence, the clusters are to be construed 
as thyrses. 

MALE STRUCTURES 

The staminate flower (fig. 1) consists, beyond its receptacle, of a calyx 
which is reduced to a minute toothed cup and of two stamens having brief 
filaments and prominent basifixed extrorse anthers. The vascular supply 
of this flower (fig. 2) consists of a cylinder of tissue originating from the 
two sides of a bract gap, emitting a cycle of a small indefinite number of 
feebly developed traces to the calyx, and then splitting into two bundles 
which ascend the connectives of the anthers to their summits. 

The anthers are of the structure usual in flowering plants. The cells 
of the endothecium duly develop ribbed walls, and the dehiscence of the 
anthers, which occurs through the usual two lengthwise clefts, is produced 
by their contraction. The tapetum is of the secretion type. The nuclei of 
the tapetal cells divide more than once, and then undergo fusions, with the 
result that just before the tapetum is absorbed its cells contain varying 
numbers of large nuclei with varying numbers of nucleoli (figs. 3, 4). 

The haploid chromosome number, observed during meiosis in the pollen 
mother cells, is 23 (figs. 5,6, 7). The pollen grains are separated by simul- 
taneous furrowing. When mature, they are four-grooved, having the 
surfaces between the grooves finely pitted, and contain a tube nucleus 
and a generative cell (fig. 8). 


THE PISTILLATE FLOWER 


The pistillate flower (fig. 9) consists, beyond its receptacle, of a cup- 
shaped calyx with a dentate margin and a compound pistil of two carpels. 
The ovary contains two locules. The septum between the locules is punc- 
tured by a small cleft near its upper end: the upper ends of the locules 
are continuous. Each locule contains two ovules which are pendant from 
the distal area of the septum. The ovary is flattened contrary to the nar- 
row septum (fig. 13); the flattening is moderate through most of the 
height of the ovary, but is greater in the upper part. The brief style is 
cylindrical. The stigma is of two lobes which are pressed together when 
the flowers are first exposed but become separate at anthesis. The stig- 
matic surface is papillose. 

The ovary bears a moderate number of peltate trichomes (figs. 15, 16) 
which are of the same nature as those which occur on leaves of Syringa 
and Ligustrum. The pedicel and flower bear also a few simple hairs, mostly 
on the margin of the calyx. 

The vascular system supplying this flower is as follows (figs. 11, 12). 
The usual cylinder of vascular tissue ascends the pedicel. The calyx con- 
tains a whorl of a varying number of feebly developed bundles which 
fade out below. This means that the stele in the pedicel supplies only the 
pistil. The stele gives rise to an outer whorl of about fourteen bundles 
including (a) two well-marked carpel-dorsals, respectively ascending the 


1960] COPELAND: FRAXINUS VELUTINA 163 


Fics. 1-21. Fraxinus velutina: 1, staminate flower <8; 2, vascular supply of 
two staminate flowers, « 40; 3, 4, mature cells of the tapetum, 720; 5, 6, pollen 
mother cells with nucleus in heterotypic metaphase, * 720; 7, pollen mother cell 
with nucleus in heterotypic anaphase, xX 720; 8, pollen grain, * 720; 9, pistillate 
flower, X 8; 10, pistil, 8; 11, pistillate flower cleared in chlorine water, * 20; 12, 
model of vascular system in lower part of the pistillate flower, & 40; 13, cross section 
of young ovary, X 40; 14, longitudinal section of young overy, * 40; 15, 16, radial 
section and surface view of scale of ovary, X 320; 17, archesporial cell of young ovule, 
xX 320; 18,19, longitudinal section of developing ovule, & 40, and nucellus of same 
showing megaspore mother nucleus in heterotypic metaphase, * 320; 20, 21, longi- 
tudinal section of ovule, 40, and nucellus of same showing megaspore mother cell, 
< 320. ca, sepal bundles; cd, carpel dorsal bundles; cl, carpel lateral bundles; cv, 
carpel ventral bundles; ovw, ovary wall bundles. 


margins of the ovary and continuing up the style into the stigmatic lobes; 
(b) a total of about eight ovary wall bundles, being about two on each 


164 MADRONO [Vol. 15 


side of each carpel dorsal; and (c) a pair of well-marked carpel-laterals 
at each margin of the septum. Above the level of the locules, the ovary 
wall bundles and carpel laterals spread apart to form two fan-like layers 
toward the respective surfaces of the flattened upper part of the ovary. 
These bundles do not enter the style, but fade out. The vascular tissue 
which ascends beyond the departure of the whorl just described takes the 
form of an attenuate cone ascending the septum of the ovary. The cone 
splits to form two bundles located toward the margins of the septum, and 
each of these, toward the summit of the septum, splits in turn into two 
bundles which diverge and turn down to supply two ovules lying in dif- 
ferent locules. 
OVULE AND EMBRYO SAC 

When the pistillate flowers are first exposed, before the stigmatic lobes 
swing apart and become receptive, one finds in each locule two immature 
ovules (figs. 13, 14) of the form of downward-pointing fingers. Each one 
contains a strand of immature vascular tissue. Each contains one hypo- 
dermal archesporial cell (fig. 17). The archesporial cells are located on 
the sides of the ovules which are away from the plane of the carpel-dorsal 
bundles: one sees them best in sections cut parallel to the septum. 

The archesporial cell is itself the megaspore mother cell. It becomes 
elongate, and the epidermis covering it is pushed up as a scanty nucellus 
(figs. 19, 21). The tissue on all sides of the nucellus grows forth to form 
an integument. The growth is greatest on the side of the nucellus toward 
the original tip of the ovule, which now becomes the chalaza. The effect of 
this growth is to turn the nucellus toward the summit of the ovary, and to 
enclose it except for a narrow micropyle leading up from it (the growth of 
the ovule is illustrated only by two little diagrams, figs. 18, 20). The 
mature ovule is somewhat flattened between the septum and the ovary 
wall. 

During the growth of the ovule as just described, spiral tracheids 
appear in the main bundle which runs down the raphe to the chalaza. At 
the same time, several additional bundles begin to undergo differentiation 
in the integument. These latter bundles, few but not of definite number, 
usually three or four, extend the length of the ovule from the end of the 
original bundle, in the chalaza, nearly to the level of the micropyle. 

While the integument is growing up about the nucellus (fig. 18), the 
meiotic divisions of the nucleus of the megaspore mother cell, and the 
accompanying cell divisions, begin to take place (figs. 19, 22-24). A T- 
shaped tetrad of megaspores is produced. The spore at the chalazal end 
is functional. 

The nucleus of the functional megaspore undergoes three successive 
divisions, while the three non-functional spores and the nucellus are 
absorbed (figs. 25, 26; the stage with eight free nuclei has not been seen). 
An embryo sac with an egg and two synergids, two polar nuclei and three 
antipodal cells, is organized (fig. 27). The antipodal cells appear mori- 
bund from the time when they are set apart, and soon disappear. During 


1960] COPELAND: FRAXINUS VELUTINA 165 


My | 
£127 Vy 


sau) 


qn: 


NO iy 
SZ 


C1 “= 


Fics. 22-37. Fraxinus velutina: 22, nucellus showing diad cells, * 320; 23, homeo- 
typic metaphase, * 320; 24, tetrad of megaspores, X 320; 25, 2-nucleate embryo sac, 
X 320; 26, 4-nucleate embryo sac, X320; 27, mature embryo sac, & 320; 28, fertiliza- 
tion, 320; 29, first division of endosperm nucleus, & 320; 30, 31, zygote and endo- 
sperm in 4-celled stage, *320; 32, 33, zygote about one month after fertilization, 
320; 34, first division of zygote, & 320; 35, four-celled embryo in many-celled endo- 
sperm, X 320; 36, dissection of lower part of fruit about two months after fertiliza- 
tion, X 20; 37, longitudinal section of seed at same stage as in fig. 36, X 80. h, 
hypostase. 


166 MADRONO [ Vol. 15 


the development of the embryo sac, the inner epidermis of the integument 
takes on the character of a jacket layer. 

The meiotic divisions appear always to take place earlier in one of the 
four ovules of the ovary than in the others (figs. 18, 19 show the beginning 
of meiosis before the integument is fully developed; figs. 20, 21 show a 
fully formed ovule in which meiosis has not begun). Through all later 
stages, one ovule is always found in a more advanced stage of development 
than the others. More ovules than one may develop complete embryo 
sacs; pollen tubes may enter more than one ovule; but only the ovule 
which developed most rapidly is capable of maintaining an embryo. 
Developed embryo sacs in other ovules undergo degeneration by a process 
of collapse which begins at the chalazal end. 

When the embryo sac is ready for fertilization, the ovule contains a 
hypostase, consisting of a small body of differentiated chalazal cells. It is 
recognizable by cell walls which are relatively retentive of the dye safra- 
nine; in sections stained with haematoxylin alone, it is not recognizable. 
It is separated from the chalazal end of the embryo sac by a few undif- 
ferentiated cells. 


FERTILIZATION, ENDOSPERM, AND EMBRYO 


Pollen tubes have been seen in the micropyles of various ovules. Their 
discharged tips, of the form of heavily staining masses, have been seen 
between pairs of synergids in which the nuclei remain recognizable: it 
appears that both synergids survive for some time after the entrance of 
the pollen tube. On several slides, one sperm nucleus has been seen near 
the egg nucleus, while the other sperm nucleus is in process of fusing with 
the polar nuclei (fig. 28). 

The endosperm is cellular from its origin. The first division of its 
nucleus (fig. 29) is followed by deposition of a transverse wall. The divi- 
sions of its daughter nuclei are followed by the deposition of walls which 
are transverse or nearly so: the endosperm passes through a stage in 
which it is a linear (or nearly linear) tetrad of cells (figs. 30, 31). During 
further multiplication, the cells of the endosperm do not become differ- 
entiated except for the crushing of some of them at the micropylar end; 
no haustoria are produced. The chalazal end of the endosperm digests or 
crushes the cells of the chalaza as far as the hypostase. For the rest, the 
endosperm does not digest the adjacent cells. It appears to press upon the 
jacket layer and the hypostase, and the integument grows along with the 
endosperm. 

Fertilization having taken place early in March, the zygote remains 
undivided until about the beginning of May, by which time the endo- 
sperm is already many-celled. Before the zygote divides, it may become 
enlarged (figs. 32,33). After it divides, the enlargement disappears; there 
is no exceptionally swollen cell at the base of the suspensor (figs. 34, 35). 
Dividing transversely, the zygote and its progeny produce a uniseriate 
filament of a dozen or more cells (figs. 37-39). The definitive embryo 


1960] COPELAND: FRAXINUS VELUTINA 167 


ARISES SC 8 eae 


“NE GENT: 


2 
4 
4 
3 


‘ 


Fics. 38-44. Fraxinus velutina: 38, 39, embryos from same collection as figs. 36 
and 37, X 320; 40, mature fruit, * 4; 41, dissection of lower part of mature fruit, 
x 20; 42, cross section of lower part of mature fruit, « 20; 43, dissection of seed 
(much longer than the one in fig. 41), & 8; 44, section of outer part of seed, seed 
coat to the left and endosperm to the right, * 320. 


originates by longitudinal divisions in several cells at the distal end of this 
filament. The proximal cells lose their stainable contents, shrink, and 


disappear. 


168 MADRONO [Vol. 15 


FRUIT AND SEED 

Between early March and late May, growth of the ovary produces a 
samara of mature size. There are great differences in rate of growth 
between different parts of the ovary and different dimensions of the parts. 
The lower part of the ovary, originally a small moderately flattened cyl- 
inder, retains this shape while growing to seven or eight times its original 
dimensions. The upper part, while undergoing slight increase in thickness, 
grows to some fifteen times its original width and fifty times its original 
length. Thus it produces the wing of the samara. The wing is not derived 
from the style, which persists, if at all, as a withered terminal scrap. 

The internal septum of the ovary, originally a wall with nearly plane 
surfaces, undergoes swelling immediately after fertilization and becomes 
fusiform. The bundles of the ovary wall become greatly enlarged by the 
differentiation of masses of fibers and form vertical ribs on the inner 
surface. Septum and ribs fill the locules, leaving scant clefts of compli- 
cated form. 

Of the four ovules which are hung from the upper part of the septum, 
three undergo no growth, but turn dark and shrivel. The fourth develops 
at its proximal end a long funiculus which holds the main body of the 
developing seed at about the middle of the height of the septum (fig. 36). 
During the growth of the funiculus, its surface is thrown into microscopic 
transverse ridges. 

The seed proper, enlarging principally after the beginning of July, 
reaches dimensions approximately half of those of the lower part of the 
fruit, that is, of the fruit apart from the wing (fig. 41). In the course of 
this growth, the seed presses into, and largely crushes, the enlarged sep- 
tum. It pushes back the funiculus, throwing it into coils. The surface of the 
mature seed is yellow to brown, shiny, and minutely papillate. 

Dissection shows the seed to have a thin coat covering an endosperm 
in which lies a large cylindrical embryo divided through the distal half 
of its length into two cotyledons (figs. 42, 43). The papillae on the surface 
of the seed are enlarged epidermal cells. The jacket layer, that is, the 
internal epidermis of the integument, remains intact. With the exception 
of the papillae, the jacket layer, and small bodies of fibers in the four or 
five longitudinal bundles, the cells of the integument are compressed and 
nearly empty. A definite continuous wall at the outer margin of the endo- 
sperm belongs to the endosperm, not to the jacket layer. The cells of the 
endosperm are packed with granules (fig. 44). These are definitely not of 
starch; they appear to be of protein. 


DISCUSSION 


Schnarf’s account of the embryology of Oleaceae consisted of scattered 
observations upon Forsythia, Jasminum, Ligustrum, and Fraxinus in the 
classic general papers of Hofmeister (1858), Warming (1878), Guignard 
(1882), Billings (1901), Juel (1915), and Dahlgren (1923, 1927). Som- 
mer (1929) had studied Fraxinus excelsior among various plants in which 


1960] COPELAND: FRAXINUS VELUTINA 169 


a distinction among the ovules of a single ovary, some continuing their 
development and others undergoing abortion, appears suddenly at a cer- 
tain stage of development. Eames (1931) included Syringa and Forsythia 
among plants in which he studied the vascular supply of the pistil. Subse- 
quent embryological studies include those of Andersson (1931) on a wide 
variety of Oleaceae, and of King (1938) and Messeri (1950) on the 
domestic olive, Olea europaea. Johnson (1941) included Forsythia among 
plants in which he studied the cytology of the male gametophyte. Fotidar 
(1942) studied the floral anatomy of Nyctanthes. Numerous counts of 
chromosomes are reported by Sax (1930), O'Mara (1930), Sax and Abbe 
(1932), Taylor (1945), and Dutt (1952); the contribution of Taylor is 
particularly interesting as including counts for Fraxinus velutina and its 
varieties, and as proposing to limit the subfamily Oleoideae to genera in 
which the basic chromosome number is 23, thus excluding Jasminum and 
Menodora. 

The observations on Fraxinus velutina here presented are in very 
nearly complete agreement with the facts as to Oleaceae in general as 
stated in the literature just cited. Andersson noted in various Oleaceae the 
peculiar tapetum, characteristic of widely scattered presumably derived 
groups, in which the nuclei divide more than once. Johnson found the 
pollen grains of Forsythia binucleate. The vascular supply of the pistil, 
alike in Syringa, Ligustrum, Nyctanthes, and Fraxinus, exhibits slight 
variations upon a common pattern which is precisely that of the typical 
bicarpellate compound pistil according to the theory of Eames. The pat- 
tern of the vascular supply to the ovules is identical in Olea and Fraxinus. 
Billings was presumably mistaken in describing, in the ovule of Fraxinus 
excelsior, a single vascular strand which descends the raphe to the chalaza 
and ascends the integument on the side opposite the raphe. In F. velutina, 
as noted, a varying small number of bundles, in positions which vary from 
one ovule to another, run up from the chalaza; Fotidar observed the same 
structure in Vyctanthes. Also, as Dahlgren suspected, Billings was surely 
mistaken in figuring an ovule in which the megaspore mother cell is 
covered by more than one layer of cells of the nucellus: this appearance 
represents an oblique section of the ovule. In most Oleaceae, the embryo 
sac is of normal type and the definitive embryo develops from several 
distal cells of a filamentous early embryo. In these points, Olea appears 
exceptional: its embryo sac is said to be of Scilla-type and its filamentous 
early embryo is very short. 

The proper location of Oleaceae in the taxonomic system is next to be 
discussed. The traditional place of the family is in an order named Con- 
tortae. The order was established by Linnaeus (1764) to include the 
plants subsequently assembled as families Apocynaceae and Asclepia- 
daceae. Of Olea and its allies, Linnaeus made a separate order Sepiariae; 
he placed Gentiana among primulaceous plants in his order Rotaceae. 
Eichler (1886) and Engler (1892) are responsible for assembling as 
order Contortae the families Oleaceae, Loganiaceae, Gentianaceae, 


170 MADRONO [ Vol. 15 


Apocynaceae, and Asclepiadaceae. In earlier presentations of the Engler- 
ian system, one finds the small tropical family Salvadoraceae placed next 
to Oleaceae; in later presentations it is dismissed from this neighborhood, 
surely correctly, since the ovules of Salvadoraceae have two integuments 
and a nucellus of more than one layer of cells (David, 1938). Wettstein 
(1908) followed Linnaeus and most pre-Englerian authors in placing 
Oleaceae in an order (he called it Ligustrales) separate from Contortae. 
Schnarf followed Wettstein, although expressing doubt that the families 
remaining in Contortae belong together as a natural group. Wettstein 
(1908) is authority for family Menyanthaceae, a segregate from Gen- 
tianaceae, and Schnarf (1931) is authority for family Buddleiaceae, a 
segregate from Loganiaceae. Tournay and Lawalrée (1952) transferred 
Menyanthaceae and Buddleiaceae from Contortae to Ligustrales. 

Schnarf, and Tournay and Lawalrée, were influenced by embryological 
knowledge, including particularly the following point. The endosperm is 
nuclear in proper Loganiaceae and Gentianaceae, and in Apocynaceae and 
Asclepiadaceae. It is cellular in Buddleioideae, Menyanthoideae, and 
Oleaceae. 

Assuming that the production of a nuclear endosperm is a primitive 
character from which the production of the cellular endosperm has 
repeatedly been derived, the presence of both types in a particular order 
or family is not by itself sufficient reason for dividing the group. We can 
interpret Loganioideae, Gentianoideae, Apocynaceae, and Asclepiadaceae 
as a natural series in which the primitive type of endosperm is retained, 
and Buddleioideae and Menyanthoideae as offshoots from it in which 
the derived type of endosperm has developed independently. This appears 
to be the idea of Moore (1947), who considers Loganiaceae to be an 
immediate ally or derivative of some primitive stock from which have 
evolved also the Tubiflorae (among which a repeated evolution of the 
cellular endosperm is evident ) and the Rubiales (which retain the nuclear 
endosperm ). 

The Buddleioideae have a celiular endosperm with haustoria (Moore 
describes these as absent in Polypremnum, but one of his figures shows 
structures to which no other term can be applied) and an embryo devel- 
oped from two cells terminal upon a three-celled suspensor (Soueges, 
1940; Moore, 1948). 

Of Menyanthoideae, the writer has learned nothing beyond what was 
known to Schnarf. A tapetum in which the nuclei divide more than once; 
an embryo sac of normal type, with fugitive antipodal cells; a cellular 
endosperm in which cell divisions beyond the first are transverse, and 
which lacks haustoria; and an early embryo of the form of a many-celled 
filament: all of these are characters in precise agreement with Oleaceae. 
To present knowledge it appears probable that the Oleaceae are derived 
from the Menyanthoideae and should be placed after that group. 


SUMMARY 
1. Fraxinus velutina Torrey, the Arizona ash, a tree of the southwest- 


1960] COPELAND: FRAXINUS VELUTINA 171 


ern United States, is a typical ash of the group having dioecious apetalous 
flowers. The flowers and their vascular systems are described. The vas- 
cular system of the pistillate flower is very nearly that of the typical 
bicarpellate compound pistil according to the theory of Eames. 

2. Staminate flowers consist of little more than two stamens with ribbed 
endothecia and tapeta in which the nuclei divide more than once and then 
undergo random fusions. The haploid chromosome number is 23. Pollen 
grains are 4-grooved, binucleate. 

3. The ovules are unitegmous and tenuinucellate. They have several 
longitudinal bundles in the integument and an obscure hypostase in the 
chalaza. The inner epidermis of the integument becomes a jacket laver. 
Of four ovules in the ovary, only one becomes a seed. 

4. The embryo sac is of normal type, the antipodal cells disappearing 
quickly. 

5. Double fertilization was observed. 

6. The endosperm is of cellular type. The first cell division is by a 
transverse wall; the second cell divisions are by walls which are trans- 
verse or nearly so. No haustoria are produced. 

7. The zygote, after remaining undivided for several weeks, produces 
a filament of many cells. The embryo proper is derived from several 
cells at the distal end of this filament. 

8. The single seed of the samara crowds aside or crushes other struc- 
tures within the ovary and becomes mature in autumn. Papillae upon its 
surface are enlarged epidermal cells. The jacket layer persists to this 
stage. A continuous wall within the jacket layer is the outer cell wall of 
the endosperm. There is a large straight dicotyledonous embryo. 

9. These observations, compared with others in the literature, tend to 
substantiate the naturalness of the family Oleaceae and the order Con- 
tortae. Among other Contortae, the Menyanthoideae appear most similar 
to Oleaceae in embryological characters. 

Sacramento City College 
Sacramento 22, California 


LITERATURE CITED 

ANDERSSON, A. 1931. Studien tiber die Embryologie der Familien Celastraceae, Olea- 
ceae und Apocynaceae. Lunds Univ. Arskr. n. f., Avd. 2, 27, no. 7: 1-110. 

Bitiincs, F. H. 1901. Beitrage zur Kenntniss der Samenentwicklung. Flora 88: 
253-318. 

DAHLGREN, K. V. O. 1923. Notes on the ab initio cellular endosperm. Bot. Notiser 
1923: 1-24. 

———. 1927. Die Morphologie der Nuzellus mit besonderer Beriicksichtigung der 
deckzellosen Typen. Jahrb. wiss. Bot. 67: 347-426. 

Davip, ELIsaABETH. 1938. Embryologische Untersuchungen an Myoporaceen, Salva- 
doraceen, Sapindaceen und Hippocrateaceen. Planta 28: 680-703. 

Dutt, M. 1952. Chromosome numbers in some ornamental jasmines. Science and 
Culture 17:527-528. 

EaMEs, A. J. 1931. The vascular anatomy of the flower with refutation of the theory 
of carpel polymorphism. Am. Jour. Bot. 18:147-188. 

EICHLER, A. W. 1886. Syllabus der Vorlesungen tber specielle und medicinisch- 
pharamaceutische Botanik. 4th ed. Berlin. 


172 MADRONO [Vol. 15 


ENGLER, A. 1892. Syllabus der Vorlesungen uber specielle und medicinisch-pharma- 
ceutische Botanik. Berlin. 

Foripar, A. N. 1942. Floral anatomy of Nyctanthes Arbor-tristis L. Jour. Indian Bot. 
Soc. 21:159-166. 

GUIGNARD, L. 1882. Recherches sur le sac embryonnaire des phanérogames angio- 
spermes. Ann. Sci. Nat. Bot. sér. 6, 13:136-199. 

HOFMEISTER, W. 1858. Neuere Beobachtungen uber Embryobildung der Phanero- 
gamen. Jahrb. wiss. Bot. 1:82-188. 

Jounson, G.W. 1941. Cytological studies of male gamete formation in certain 
angiosperms. Am. Jour. Bot. 28:306-319. 

JureL, H. O. 1915. Untersuchungen uber die Auflosung der Tapetenzellen in den 
Pollensacken der Angiospermen. Jahrb. wiss. Bot. 56:337-364. 

Kinc, J. R. 1938. Morpholegical development of the fruit of the olive. Hilgardia 
11:437-458. 

Linnaeus, C. 1764. Genera plantarum. ... 6th ed. Stockholm. 

MEsseErI, ALBINA. 1950. Alcuni dati sulla embriologia ed embriogenesi di ‘Olea 
europaea” L. Nuov. Giorn. Bot. Italiano 57:149-169. 

Moore, R. J. 1947. Cytotaxonomic studies in the Loganiaceae. I. Chromosome num- 
bers and phylogeny in the Loganiaceae. Am. Jour. Bot. 34:527-538. 

. 1948. Cytotaxonomic studies in the Loganiaceae. II. Embryology of Poly- 
premnum procumbens L. Am. Jour. Bot. 35:404—410. 

Munz, P.A. and J. D. LaupERMILK. 1949. A neglected character in western ashes. 
El] Aliso 2:49-62. 

O’Mara, J. 1930. Chromosome number in the genus Forsythia. Jour. Arnold Arb. 
11:14-15. 

Pratt, M. B. 1922 ? Shade and ornamental trees of California. Sacramento ? 

SAx, K. 1930. Chromosome number and behavior in the genus Syringa. Jour. Arnold 
Arb. 11:7-14. 

and E. C. Appr. 1932. Chromosome numbers and the anatomy of the sec- 

ondary xylem in the Oleaceae. Jour. Arnold Arb. 13:37-48. 

ScHNARF, K. 1931. Vergleichende Embryologie der Angiospermen. Berlin. 

SOMMER, BERTA-SIBYLLE. 1929. Uber Entwicklungshemmungen bei Samenanlagen. 
Flora 124:63-93. 

SouEcEs, R. 1940. Embryogénie des Loganiacées. Développement de l’embryon chez 
le Buddleia variabilis Hemsley. Compt. Rend. 211:139-140. 

Taytor, H. 1945. Cyto-taxonomy and phylogeny of Oleaceae. Brittonia 5:337-367. 

Tournay, R., and A. LAWALREE. 1952. Une classification nouvelle des familles ap- 
partenant aux ordres des Ligustrales et des Contortées. Bull. Soc. Bot. France 
99:262-263. 

von WETTSTEIN, R. 1901-1908. Handbuch der systematischen Botanik. 2 vols. Leipzig 
and Vienna. 


A NEW SILENE FROM NORTHWESTERN CALIFORNIA 
A. R. KRUCKEBERG ! 


Long past is the era in California botany when a collector could count 
among his season’s haul a good proportion of undescribed species. Most 
areas of the state are sufficiently well known so as to limit the likelihood 
of uncovering anything new. Nowadays, range extensions, records of new 
adventives, and the discovery of some inconspicuous annual that fails to 


1 Supported by funds from the State of Washington Initiative No. 171 and by the 
National Science Foundation, Grant G-1323. 


1960] KRUCKEBERG: SILENE L/S 


match any known relative, serve to satisfy the field botanist’s taste for 
novelties. When a hitherto undescribed perennial which is an element of 
a stable plant community is found, it is of more than passing interest. 
Such discovery is most likely in the relatively little explored mountain- 
ous terrain of northwestern California. Szlene marmorensis Kruckeberg, 
described below, is one of these latter day discoveries, having been col- 
lected in 1954 by Dr. C. Leo Hitchcock along the forested slopes of the 
Marble Mountains in southwestern Siskiyou County. 

For the past eight years, I have been maintaining a collection of living 
plants of North American species of Silene. The plants have been grown 
for purposes of observation under uniform conditions, for determining 
chromosome number (Kruckeberg 1953, 1960), and for assessing degree 
of genetic relationships by means of interspecific hybridization (Krucke- 
berg 1954). The genus is represented in California by twenty native 
species (sensu Hitchcock and Maguire 1947), nearly all of which are well 
represented in herbaria. Moreover, most of them are readily distinguish- 
able and thus stand out as clearly defined species. Having worked with 
living plants of all the Californian species as well as most of the known 
species occurring elsewhere in North America, I was genuinely impressed 
with a specimen that did not fall into place with any known species. 

In general habit, Silene marmorensis might be confused with the Sierra 
Nevadan S. verecunda Wats. subsp. platyota (Wats.) H. & M., both hav- 
ing long, flexuous stems and short, lanceolate leaves. In inflorescence and 
floral characters, though, the new species combines features of at least 
three species—S. campanulata Wats., S. lemmoniu Wats., and S. bridgest 
Rohrb.—all of which share, as well, the common characteristics of an 
ovoid, cartilaginous capsule and large black seeds; in addition, all four 
species are tetraploid (2n — 48). The new species appears most closely 
related to S. bridgesi of the yellow pine belt in the Sierra Nevada, owing 
to the close correspondence of the two species in inflorescence and flowers. 
However, in S. marmorensis, the flowers are not pendant, nor do the 
proportions of its calyx, petals, and style match those of S. bridgesit. 

The suite of characters which defines S. marmorensis can be sum- 
marized as follows. The several wiry, weakly ascending stems bear 5—7 
pairs of uniformly short, lanceolate leaves; the open, lax, glandular 
inflorescence bears 4—6 pairs of lateral cymules, with the cymules con- 
sisting of single flowers in wild plants and the lateral flowers apparently 
abortive; the flowering calyces are elongate, ovate-lanceolate, and are 
borne divaricately on thin, wiry pedicels. The pinkish petals have a 
simple, bifid blade at the base of which are the two laterally divaricate, 
erose auricles; the appendages are short and rather broad. No one of 
these features is specific for a western Silene, but in ensemble, they con- 
trive to give a picture of a rather delicate, unassuming grace and unique- 
ness to the plant (figs. 1-3). 

Having examined only two collections of S. marmorensis, it would be 
rash to attempt a delimitation of its distribution and habitat preference. 


174 MADRONO [Vol. 15 


The two specimens cited below were collected along the steep, winding 
forest road leading northeast up to Camp Three from the confluence of 
the Salmon and Klamath rivers at Somes Bar. The terrain is steep, 
forested mountainside with a south to southwest exposure. A topotype 
collection (Kruckeberg 4023) grew in loose talus of gabbroic rock in a 
fairly open stand of Douglas fir, black oak, and madrone. A list of asso- 
ciated species is appended to the species description. 

Silene marmorensis sp. nov.” Planta perennis tenui radice; caudice 
caulibus compluribus tenuibus, 2.5-4.0 dm. longis, puberulentis, supra 
glandulosis; foliis caulinis 5—7 paribus, fere eadem magnitudine, lanceo- 
latis, 3.0-4.5 cm. longis, 0.3—0.5 cm. latis, scabrido-pubescentibus; brac- 
tis reductis, lineari-lanceolatis, glandulosis; inflorescentibus terminalibus, 
10-20 cm. longis, fere simplicibus, cymulis 5—7 iugis, pedicellis 7-10 mm. 
longis, filiformibus, glandulosis; calyce 13 mm. longo, anguste elongato- 
ovato, glandulosc, minus valide 10-nervo, lobis late lanceolatis, 3 mm. 
longis; calyce in fructu campanulato; corolla supra ex rubro pallea, infra 
subviridilurido (galbino), ungue 8-10 mm. longo, glabrato, sursum 
latiore, auriculo parvo eroso in utroque summae latere, lamina 4-6 mm. 
longa oblongata, alte bilobata, lobis integris vel in apice tenuiter erosis, 
appendicibus 2, oblongatis; staminibus exsertis, in tubo compressis, 
filamentis 11-13 mm. longis, glabratis, polline subfusco; stipitibus 3—4 
mm. longis, puberulentis; stylis 3, 10-12 mm. longis, filiformibus, papillis 
stigmatum paucis et tantummodo in apice styli; ovariis glabratis ovatis 
maturitate et ligno-cartilagineis et 5-dentatis; seminibus 2.5 mm. longis, 
nigris, fulgentibus, tuberculis ex ordinibus brevibus conicis. 

Perennial, from a long slender taproot, the multicipital crown bearing 
several underground, erect branches, each of which terminates above 
ground in a slender stem, 2.5—4.0 dm. long, stems simple, purplish and 
eglandular-pubescent at base, retrorsely glandular-pubescent above, espe- 
cially on branches of inflorescence; the 5—7 pairs of cauline leaves similar 
in size and shape, reduced only in the inflorescence, lanceolate, 3.0—4.5 
cm. long, 0.3—0.5 cm. wide, sparsely scabrid-pubescent on both surfaces, 
the leaf-like bracts of inflorescence progressively reduced upwards, short- 
lanceolate, glandular; inflorescence terminal, 10—20 cm. long, simple, or 
with 1-2 branches, each bearing 5—7 pairs of cymules (the lateral flowers 
of each cymule apparently abortive in field material) ; pedicels 7-10 mm. 
long, filiform, glandular; calyx in flower indistinctly 10-nerved, glandu- 
lar, narrowly elongate-ovate, slightly constricted at base, about 13 mm. 
long, becoming campanulate through distension by the maturing ovary, 
the teeth ovate-lanceolate, short-acuminate, 3 mm. long, somewhat mem- 
branous in the sinuses, margins of teeth densely long-ciliate; corolla pale 
pink above, greenish yellow beneath, the claw 8-10 mm. long, glabrous 
throughout, slender at base widening above, with a small erose, angular 
auricle at either side of summit, the blade 4-6 mm. long, oblong, bilobed 


*The Latin diagnosis kindly prepared by W. M. Read, Professor of Classics, 
University of Washington. 


1960] KRUCKEBERG: SILENE 175 


SLUBSE BECHOrecnl« Prnexrmweren ts 


" TYPE 


4 3AYER 3S 


FLORA OF CALIF ORHIA 


QILENE VEREQUBDA SSK. PLATYOTA 
{Se wathe} H. & Ne 


SISKIYOU CD: | WILE BORTH OF SOMES GAR 
OH ROAD TH Cewr THOLE » 
PLANTS BITH TAPROOT, PETALS Finn < 


By Le HiTeocK B25 PURE Be, I9Sa 


Fic. 1. Type specimen of Silene marmorensis, Hitchcock 20221 (WTU 179156), xX %4. 


over one-half its length, the lobes oblong, entire to slightly erose at tip, 
appendages two, broadly oblong, truncate, the free margins entire; 
stamens slightly exserted, crowded at throat, the filaments 11-13 mm. 
long, glabrous throughout; pollen tawny brown in color; carpophore 3—4 


176 MADRONO [ Vol. 15 


Fic. 2. Silene marmorensis: a, plant grown in greenhouse (photo by W. Martin, 
Still Photo Unit, University of Washington) ; b, seed, X ca. 20. 


mm. long, retrorsely puberulent; styles three, 10-12 mm. long, filiform, 
nearly straight, the stigmatic papillae few and congested at tip; ovary 
glabrous, at maturity ovoid with walls woody-cartilaginous, opening with 
five teeth; seeds about 2.5 mm. long, black, shiny, with concentrically 
longitudinal rows of short conical tubercles. 2n = 48. Figs. 1-3. 

Type. Siskiyou County, California: 1.0 mile north of Somes Bar on 
road to Camp Three, June 22, 1954, C. L. Hitchcock 20221 (WTU 
179156); another specimen (topotype) from 5.5 miles above Somes Bar 
on road to Camp Three, growing in loose talus of gabbroic rock, A. R. 
Kruckeberg 4023 (WTU 172672). Some of the vegetation associated 
with Kruckeberg 4023 is as follows: Pseudotsuga menziesti, Quercus 


1960] KRUCKEBERG: SILENE 177 


Fic. 3. Flower of Silene marmorensis: a, single whole flower; b, carpophore with 
attached single petal and stamen. All x 7. 


kelloggii, Arbutus menziesii, Cornus nuttalliu, Quercus chrysolepis, Acer 
macrophyllum, Pinus lambertiana, P. ponderosa, Lithocarpus densiflora, 
Corvlus californica, Ceanothus integerrimus, and Cercis occidentalis ; also 
a sparse covering of such herbaceous plants as Cynoglossum occidentale, 
Poa sp., Galium sp., Stephanomeria sp., Polystichum lemmonu, P. munt- 
tum, Pteridium aquilinum, Hieracium albiflorum, Eriophyllum lanatum, 
Iris sp., and Smilacina racemosa. 

Department of Botany, 

University of Washington, 

Seattle 5, Washington 

LITERATURE CITED 


Hitcucock, C. L. and B. Macurre. 1947. A revision of the North American species 
of Silene. Univ. Wash. Publ. Biol. 13: 1—73. 

KRUCKEBERG, A. R. 1954. Chromosome numbers in Silene (Caryophyllaceae): I. 
Madrono 12: 238-246. 

———. 1955. Interspecific hybridizations of Silene. Am. Jour. Bot. 42: 373-378. 

-. 1960. Chromosome numbers in Silene (Caryophyllaceae). II. Madrono 15: 

in press. 


178 MADRONO [ Vol. 15 


FREED HOFFMAN 
1880-1959 


The botanical career of Mr. Freedom W. Hoffman, who died at his 
home near Guerneville, California, 13 November 1959, spanned a fifty- 
year period. Freed was born at Knights Landing, Yolo County, 30 
January 1880, where he lived until he went away to school at about the 
age of fifteen. As his mother was of French descent, Freed learned, as"a 
child, to speak French and he retained his fluency and interest in this 
language throughout his life. Following his graduation from Chico Normal 
School he studied art in New York City for several years. Upon his return 
to California he began a teaching career in which he achieved considerable 
success for something over a decade. 

On 24 August 1907, Freed married Jemella Gertrude Peugh. Through- 
out the forty-seven years of their married life, Jimmy regularly accom- 
panied Freed on trips into the remote back country. In the early days 
such trips were made with burro or mule, while later a Jeep served a 
similar purpose. 

While at Berkeley soon after his marriage, Freed became principal of 
the LeConte School, to which he often referred in later years as the first 
Junior High School in America. At about this time Freed studied with 
Professor Setchell and Professor Gardner in the Botany Department at 
the University of California at Berkeley. He mentioned to me several 
times Dr. Gardner’s offer of a teaching assistantship which, while it 
tempted him, was rejected in favor of a teaching position at San Francisco 
Normal School. 


1960 | MORRISON: FREED HOFFMAN 179 


I first met Freed and Jimmy at their home near Guerneville in the 
early summer of 1941. Freed had sent some interesting specimens of 
Streptanthus to the Herbarium at the University of California, Berkeley, 
for identification. In order to meet the collector and see the populations 
of plants, I drove to Guerneville. From the beginning of our eighteen- 
year friendship I was charmed by Freed. His slow, patient, deliberate 
approach to problems made a real impression on me. 

Because I was interested in Streptanthus and its distribution in relation 
to serpentine soils, because I enjoyed being with Freed and Jimmy, and 
because I could collect abundant fresh flowering plant material near 
Guerneville for class use, I went back to the Hoffmans’ several times dur- 
ing the summer of 1941. On one occasion Freed and I spent several days 
beginning the construction of a cabin on a remote hunting claim, which 
Freed had proven to be still part of the public domain in 1910 and 1911 
even though it had previously changed hands several times in land deals. 
Careful search of land office records and many weekends spent surveying 
had finally enabled him to file on the quarter section as a hunting claim. 
Its chief value lay in the existence of a spring not far below a ridge top. 
Freed and I hunted deer, fruitlessly, in the early mornings and the early 
evenings. During the day we began a cabin to replace the old one built by 
Freed in 1911. As we leisurely cut and notched the sill logs, Freed 
recounted, interspersed with discussions of the Pythagorean theorem, his 
reasons for leaving teaching to become an orchardist. 

Freed Hoffman was a man with a very considerable artistic talent. His 
oils and watercolors with which their home and guest cottage were hung 
made a lasting impression on all who saw them. The intricate woodcarving 
on the massive lauan loom which he built for Jimmy was still another 
evidence of his creative ability. As a carpenter, stone mason, botanist, 
botanical artist, Freed’s accomplishments were of professional quality. 
Certainly his abilities as a teacher were equally great. Yet he resigned his 
teaching position, left a career for which he seemed well fitted, and still 
in his thirties, took over the management of the extensive orchards owned 
by himself and Jimmy near Guerneville. 

With brush and palette, with hammer and saw, with pruning hook and 
picking basket and ultimately with plant press and seed bed, Freed found 
that he could see and sense the results of his labors almost immediately, 
while in teaching often many years passed before results were evident. 
Freed had the kind of patient sensitivity that would lead him to cut down 
his fruit picking speed by half in order not to discourage completely a 
youngster during his first day on the ladder, but he simply could not wait 
the many years to be greeted by a former pupil, now a mature man, who 
might say, ““You probably don’t remember me, Professor Hoffman, but 
you taught me geometry. .. .” 

With the realization in 1941 that serpentine outcroppings often sup- 
ported unusual populations of plants, Freed began a series of botanical 
trips which eventually brought his collections over the 4000 mark. Among 


180 MADRONO [Vol. 15 


his collections from remote and little-known serpentine areas is the type 
specimen of Haplopappus ophitidis (J. T. Howell) Keck. An Allium col- 
lected by Freed is likely to be the type of a new species. Especially in the 
genus Streptanthus, in which he published two new species in 1952, 
Freed’s numerous collections have increased greatly our knowledge of 
variation and geographic distribution. 

When World War II ended and gasoline, as well as new vehicles, 
became readily available, Freed purchased a Jeep in which he and Jimmy 
traveled widely in search of serpentine and “‘Streps.”’ Jimmy’s death in 
June, 1953, following their return from an extensive collecting trip in the 
Southwest, was a blow from which Freed found it almost impossible to 
recover. A trip to the Piedmont of North Carolina to visit Jimmy’s rela- 
tives and the thoughtful solicitude of friends finally restored in Freed his 
former interests. 

On 7 April 1955 Freed married Blanche Lenora Griden, who survives 
him. Blanche’s lively interest in Freed’s botanical studies and her devoted 
care during the trying time of Freed’s stroke and his lengthy and arduous 
convalescence have endeared her to those of us who came to know her 
through Freed. 

Freed’s ties with the profession of botany were primarily with members 
of the California Botanical Society and the personnel of the Herbarium 
at Berkeley. He corresponded rather regularly with Bacigalupi, Carter, 
Kruckeberg, McMillan, Mason, Morrison, and others interested in 
serpentine, Streptanthus, or both. His collections, his watercolor sketches, 
especially of Streptanthi, and his voluminous notes on various sections of 
this genus are on deposit in the Herbarium of the University of California 
at Berkeley —JouNn L. Morrison, State University, College of Forestry, 
Syracuse University. 


CLEARED CARDIOCARPON LATE-ALATUM LESQ., 
CORDAITEAN SEEDS FROM MICHIGAN! 


J. F. Davipson 


Arnold (1948) described Spermatites cylix from the Big Chief No. 8 
mine at St. Charles, Michigan, as appearing to be the apical portion of a 
very large spore. The present account may throw some light upon the 
nature of the object so designated, while extending our knowledge of 
the material previously identified (Arnold, 1949) as Cardiocarpon late- 
alatum Lesq. 

The Cordaitean seed that Lesquereux described as Cardiocarpon late- 
alatum (1879, Pl. LXXXV, figs. 46, 47; 1880, p. 568) is a small, rounded, 
slightly cordate body, about 9 mm. wide and 10 mm. long. The nucule, 


1 This work was financed in part by a grant from the University of Nebraska 
Research Council. 


1960} DAVIDSON: CARDIOCARPON 181 


as figured, is about 5 mm. wide, and is surrounded by a marginal wing 
which varies from 2 to 3 mm. in width. The apical portion of the wing 
is only slightly prolonged. Lesquereux expressed uncertainty about the 
validity of the distinction between this species and C. zonulatus and C. 
simplex, all three being from the sub-conglomerate at Pittston, by saying: 
“Perhaps these three forms, separated as species, represent the same, 
although the differences appear evident” (1880, p. 569). Of these three 
species, however, the specimens from Grand Ledge, Michigan, which are 
discussed below, show the closest resemblance to Lesquereux’ figures 46 
and 47 which represent C. late-alatum. While Lesquereux was familiar 
with Brongniart’s work in which the name Cardiocarpon was used for 
silicified seeds, he based his own separations on characters expressed in 
compressions. 

In 1955, large numbers of Cardiocarpon seeds were collected at Grand 
Ledge, Michigan. These occurred in the shale immediately below what 
Kelly (1933) designated as “‘Cycle A” in his Pennsylvania stratigraphy. 
In the dried shale, the seeds could be studied only as compressions, but 
when freshly-collected shale was submerged in water the shale immedi- 
ately started to disintegrate and some of the seeds, as well as other plant 
remains, could be recovered. 

An attempt to clear the seeds with concentrated nitric acid and potas- 
sium chlorate (Schultz’s solution) and 1 per cent ammonium hydroxide 
resulted in quite opaque preparations (fig. 1). This was apparently due 
to the intrusion of shale within the layers of the integument, and was 
eliminated by soaking the seeds in hydrofluoric acid previous to clearing. 
Such seeds were dehydrated in alcohol before mounting permanently in 
Diaphane. 

In addition to the more or less complete seeds, several other isolated 
fragments were cleared. Some of these fragments were portions of the 
nucellar region of the seed, which showed no evidence of the previously- 
surrounding integument tissue. During the clearing and mounting process, 
the nucellar portions showed a strong tendency to become separated from 
their enveloping integuments. Since the cuticle of the integument is more 
delicate than that of the nucellus, it is logical that the latter would 
be occasionally preserved after the disintegration of the surrounding 
integument. 

With the variation evident in these seeds, together with the variation 
originally recognized by Lesquereux, we are faced with two alternatives 
as regards the disposition of the specimens within our nomenclatural 
system. If we accept the names proposed by Lesquereux as denoting three 
species of Paleozoic seeds, then we can validly apply his names only to 
those specimens which agree exactly with his figured and described types. 
This implies that the vast majority of specimens which do not so agree 
will have to be described as new species. This might well be the case here. 
The alternative deals with probabilities. Since Lesquereux stated that his 
specimens from the same habitat were possibly conspecific, and since 


182 MADRONO [ Vol. 15 


CARA RC ROR ERTL NY NU EIR NOP MOOT TROP ROLLE LEER PER Sanne nconane te 


; ; 


Fics. 1-6. Cardiocarpon late-alatum Lesq. (Circles are 18 mm. in diameter.) 
Fic. 1. Mature seed. (Without treatment with hydrofluoric acid, the intruded shale 
obscures detail. Other figures are cleared after treatment with hydrofluoric acid. 
Figures 2—5 show gradually increasing size of nucellus). Fic. 2. Nucellus, although 
displaced in clearing, shows apical beak. Fic. 3. Nucellus in position. Fic. 4. Displaced 
nucellus, showing heavily cutinized basal region. Fic. 5. Nucellus found free in the 
shale. The integument was not preserved. Fic. 6. Portion of the base of a nucellus, 
found free. (This is the kind of structure described as Spermatites cylix Arnold, the 
type of which was studied in comparison.) 


1960} DAVIDSON: CARDIOCARPON 183 


comparable material has been found in another single habitat at Grand 
Ledge, it is probable that the variations encountered represent slight 
differences in the preservation process, and differences in the ages of the 
seeds when shed. The latter point is illustrated in the figures, which show 
variation not only in the over-all size of the seeds, but also in the size of 
the nucellar region. Were abortion involved, one might reasonably expect 
to find a series of small, aborted seeds and another series of more or less 
mature seeds, without the intermediate sizes. 

Of the foregoing alternatives, the latter appears the more logical, and 
less apt to result in a confusion of names. In the material collected at 
Grand Ledge, some of which is figured here, it is assumed that the varia- 
tion encountered represents differences in the maturity of individual 
seeds, and differential preservation of conspecific material. Thus it is here 
all referred to Cardiocar pon late-alatum Lesq. 

The seeds from Grand Ledge are flattened, circular to ovate in outline, 
9-15 mm. long by 9-11 mm. broad. The basal region shows an indenta- 
tion at the point of attachment, which extends almost to the swollen basal 
stalk of the nucellus, while the distal end appears as a deeply bifid beak. 
The surface of the integument appears to be composed of roughly isodia- 
metric cells, approximately 45 microns in diameter, except for those of 
the wing, which are about 20-25 microns broad and 60—100 microns long. 
The wing starts as a narrow band about 0.5 mm. wide near the base of the 
seed, and gradually increases in width upward to an observed maximum 
of 2.0 mm. In these specimens, the integuments, with the exception of the 
wings, were filled with clay, apparently bound with silica, since hydro- 
fluoric acid dispersed it. 

The nucellar body is very heavily cutinized, ovoid to globose, 2—7 mm. 
long, and equaily broad. At the proximal end, a compact tissue of heavily- 
walled cells forms a saucer-shaped base (figs. 4, 5) which in turn arises 
from a short, 1 mm. long cylinder of similar cells in which no vascular 
tissue is apparent. The base of this cylindrical stalk is somewhat swollen, 
the cell walls are thinner, the cells are slightly larger and have a glandular 
appearance. The tendency for the nucellar portion to separate from the 
integument is shown in figures 2 and 4, while figure 5 shows the heavily 
cutinized base of the nucellus beginning to break away from the upper 
portion, in a specimen found without the surrounding integument. In 
figure 5 also may be seen the region of attachment of the nucellar stalk 
to the integument. Figure 6, which is one of the fragments found in the 
shale and cleared, is merely the basal portion of the nucellar region of a 
seed. 

Although the apex of the nucellar body appears to be rounded in most 
specimens, closer examination shows that the specimens are incomplete. 
The smallest (fig. 2) and the largest (fig. 1) individuals show a definite 
attenuation at the apex such as might be expected to lead to a pollen 
chamber. 

The specimen shown in figure 6 was compared with the type material 


184 MADRONO [Vol. 15 


of Spermatites cylix Arnold, and they appear to be conspecific. Hence, 
Spermatites cylix probably refers to the basal portion of a nucellus from 
a Cordaitean seed comparable to Cardiocar pon late-alatum. 


Department of Botany, 
University of Nebraska, Lincoln 


LITERATURE CITED 


ARNOLD, C. A. 1948. Some cutinized seed membranes from the coal-bearing rocks of 
Michigan. Bull. Torrey Club 75: 131-146. 

. 1949. Fossil flora of the Michigan coal basin. Contr. Mus. Paleont. Univ. 
Mich. 7: 131-269. 

KeELty, W. A. 1933. Pennsylvania stratigraphy near Grand Ledge, Mich. Journ. Geol. 
41: 77-89. 

LESQUEREUX, L. 1879. Atlas to the coal flora of Pennsylvania. Second Geol. Surv. Pa. 
Harrisburg. 

. 1880. Coal flora of Pennsylvania. Second Geol. Surv. Pa. Harrisburg. 


THE BASIC CHROMOSOME NUMBER OF THE 
GENUS NEPTUNIA 


(LEGUMINOSAE-MIMOSOIDEAE) 
B. L. TURNER AND O. S. FEARING 


The genus Neptunia is composed of about ten or eleven species of 
annual and perennial herbs. Its members are widely distributed in the 
tropical and subtropical regions of the world. Five species are endemic 
to the Old World (three in Australia, two in India); two are cosmopoli- 
tan, occurring in wet habitats, principally in tropical regions; and three 
or four are confined to North and South America. 

The region with the greatest number and diversity of taxa appears to 
be Texas and adjacent Mexico where four or five species are represented 
(Turner, 1951). From a standpoint of floral morphology, this area also 
retains one of the least modified species in the genus (Neptunia lutea). 

The first chromosome count reported for a species of the genus was by 
Dnyansagar (1952). He reported a number of n = 18 from sectioned 
anther material of the Indian species, NV. triquetra. However, the camera 
lucida drawing documenting this count appears to show 18 somatic 
chromosomes and is perhaps but a portion of the complement of a single 
somatic cell of premeiotic ‘‘mother cell’ tissue. 

Turner and Beaman (1953) reported counts for three unnamed Ameri- 
can taxa of Neptunia as 2n = 28. Their counts were obtained from 
somatic cells of sectioned root tip material. The only other report for the 
genus has been that of Frahm-Leliveld (1953) who listed an aproximate 


1 All of the described taxa in the genus, except this species, have some flowers with 
anantherous staminodia modified into petaloid structures. NV. lutea has flowers with 
the stamens all alike and anther-bearing. 


1960] TURNER AND FEARING: NEPTUNIA 185 


TABLE 1. SPECIES OF NEPTUNIA EXAMINED FOR CHROMOSOME NUMBER 


Species Seed Source and Chromosome 
Voucher Collection Number 


—__——. 


*Neptunia dimorphantha Domin AusTRALIA (Seed communicated 2n = 28 (fig. 3) 
by Div. of Plant Industry, 
C.S.I.R.O., Canberra City). Q227 


*Neptunia gracilis Benth. AUSTRALIA (As above) C884 2n = 56 
*Neptunia monosperma Benth. AvsTRALIA (As above) W652 2n — 28 (fig. 2) 
Neptunia plena (L.) Benth. INDONESIA (Reported by 2n = 78? 
Frahm-Leliveld, 1957) 
Neptunia triquetra Benth. Inp1a (Reported by n—18? 
Dnyansagar, 1952). 
*Neptunia prostrata (Lam.) Inp1A. Raipur (Seed communi- 2n=—56 (fig. 1) 
Baill. cated by Dr. V. R. Dnyansagar). 
Turner s.n. 
*Neputnia lutea (Leavenw.) Texas. Galveston County: 2n = 28 fig. 5) 
Benth. Turner 2189 
*Neputnia lutea (Leavenw.) Texas. Galveston County: 2n = 28 
Benth. Turner 2923 
*Neptunia pubescens var. Texas. Galveston County: 2n = 28 
floridana (Small) Turner Turner 2194 
*Neptunia pubescens var. Texas. San Patricio County: 2n = 28 (fig. 4) 
lindheimeri (B.L. Robinson) M.C. Johnston 541338 
Turner 


* Indicates new report for the genus. 


count for V. plena as 2n = + 72; ina later paper Frahm-Leliveld (1956) 
in reporting the same species, apparently settled on the number 2n = 78, 
though the drawing documenting this count is not easily interpreted. 

Because of the differing base numbers for the genus reported by these 
workers, and because of the known unibasic nature of most genera in the 
Leguminosae, the present authors have reinvestigated the previous re- 
ports for the taxa reported by Turner and Beaman and in addition have 
investigated four species previously unreported.” 

MATERIALS AND MeEtHops. Chromosome counts listed as new in the 
present paper (Table 1) were made from root tip cells of germinating 
seeds using a squash technique outlined by Turner and Fearing (1959). 
Polyploid cells were noted in the tissue of all the taxa examined, though 
diploid cells appeared to be more common and this is the number given 
in Table 1. 

RESULTS AND Discussion. Altogether, counts for eight of the approxi- 
mately ten species in the genus have been reported (Table 1). Except for 
the doubtful count of n = 18 for N. triquetra and the count of N. plena 


* Attempts to obtain seed of the controversial NV. triquetra have been unsuccessful. 
The authors are grateful to Dr. V. R. Dnyansagar who so kindly furnished the seed 
of N. prostrata used in the present study. 


186 MADRONO [ Vol. 15 


2 x ® 
© @ & gr ® a ~ fi 
Pr oofe a es eb .8 - 
<P, ce ; te_ &y ” 
= . ye od a @ 
° p a is & e ¢ % 
o 8 o ® a . os 
oo ; Ps 
*e 
| 2 6 2 
| oe 
S > " ® 6@ y ~ 
Wer * ot ea Wry’) nate . 
@ 0 ae 
- be @ 6 ¢ 8 We 
© € 
tly a ® 96 
e Wa no ° 
3 4 5 


Fics. 1-5. Camera lucida drawings of the mitotic chromosomes in Neptunia spp.: 
1, N. prostrata (2n = 56); 2, N. monosperma (2n = 28), late metaphase, 2 pair of 
chromosomes have already separated and are shown in the “unpaired” condition; 
3, N. dimorphantha (2n = 28); 4, N. pubescens var. lindheimeri (2n = 28), pro- 
phase; 5, N. lutea (2n = 28). (X ca. 1200.) 


(2n= 78), all reported counts have been on a base of x = 14. The species 
here reported are from taxa occurring on several continents, and one of 
them, V. lutea, has a “primitive” floral structure and occurs in a region 
where several diverse taxa are found. These facts make a base number of 
x = 14 for the genus seem more plausible than that of a multibasic pat- 
tern, particularly since NV. triquetra and N. plena are not especially dif- 
ferent morphologically from species with known counts of 2n = 28. In 
addition it might be noted that related genera of the tribe Adenanthereae, 
in which Neptunia is usually included, are also on a base of x = 14 (e.g. 
Prosopis and Dichrostachys, Darlington and Wylie, 1956). 


SUMMARY 


Chromosome numbers for six species of Neptunia, all on a base of 
x — 14, are presented for the first time. These counts were obtained from 
diverse species which occur naturally in Australia, India and Texas. In 
view of the known constancy of base numbers for most genera of the 
Leguminosae, and in view of the poor documentation for differing base 


1960] WEBBER: YUCCA 187 


numbers reported by other workers, it has been concluded that x = 14 is 


probably the correct base chromosome number for the genus. 
The Botany Department, and 
Plant Research Institute, 
University of Texas, Austin 


LITERATURE CITED 


DaruincTon, C. D., and A. P. Wy te. 1956. Chromosome atlas of flowering plants. 
MacMillan Co., New York. 519 pp. 

DNYANSAGAR, V. R. 1952. Embryological studies in the Leguminosae. IV. A contribu- 
tion te the embryology of Neptunia triquetra Benth. Indian Acad. Sci. Proc., 
Sect. B. 36: 1-11. 

FRAHM-LELIVELD, J. A. 1953. Some chromosome numbers in tropical leguminous 
plants. Euphytica 2: 46-48. 

. 1957. Observations cytologiques sur quelques Légumineuses tropicales et sub- 
tropicales. Rev. Cytol. et Biol. Veg. 8: 273-287. 

TuRNER, B. L. 1951. Revision of the United States species of Neptunia (Leguminosae). 
Am. Midl. Natur. 46: 82-92. 

— —., and J. H. Beaman. 1953. Chromosome complements in Desmanthus (Legu- 
minosae). Field and Lab. 21: 47-50. 

. and O. S. FrEarinc. 1959. Chromosome numbers in the Leguminosae. II. 

African species, including phyletic interpretations. Am. Jour. Bot. 46: 49-57. 


HYBRIDIZATION AND INSTABILITY OF YUCCA 


JoHN MILTON WEBBER! 


The contention that hybridization is largely responsible for the wide- 
spread variability of southwestern yuccas (2,4,9)* is supported by the 
following facts. 1) Cross-pollination is enhanced by the yucca’s depend- 
ence upon the yucca moth for pollination. 2) Two or more species fre- 
quently occur in mixed stands or near each other. 3) The karyotypes of 
all species are strikingly similar (cf. 1). 4) Inter- and intra-specific pol- 
linations produce equally abundant seed. 5) There is little difference in 
the degree of relationship and the ability to hybridize. 6) Many variants 
exhibit specific characteristics of two or more species. 7) Apparent 
hybrids are frequently more fertile than ‘“‘typical” species. 8) Progenies 
of apparent hybrids are composed of two or more types. These facts, 
however, pertain only to putative hybrids and the conditions favoring 
hybridization. Although a few garden and artificial hybrids (3,8) have 
been cited, their characteristics, behavior, and fertility have not been 
recorded. The present study of artificial hybrids indicates that yuccas are 
genetically similar and that hybridization among native plants is common. 


1 Agronomist, Crops Research Division, Agriculture Research Service, U.S. D.A., 
Berkeley, California. Work done in cooperation with Botany Department, Univer- 
sity of California, Berkeley, California. 


2 Numbers in parentheses refer to Literature Cited. 


188 MADRONO [Vol]. 15 


The seeds of the species involved in the hybirds were collected in the 

following localities: 

Yucca glauca Nutt., Grant, New Mexico. 

Yucca elata Engelm., White Sands National Monument, New Mexico. 
Yucca constricta Buckl., Big Spring, Texas. 

Yucca schidigera Roezl ex Ortgies, Riverside, California. 

Yucca arizonica McKelv., Nogales, Arizona. 

Yucca neomexicana Woot. & Standl., Kenton, Oklahoma. 

The pollinations were made at Riverside, California (9), and the 
hybrids were grown in the University of California Botanic Garden, 
Berkeley, California. Specimens of the hybrids are in the University of 
California Herbarium, Berkeley, California. 


FERTILITY AND MEIOTIC BEHAVIOR 


The percent of F, fruit obtained from self-pollinations and the via- 
bility of F,; and Fs seed are given in Table 1. 

The microsporocyte divisions of the hybrids were identical or strik- 
ingly similar. Each consistently formed 5 large and 25 small bivalents 
and exhibited few or no irregularities in either the first or the second 
division. All tetrads appeared normal, and only 6 percent of the matured 
grains were abortive. 


TABLE 1. PERCENT OF Fy} SELFED FRUIT AND VIABILITY OF Fy AND F2 SEED 


Percent Percent of Fy Percent 
germination fruit from germination 
of Fj seed* self-pollinations* of Fe seed 
VE OULUC ay COL ee 84 88 86 
Y.constricta X Y. schidigera ................ 64 72 ie 
Y.arizonica X Y. neomexicana .............. 54 24 68 


VY. Gyt20n1Cd XV SIOUCE a, 62 32 ia 


* Fruit percentages based on 25 pollinations and seed germination based on ger- 
mination test of 50 seeds. 


CHARACTERS INDICATIVE OF DERIVATION 


The majority of characters of the hybrids are either intermediate in 
nature or approach those of the parents. The most helpful characteristics 
in recognizing the derivation of the hybrids are the following: 

Yucca glauca X Y. elata: the low height of the inflorescence (7.05 cm. 
above foliage) and the greenish, swollen styles approximate those of Y. 
glauca, while the large head of leaves and the paniculate inflorencence 
resemble those of Y. elata. The hybrid is very similar to Y. intermedia 
McKelv. var. ramosa McKelv. (5,p.120, pl. 46) and to plants reported 
possibly to be Y. elata X Y. glauca hybrids (9, p. 63, pl. 41). 

Yucca constricta X Y. schidigera (figs. 1, 2): the non-fleshy fruit 
(3.20 cm. thick, 6.35 cm. long) and large, angular seeds (7.20 by 11.1 
mm.) are characteristic of Y. constricta, while the indehiscent fruit and 


1960] WEBBER: YUCCA 189 


iz 
fy 
re 


Fics. 1-4. Yucca parents and hybrids. Fic. 1. Habit of Yucca constricta « Y. 
schidigera. Fic. 2. Yucca constricta X Y. schidigera and parents: a, pistil and stamens 
of Y. constricta; b, pistil and stamens of Y. schidigera; c, pistil and stamens of hybrid ; 
d, capsule of hybrid; e, cross section of hybrid capsule; f, seeds of Y. schidigera; g, 
seeds of hybrid; kh, seeds of Y. constricta. Fic. 3. Habit of Yucca arizonica = Y. 
neomexicana. Fic. 4. Yucca arizonica X Y. glauca and parents: a, pistil and stamens 
of Y. glauca; b, pistil and stamens of hybrid; c, pistil and stamens of Y. arizonica; d, 
fruit of hybrid; e, seeds of Y. glauca; f, seeds of hybrid; g, seeds of Y. arizonica. 


190 MADRONO [ Vol. 15 


thick, rough seeds containing ruminate endosperm are characteristic of 
Y. schidigera. The only similar fruits found on native plants (9, p.56, 
pl.34) occurred on plants belonging to the Y. glauca alliance, but these 
fruits contained typical capsular seeds. 

Yucca arizonica X Y. neomexicana (fig. 3) and Y. arizonica X Y. 
glauca (fig. 4) are fairly similar. They differ chiefly in the leaf blades 
being mainly concavo-convex and the flowers globose in Y. arizonica X 
Y. neomexicana, while the leaf blades are largely plano-convex and the 
flowers campanulate in Y. arizonica X Y. glauca. The most significant 
features of these hybrids are the following: 1) the non-fleshy, indehiscent 
fruits and the large, angular, thick, rough seeds containing a ruminate 
endosperm (fig. 4), which characterize the dehiscent-indehiscent origin 
of the hybrid; 2) the exceptionally long, conical ovary (3.80 cm.) and 
fruits (10.3 cm.) (fig. 4), which separate the hybrids from Y. constricta 
x Y. schidigera and possibly characterize all hybrids between species of 
the Baccatae series of McKelv. and dehiscent fruit species; and 3) the 
paniculate-racemose nature of the inflorescence proper (lower half 
cuneiform, upper half racemose) (fig. 3), which probably characterizes 
hybrids between plants with typical paniculate and racemose inflores- 
cences. Although no native plants exhibiting the first two of the preceding 
features have been reported, those with paniculate-racemose inflores- 
cences are fairly common. Such inflorescences are characteristic of Y. 
utahensis McKelv. and VY. intermedia, and they are common among 
native yuccas reported to be possible hybrids (9, pp.56—68, pl.31, 33, 40). 


YUCCA CONSTRICTA X Y. SCHIDIGERA Fy 


Second generation seedlings of only Y. constricta X Y. schidigera have 
been grown. Leaf variations between individuals of a year-old popula- 
tion are as follows: 1.40 cm. wide and 12.0 cm. long to 0.63 cm. wide and 
21.3 cm. long; thick, rigid and straight to thin, flexible and falcate; 
light green to dark green; and thin, entire margin to corneous, denticulate 
margin. The leaves of several of the seedlings are considerably broader 
than those of equally as old seedlings of such broadleaved species as Y. 
faxoniana (Trel.) Sarg., Y. supicola Scheele, and Y. gilbertiana (Trel.) 
Rydb. Although the seedling leaves of Y. schidigera have denticulate 
margins they become filiferous several months before they are a year old. 
Corneous, denticulate margins are characteristic of matured leaves in 
species of the sections of Hesperoyucca, Clistocarpa, and the series Rupi- 
colae of McKelv. 

DISCUSSION 


The normal meiotic behavior and the high fertility of the hybrids 
indicate that the chromosomes of the parental species are homologous 
and differ only with respect to certain genes. Furthermore, since the 
similar karyotypes of yuccas suggest parallel speciation, it is very likely 
that there is a considerable degree of genetic affinity between the majority 
of species. Under these conditions it appears that the major barrier to 


1960 | WEBBER: YUCCA 191 


interbreeding among the native plants of Yucca is spatial isolation. Al- 
though these suppositions are supported by putative, natural hybrids 
involving many species, several species are not included. It is very 
probable, however, that hybrids involving the latter species have not 
been recognized, or that barriers other than genetic affinity or geo- 
graphical separation occur. 

The fact that no apparent hybrids involving Y. schottu Engelm. have 
been reported is unquestionably due to the late flowering season of this 
species. Although similar flowering barriers occur between several other 
species, usually the flowering period within a group of associated species 
overlaps. The failure of Y. arizonica X Y.neomexicana and Y. arizonica 
xY. glauca to fruit freely was probably due to the long style and closed 
stigma lobes, which commonly prevented fertilization. It is likely that 
the capitate stigma of Y. whipplei Torr. is a structural barrier and that 
similar barriers exist in other species. 

Although dehiscent- and indehiscent-fruited species are frequently as- 
sociated, no reputed natural hybrids between them have been reported. In 
general appearance Y. constricta X Y. schidigera resembles an indehis- 
cent, baccate-fruited yucca, while Y. arizonica X Y.neomexicana and Y. 
arizonica X Y. glauca resemble a capsular-fruited yucca. If these hybrids 
were admixed with wild plants, they would undoubtedly be considered 
hybrids between baccate-fruited species and capsular-fruited species 
respectively. Their true origin could not be determined without careful 
examination, unless fruits and seeds were available. 

The genera Yucca and Agave have identical karyotypes (6,10) and a 
similar distribution (1,8,9), and both are highly unstable (1,7,9). Al- 
though much of the diversity in Agave is attributed to hybridization, 
considerable is due to polyploidy, mainly allotetraploidy. The polyploids 
in Agave have a wider distribution and, in general, a greater vegetative 
development than the diploids (1). No polyploids have been reported in 
Yucca. Both thick, succulent-leaved species (VY. faxoniana, Y. torrevi 
Shafer.) and thin, flaccid-leaved species (Y. constricta, Y. glauca) are 
diploids (6). Furthermore, the distribution of these diploids is as exten- 
sive as the combined diploid-polyploid distribution of Agave. Yucca ex- 
tends from south-central Mexico to South Dakota (8,9), whereas Agave 
extends from northern South America to Utah and Nevada (1). 


LITERATURE CITED 


1. GRAMIcK, E. B. 1944. A karyosystematic study of the genus Agave. Am. Jour. 
Bot. 31: 283-298, illus. 

2. HarnEs, L. 1941. Variations in Yucca Whipplei. Madrono 6: 33-45, illus. 

3. Moton, G. 1914. Monograph, Le Yucche. Published in Italian. 

.McKetvey, S. D. 1938. Yuccas of the Southwestern United States. Part One, 
150 pp., illus. Jamaica Plain, Mass. 

. 1947. Yuccas of the Southwestern United States. Part Two, 192 pp., 

illus. Jamaica Plain, Mass. 

. and K. Sax. 1933. Taxonomic and cytological relationships of Yucca and 

Agave. Jour. Arnold Arb. 14: 76-81, illus. 


aN 


OL 


192 MADRONO [Vol. 15 


7. STANDLEY, P. C. 1920-1926. Trees and shrubs of Mexico. Contr. U.S. Nat. Herb. 
23: 1-1721, illus. 

8. TRELEASE, W. 1902. The Yucceae. Missouri Bot. Gard. Ann. Rept. 13: 27-133, 
illus. 

9. WEBBER, J. M. 1953. Yuccas of the Southwest. Agr. Monograph No. 17, U.S. Dept. 
Agr. 97 pp. illus. 

10. WHITAKER, T. W. 1934. Chromosome constitution in certain monocotyledons. 
Jour. Arnold Arb. 15: 135-143, illus. 


REVIEW 


The Physiology of Forest Trees, a Symposium held at the Harvard Forest, April, 
1958, under the auspices of the Maria Moors Cabot Foundation. Edited by KENNETH 
V. THIMANN with the assistance of WILLIAM B. CRITCHFIELD and Martin H. ZimM- 
MERMAN. xvi-+ 678 pp., illustr. The Ronald Press. New York, N.Y. $12.00. 

Although principles of plant physiology are the same for all forms of plant life, 
the methods of research will differ depending on what kind of plants are used in 
experimental work. Truly, it is a great difference in applying plant physiology to cul- 
tivated annuals, such as barley or oats on one hand or to forest trees, that may be 
several hundred years old and many feet tall, on the other. Plant physiologists, work- 
ing with forest trees have felt for a long time a need for a get-together to discuss their 
common problems. Dr. Kenneth V. Thimann, Professor of Biology, Harvard Univer- 
sity, was responsible for organizing the first International Symposium on The Physi- 
ology of Forest Trees. The symposium was held under the auspices of the Maria 
Moors Cabot Foundation. Over thirty scholars from several European countries, 
Canada and United States gathered at the Harvard Forest, Petersham, Massachusetts, 
in April 1957. The topics discussed included: Water relations and sap movement; 
Photosynthesis; General Biochemistry; Mineral nutrition; Translocation; Root 
Growth and other phenomena; Photoperiodism and Thermoperiodism; and Repro- 
duction. The papers were edited by Dr. Thimann, with the assistance of Dr. William 
B. Critchfield and Dr. Martin H. Zimmermann, and published in one volume. Pub- 
lication of this volume signifies, if not the birth, at least a formal recognition, of a 
new branch of Plant Physiology. 

The import of this book on the further development of Forest Tree Physiology 
will be felt for a long time—N. T. Mrrov, Pacific Southwest Forest and Range 
Experiment Station, Berkeley, California. 


NOTES AND NEWS 


ALPHABETICAL List OF FAMILIES FOR Munz anv Keck. An alphabetical list of 
families, giving the page on which each family begins, is available for Munz and 
Keck, A California Flora. It is intended for pasting to the inside of the back cover. 
Copies may be had by sending a request for the number desired, together with a 
stamped self-addressed envelope, to Rancho Santa Ana Botanic Garden, 1500 North 
College Avenue, Claremont, California. 


Some publications of interest follow: 

Origin of Primary Extraxylary Stem Fibers in Dicotyledons, by Amélie Blyth. 
University of California Publications in Botany 30 (2): 145-232, pls. 1-23. 1958. 
sh Lye 

Secondary Phloem of Calycanthaceae, by Vernon I. Cheadle and Katherine Esau. 
University of California Publications in Botany 29 (4): 397-510, pls. 60-67, 109 
figs. in text. 1958. $2.25. 

Ontogeny of the Inflorescence and the Flower in Drimys winteri var. chilensis, 
by Shirley Cotter Tucker. University of California Publications in Botany 30 (4): 
257-336, pls. 24-33, 43 figs. in text. 1959. $1.50. 


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. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume [7 1916-1929") oe at es 9200 
Volume IT, 1930-1934 . . ... . 5.00 
Volume II], 1935-1936 . . .°2 72 “3:00 
Volume IV, 1937-1938 . 2 . «> 5 2 5.00 
Volume V, 1939-1940 . . .. . . 5.00 
Volume VI, 1941-1942 3)... _.% 2) 35.00 
Volume VII, 1943-1944 . . . . . . 5.00 
Volume VIII, 1945-1946 .... . 5.00 
Volume IX, 1947-1948 . . ... . 6.00 
Volume X, 1949-1950 . .... . 7.00 
Volume XI, 1951-1952. . . . . . 8.00 
Volume XII, 1953-1954 . . . . . . 8.00 
Volume XIII, 1955-1956... . . 8.00 
Volume XIV, 1957-1958 .... . 8.00 
Single numbers, 

Volumes land 2 ~~ <9. 45.29 50 
Succeeding numbers, 

(except Volume XI, No.2) . . . . 1.00 

Volume Xa, (No: 2 225 2 ae ee eo 


The subscription price of MapRoNo is $6.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


—— 
VOLUME 15, NUMBER 7 JULY, 1960 


Contents 


PAGE 
SANICULA DESERTICOLA, AN ENDEMIC OF Baja CALI- 
FORNIA, Peter H. Raven and Mildred E. Mathias 193 


A NEw SPECIES OF VALERIANA FROM BRAZIL, 
Frederick G. Meyer 197 


STUDIES IN WESTERN VIOLETS, IX. MISCELLANEOUS 
SPECIES IN THE SECTIONS NOMIMIUM AND CHAMAE- 


MELANIUM, Milo S. Baker 199 
CHROMOSOME NUMBERS IN SILENE (CARYOPHYLLA- 

cEAE). II, A. R. Kruckeberg 205 
A NEw SPECIES OF ZINNIA FROM MEXIco, 

A. M. Torres 215 
A CoMMENT ON COLD SUSCEPTIBILITY OF PONDEROSA 

AND JEFFREY PINES, Willis W. Wagener 217 
DocUMENTED CHROMOSOME NUMBERS OF PLANTS 219 


Review: Mont E. Lewis, Carex—its Distribution and 
Importance in Utah (Jack Major) 221 


NOTES AND NEws: EDWARD LEE GREENE CORRESPOND- 
ENCE, Robert P. McIntosh; UROSPERMUM PICROIDES 
IN BERKELEY, Annetia Carter 222 


A WEST AMERICAN JOURNAL OF BOTANY 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


Entered as second-class matter at the post office at Berkeley, California, January 29, 
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription 
price $6.00 per year. Published quarterly and issued from the office of Madrofio, 
Herbarium, Life Sciences Building, University of California, Berkeley 4, California. 


BOARD OF EDITORS 


HERBERT L. MAson, University of California, Berkeley, Chairman 
EpcAr ANDERSON, Missouri Botanical Garden, St. Louis 
LyMAN BENSON, Pomona College, Claremont,. California. 
HERBERT F. COPELAND, Sacramento College, Sacramento, California. 
Joun F. Davipson, University of Nebraska, Lincoln. 

IvAN M. JoHNsTON, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MitprepD E. Maruias, University of California, Los Angeles 24. 
MarION OwNBEY, State College of Washington, Pullman. 

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: Herbert G. Baker, Department of Botany, University of California, 
Berkeley, California. First Vice-president: Malcolm A. Nobs, Carnegie Institution of 
Washington, Stanford, California. Second Vice-president: Herbert F. Copeland, Sac- 
ramento City College, Sacramento, California. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley, California. Corre- 
sponding Secretary: Francia Chisaki, Department of Botany, University of California, 
Berkeley, California. Treasurer: John Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1960] RAVEN & MATHIAS: SANICULA 193 


SANICULA DESERTICOLA, AN ENDEMIC OF 
BAJA CALIFORNIA 


PETER H. RAVEN AND MILprReEpD E. Matutas! 


One of the more restricted species of northern Baja California is Sani- 
cula deserticola Bell, known from a few disjunct populations at the north- 
ern margin of the Sonoran Desert. One population is known from near E] 
Marmol] at the headwaters of the Arroyo de San Fernando, while others 
are in the yellow hills northwest of Rancho Arenoso and near Rancho 
Aguajito, both in the drainage of the Arroyo del Rosario. At the last- 
mentioned locality, it was found growing abundantly on the northwest- 
facing slopes of a yellow conglomerate hill, 3.6 miles west of Rancho 
Aguajito (Raven, Mathias, and Turner 12,678), associated with Rosa 
minutifolia, Euphorbia misera, Yucca whip plei eremica, Idria columnaris, 
Eriogonum fasciculatum, E. scalare, Encelia californica var. asperifolia, 
Calandrinia maritima, Harfordia macroptera, Brodiaea pulchella, Lavia 
platvglossa, and Filago californica, as well as two species of Agave and one 
each of Dudleya, Mammillaria, Echinocereus, Echinocactus, and Opuntia. 
This curious mixture of characteristic members of the California flora 
and such species as /dria columnaris, restricted to the Sonoran Desert, 
clearly demonstrates the unique ecological position of Sanicula deserticola. 

The subfamily Saniculoideae of the Umbelliferae, with some 260 
species, like the other subfamilies Hydrocotyloideae and Apioideae, has 
apparently had a long and independent evolutionary history. The distri- 
bution of the extant genera of Saniculoideae shows clearly that they have 
developed within the Arcto-Tertiary Geoflora and have been associated 
with it for a long time, perhaps since late Mesozoic time when this Geo- 
flora is first recognized in the fossil record. Several of the genera in this 
subfamily are restricted to areas of Arcto-Tertiary-derived deciduous 
forest in eastern Asia. Others range south along mountain chains to 
Africa. The genus Eryngium is world-wide in distribution, whereas the 
genus Sanicula is exceedingly widespread in the Northern Hemisphere, 
with some of its species occurring also in the Southern Hemisphere. Shan 
and Constance (1951) considered the section Sanicula (Sanicla), with 
about one-third of the species of the genus Sanicula, the main trunk of the 
genus. Some species of this section are widespread in Eurasia, and their 
present distribution suggests development of the section from a northern 
stock with subsequent southerly migrations. In North America S. mari- 
landica L. and S. trifoliata Bickn., which Shan and Constance considered 
probably the least advanced species, occur as common associates of the 
eastern deciduous forests. The distribution of this section is therefore 
closely related to that of the Arcto-Tertiary Geoflora. 


1 The authors would like to acknowledge the helpful suggestions of Dr. Harlan 
Lewis in the preparation of this paper. 


Maprono, Vol. 15, No. 7, July 13, 1960. 


194 MADRONO [Vol. 15 


In western North America, the genus Sanicula is represented by sec- 
tion Sanicoria, which Shan and Constance showed was probably derived 
from members of section Sanicula. Sanicula deserticola is one of fourteen 
species comprising section Sanicoria, which is the most diverse within the 
genus in vegetative and reproductive characters. The development of the 
Madro-Tertiary Geoflora in early Tertiary time in western North Amer- 
ica involved a segregation of species from the dry margins of the tropics 
and from the Arcto-Tertiary Geoflora,—a segregation fostered by the 
ever-increasing influence of aridity, particularly the loss of summer rain, 
and of more extreme temperature variation over much of the West (Axel- 
rod, 1958). Increasing environmental diversity in this area has resulted 
in large measure from the continuation of this process of progressively 
increasing aridity, and the evolution of section Sanicoria was doubtless 
correlated with the development of this climatic and topographic diversity. 
Bell (1954) showed that the different diploid species of section Sanicoria 
differ in their environmental preferences. The existence of localized 
species in specialized environmental situations is unique to this section. 
Thus Sanicula peckiana F. Macbr. occurs only on serpentine, S. saxatilis 
Greene occurs only on volcanic or serpentinized rocks, S. arctopoides H. 
& A. occupies coastal bluff and dune habitats, and S. maritima Kell. is 
a local species confined to moist adobe soil near the coast. Although some 
species of the section are less sharply differentiated ecologically, they 
are, as a Class, plants of relatively xeric habitats in chaparral and various 
oak-conifer woodland associations derived from the Madro-Tertiary 
Geoflora. One species, S. graveolens Poepp., ranges widely north and 
east, and others occur in suitable sites north along the Pacific Coast, but 
most of the species of section Sanicoria are members of the California 
flora in the broad sense as delimited by Howell (1957). The occurrence of 
two mesophytic species of Sanicula on the western coast of South America 
is, we believe, the product of relatively recent long-range transtropical 
dispersal of the type discussed by Grant (1959) and by Raven and 
Lewis (1959). 

On morphological grounds, the species most closely related to Sanicula 
deserticola is S. bipinnatifida Dougl., but nevertheless the two are amply 
distinct. Sanicula bipinnatifida is found colonially in open rocky grass- 
land from Washington south to the northern edge of Baja California, in 
regions with average annual precipitation ranging from about 12 to 40 
inches. The populations of S. deserticola are about 150 miles south of 
the range of S. bipinnatifida in a region with an average annual precipita- 
tion that ranges from perhaps 2 to 7 inches (fig. 1). It is therefore found 
in a habitat which with respect to aridity is very extreme for members of 
section Sanicoria and for the genus Sanicula as a whole. In its native 
habitat S. deserticola probably flowers whenever it has sufficient water. 
This is suggested by its flowering response when grown at the University 
of California, Los Angeles, where it flowered twice a year when supplied 
with abundant water. 


1960 | RAVEN & MATHIAS: SANICULA 195 


me memes ee we 
. 
~ 


—— 
a 


ed 


corr 


30 ae 


C==21 S. bipinnatifida 
+ +§&, deserticola 


“+= Margin of Sonoran Desert 
(after Shreve) 


Fic. 1. A portion of western North America, showing ranges of Sanicula deserticola 


and S. bipinnatifida and approximate line of demarcation between desert and chapar- 
ral in Baja California. 


196 MADRONO [Vol. 15 


Rodriguez (1957) found that the vessels of Sanicula deserticola have 
the smallest mean length for any member of the family that he examined; 
this, together with its long taproot and clumped habit, indicate its ad- 
vanced position compared with S. bipinnatifida. One may reasonably infer 
that populations ancestral to both became differentiated at the southern 
margins of their distribution in response to an arid climate. The disjunct 
populations of S. deserticola may have been somewhat more continuous 
during pluvial periods of the Pleistocene, but at which time period this 
species became spatially and genetically isolated from S. bipinnatifida 
cannot be demonstrated. 

The area of Sanicula deserticola lies along the southern margins of 
the transition area between the Californian chaparral and the Sonoran 
Desert, as defined by Shreve (1936). Shreve noted that species which 
are endemic in the transition area are preponderantly of northern rela- 
tionship, while the plants of the desert area are more sharply confined to 
their own formation. He explained this relationship by the fact that the 
only requirement for the long southward extension of a chaparral species 
is a relatively moist habitat, however restricted in area this may be, while 
the requirements for northward extension of desert species are more com- 
plex. The approximate boundary between the two areas is shown in 
figure 1. Sanicula deserticola is certainly such a species of northern affini- 
ties. A parallel distributional pattern and relationship are found in the 
shrubby Salvia chionopeplica Epling; this species also consists of a few 
disjunct populations in north-central Baja California, and it is closely 
related to other species of section Audibertia, such as Salvia leucophylla 
Greene. The distribution of most species of Salvia sect. Audibertia is asso- 
ciated with the California flora and therefore similar to that of Sanicula 
sect. Sanicoria. 

Within the genus Sanicula, the section Sanicoria, apparently stemmed 
from ancestors adapted to relatively mesic sites within the area of the 
Arcto-Tertiary Geoflora and occupied successively drier and drier habi- 
tats offered by the expansion and differentiation of the Madro-Tertiary 
Geoflora. The species of section Sanicoria that occupies the most xeric 
habitats is Sanicula deserticola. Relatively few perennial Umbelliferae 
occur in such habitats, and most of those that do are members not of 
Saniculoideae, but of the larger and more diverse subfamily Apioideae. 

Department of Botany 


University of California 
Los Angeles, California 


LITERATURE CITED 


AXELROD, D. I. 1958. Evolution of the Madro-Tertiary Geoflora. Bot. Rev. 24:433-509. 

BELL, C. R. 1954. The Sanicula crassicaulis complex (Umbelliferae). Univ. Calif. Publ. 
Bot. 27:133-230. 

GRANT, V. 1959. Natural history of the phlox family. Vol. I. Systematic botany. The 
Hague: Martinus Nijhoff. 

Howe tt, J.T. 1957. The California flora and its province. Leafl. West. Bot. 8:133-138. 


1960] MEYER: VALERIANA 197 


Raven, P. H., and H. Lewis. 1959. The relationship of clarkias from two continents. 
Brittonia 11:193-205. 

RopricueEz, R. L. 1957. Systematic anatomical studies on Myrrhidendron and other 
woody Umbellales. Univ. Publ. Bot. 29:145-318. 

SHAN, R. H., and L. Constance. 1951. The genus Sanicula (Umbelliferae) in the Old 
World and the New. Univ. Calif. Publ. Bot. 25:1-78. 

SHREVE, F. 1936. The transition from desert to chaparral in Baja California. Madrono 
3:257-264. 


A NEW SPECIES OF VALERIANA FROM BRAZIL 
FREDERICK G. MEYER 


Valeriana glechomifolia sp. nov. Herba perennis omnino puberula, 
longe repens; caulis tenuis foliosus; folia opposita, laminis suborbiculari- 
bus vel orbiculari-reniformibus, crenato-dentatis, petiolis 0.6—1.4 cm. 
longis; inflorescentia erecta, 4-10 cm. longa; flores hermaphroditi; corol- 
la infundibuliformis 2—2.5 mm. longa; achaenia oblonga vel elliptica ali- 
quantulum ampulliformia ubique puberulenta; calycis limbus brevicu- 
puliformis plus minusve dentatus. 

Uniformly puberulent long-creeping perennial, rooting at the nodes; 
stems slender, terete, about 1 mm. in diameter, leafy, the internodes 1—2.5 
cm. long; leaves opposite, erect or ascending, the blades suborbicular to 
orbicular-reniform, 0.6-1.5 cm. wide, sometimes truncate at the base, uni- 
formly crenate-dentate, the petioles 0.6—-1.4 cm. long; inflorescence an 
aggregate or compound dichasium, erect, 4-10 cm. long, arising on a 
slender stalk from leaf axils along the creeping stems, with 1-3 pairs of 
leaves, the terminal dichotomies about 1—3 cm. wide in anthesis, later more 
diffuse, the bracts 1-3 mm. long, more or less spathulate, the flowers her- 
maphroditic; corolla infundibuliform, 2—2.5 mm. long, glabrous, the tube 
gibbous, the lobes 5, spreading, slightly unequal; stamens 3, exserted, 
2-lobed; style 3-lobed; achenes oblong to elliptic, about 1.5 mm. long, 
somewhat ampulliform, more or less oblique, uniformly puberulent; 
calyx-limb short-cupuliform, more or less dentate. 

Specimens examined. Brazit. Santa Catarina, Mun. Bom Retiro: 
Campo between Fazenda Campo dos Padres and Fazenda Santo Antonio, 
Campo dos Padres, alt. 1400-1650 m., November 21, 1956, L. B. Smith & 
R. Klein 7800 (type US); same locality, January 24, 1957. L. B. Smith & 
R. Reitz 10383. 

The combination of creeping habit and leaves that resemble those of 
Glechoma hederacea quickly distinguishes V. glechomifolia from all other 
New World valerianas. The fruit of V. glechomifolia allies it with other 
Brazilian valerianas with a coronate calyx-limb in the group with V. sali- 
carifolia, V. chamaedryfolia, V. foliosa, and V. eichleriana, but the uni- 
formly puberulent and more or less oblique achenes of V. glechomifolia 


198 MADRONO [Vol. 15 


Fic. 1. Valeriana glechomifolia: A, habit, * 34; B, inflorescence and single leaf, 
< 3; C, fruit, flower, and stamens, 10, Smith & Klein 7800. 


differ from the uniformly glabrous and regular achenes of the aforemen- 
tioned Brazilian species. 

Students of South American Valeriana reserved the segregate genus 
Phyllactis for species with coronate calyx. My own studies previously 
on North American species and now on those of South America indicate 
that single character differences, especially floral differences, are insuf- 
ficient in differentiating segregate genera. Indeed, the specialized calyx, 
either coronate or pappus-like, and more especially the sculpturing of the 
cypselate achene combine with vegetative characters in the differentia- 
tion of species of South American Valeriana. 

This interesting discovery by Lyman B. Smith of the Smithsonian Insti- 


1960 | BAKER: WESTERN VIOLETS 199 


tution, Washington, D.C., is one of a long series of new plants discovered 
on Dr. Smith’s fruitful collecting trip to Santa Catarina in 1956-57. 
Crops Research Division, Agricultural Research Service 
U.S. Department of Agriculture 
Beltsville, Maryland 


STUDIES IN WESTERN VIOLETS, IX. 
MISCELLANEOUS SPECIES IN THE SECTIONS NOMIMIUM 
AND CHAMAEMELANIUM 


Mito S. BAKER 


This paper treats four taxa of Viola in the sections Nomimium and 
Chamaemelanium—a newly described species, a change of status from 
species to subspecies for a second taxon, observations confirming the 
specific status of a third taxon, and a newly described subspecies. In addi- 
tion to my own specimens at the North Coast Herbarium, I have cited 
specimens from the United States National Herbarium, the New York 
Botanical Garden, the California Academy of Sciences, and the University 
of California Herbarium; to the curators of these latter herbaria I express 
my appreciation. 


Viola Aliceae sp. nov. Herba exigua omnino puberulenta cauli supra- 
terrano brevissimo suppressoque et rosella basali foliorum ac floribus uno 
duobusque folias parum excedentibus instructa cauli subterreno gracili 
bracteas squamas simulantes gerente, tota ex radice fusiformi longitudine 
variabili (ut apud specimina typica videtur) crescens; folia oblongo-ovata 
vel elliptica obscure undulato-dentata decurrentia 2 vel 3 cm. longa 
dimidio lata petiolis laminas longitudine aequantibus; stipulae incon- 
spicuae lanceolatae marginis laceratis; flores caesii longitudine 1 cm. 
parum excedentes petalis angustis eis lateralibus aliquantulo barbatis; 
stigma ut id Violae aduncae ebarbatum; et capsula et semina hucusque 
ignoti. 

A small plant, 5 to 11 cm. high, caulescent but the aerial stem unde- 
veloped, bearing a rosette of leaves and one or two flowers slightly above 
the leaves; finely puberulent throughout; subterranean stem slender with 
scale-like bracts; taproot variable in length (as in type sheet); leaves 
long-ovate to elliptic, obscurely undulate-dentate, decurrent, 2 to 3 cm. 
long and half as wide, on petioles about as long; stipules inconspicuous, 
lanceolate with lacerate margins; flowers lavender, slightly more than 1 
cm. long; petals narrow, the lateral slightly bearded; stigma like that of 
Viola adunca but without beards; capsule and seeds unknown. Figs. 1, 2. 

Type. Mexico. Near kilometer 34 post on Mexico City—Cuernavaca 
highway, altitude 9000 feet, 4. Y. and J. E. Wilcox in 1948 (UC 1,200,- 
778). Topotype. A. Y. and J. BE. Wilcox 22, 1946. Viola Aliceae should be 


200 MADRONO [Vol. 15 


assigned to Section Nomimium Ging., although the stigma and style are 
somewhat different from other members of this section. 

When making their second collection of the violet in 1948, the one 
cited above as the type, Mr. and Mrs. Wilcox dug deeply to investigate 
the character of the root system. In 1952 they returned to the type 
locality, but a diligent search of several hours failed to reveal any plants 
of this violet. 

In some cases the underground stem measures only a few centimeters, 
and could represent the normal growth from a seed over a few years, such 
as the upper right hand plant (topotype, Wilcox 22) on the type sheet. In 
other cases the vertical subterranean stem is very slender and possesses a 
number of scale-like bracts at more-or-less regular distances apart, such 
as the plant on the upper-left-hand side of the type sheet (type collection, 
Wilcox in 1948). How such a stem was formed is not clear. It might be 
formed by underground stolons from another plant such as that repre- 
sented by the topotype. However, I do not know of such a case in any 
other species of Viola. 

Two other collections of Viola resemble V. Aliceae in having similar 
slender underground stems as well as similar style and stigma, but they 
differ from that species in being wholly glabrous [Quebrada Honda, Du- 
rango, Palmer 227 (US, NY); Sierra Madre near Chihuahua, Townsend 
& Barber 94 (US, NY)]|. Their exact relationship to V. Aliceae cannot 
be determined at this time. Some of the plants on the Palmer sheet, how- 
ever, represent another species unrelated to V. Aliceae. 


VIOLA LOBATA Benth. subsp. psychodes stat. nov. V. psychodes Greene, 
Pittonia 3:318. 1898. 


In 1898, E. L. Greene proposed the name V. psychodes for a violet col- 
lected near Waldo, Oregon. Since then, this taxon has been collected at 
many locations in California. In general V. psychodes resembles V. lobata, 
which is abundant in California. The differences between these two taxa 
do not seem of sufficient magnitude to merit retention of both of them as 
species; therefore I propose treating V. psvchodes as a subspecies of the 
common IV. /obata Benth, section Chamaemelanium Ging. Both have simi- 
lar root systems, a naked stem with leaves and flowers borne near the 
summit, leaves that may be variously lobed or even entire, and similar 
pistils (figs. 3-6). Plants of both taxa have the habit of producing a 
single radical leaf for the first year. 

There are, however, some differences between the two, the most sig- 
nificant being that subsp. psvchodes is glaucous throughout, while V. 
lobata is puberulent. Another difference is leaf thickness—the average for 
V. lobata is 0.0036 inch, while for subsp. psychodes it is 0.0057 inch. The 
measurements for these averages were obtained from leaves of ten collec- 
tions of each taxon, using a Starrett micrometer which is accurate to one 
ten thousandths of an inch. 

It may be of interest to note that several of the collections of subsp. 


1960 | BAKER: WESTERN VIOLETS 201 


Fics. 1-9. Viola. Fics. 1, 2, V. Aliceae: 1, habit, * 34; 2, pistil, side view. Fics. 
3, 4, V. lobata subsp. lobata: 3, pistil, side view; 4, pistil, dorsal view. Fics. 5, 6, 
V. lobata subsp. psychodes: 5, pistil, side view; 6, pistil, dorsal view. Fics. 7, 8, 9, 
V. oxyceras: 7, flower, X 1%; 8, pistil, side view; 9, pistil, dorsal view. All pistil illus- 
trations * 9. 


202 MADRONO [Vol. 15 


psvchodes were made in serpentine areas, but I have not been able to 
ascertain whether or not the plant is restricted to serpentine soil. 


Representative specimens. OREGON. JosEPHINE County: Little Rock Creek, 2 
miles southwest of O’Brien, Constance & Rollins 2995 (NCH, UC) ; open wocds near 
Waldo, Oregon, April 20, 1887, Howell (UC); near Kirby, Baker 5526 (NCH). 
CALIFORNIA. Hearst Castle [Sacramento Canyon], July 22, 1902, Setchell & Dobie 
(UC) ; serpentine on west side of Sacramento River Canyon, Wagnon 1616 (NCH). 
Butte County: east of Oroville, west of Brush Creek Ranger Station, Cantelow 
4527 (CAS). DEL Norte County: on serpentire, Gasquet, Baker 211 (NCH), Baker 
285 (NCH); near Gasquet, Tracy 10011 (UC); headwaters Shelly Creek, Oregon 
Mountain, Hoffman (NCH). Prumas County: Highway 39 east of summit near 
Mineral, Baker 8104 (NCH). SuHasta County: Delta, Applegate 5396 (NCH); Cas- 
tella, June 19, 1923, Bethel (CAS); Castle Rock, Ripley & Barneby 9642 (CAS); 
on serpentine, Dunsmuir, Baker 8050 (CAS, UC), Hall & Babcock 4031 (UC), Heller 
7927 (UC); on serpentine, Shasta Retreat, July 4, 1911, Condit (UC); Mount Eddy 
(some genes of V. lobata Benth.), June 1, 1946, Parker (UC). Sisktyvou County: 
Rainbow Ridge above Sulloway Creek, about 1% miles west of Mount Shasta City, 
June 13, 1936, Babcock & Stebbins 2000 (UC) ; near Mount Shasta City, Baker 5525 
(NCH) ; serpentine on east slope Scott Mountain, Hoffman 2434 (NCH); 6% miles 
north of West Brand Road, Happy Camp, Hoffman 2572 (NCH); trail 1 mile west 
of Lookout, Clear Creek, Hoffman 3514 (NCH). Trinity County: near Granite 
Peak, Baker 205 (NCH); Minersville to Trinity Center, Eastwood & Howell 4912 
(CAS); between Baylers and Trinity Center, Hoffman 2420 (NCH); Trinity Alps 
Resort, Cantelow 995 (CAS); North Fork Mountain, 3500 feet, Hoffman 3504 
(NCH); near boundary of Trinity and Siskiyou counties, Scott Mountain, Howell 
13618 (CAS); north slope of Scott Mountain, 3700 feet, Cantelow 1453 (CAS) ; 
south slope of Scott Mountain, Cantelow 1452 (CAS); Scott Mountain Road, 11% 
miles north of Carrville, June 9, 1939, Cantelow (CAS); Nash Mine near Carrville, 
June 23, 1931, Van Dyke (CAS); 17 miles north of Carrville, May 21, 1936, Can- 
telow (CAS); 3 miles north of Carrville, May 21, 1936, Cantelow (CAS). 


VIOLA OXYCERAS Greene. On the Pacific Coast of North America there 
are two distinct species of caulescent blue-flowered violets in the section 
Nomimium Ging.: V. adunca J. E. Smith, which is found from sea level 
to a fairly high altitude, and V. oxyceras Greene, which grows in the 
mountains down to about 4000 feet. The latter was considered by Sereno 
Watson to be only a variety of what we now call V. adunca |V. canina 
var. oxyceras Wats., Brew. and Wats., Bot. Calif. 1:56. 1876]. This view- 
point may have arisen because in some localities interbreeding between 
these two species has produced intermediate forms; at most localities, 
however, the pure types remain distinct. 

Aside from their similarity in flower color, the two have little in com- 
mon. The leaves of V. adunca are ovate, more or less subcordate at the 
base, and have a pointed apex; those of V. oxyceras are much thinner, 
sometimes wider than long, with a cuneate or truncate base (never cor- 
date), and are rounded at the apex. The foliage of V. adunca shows some 
puberulence (except for occasional glabrous races), while that of V. oxy- 
ceras is without exception entirely glabrous. The form of the flower in 
these two is different; the flowers of V. adunca are larger, having the 
petals spread in approximately one plane, and having a rather blunt spur; 
those of V. oxyceras are smaller, the upper four petals lie mostly in one 


1960] BAKER: WESTERN VIOLETS 203 


plane, but the lower petal lies below in another plane (fig. 7). The beak 
of the style in V. adunca, although always directed essentially downward, 
may be pointed forward, or may be at right angles to the axis of the style, 
or may even assume a still more reflexed position, and it has always a 
smaller diameter than the head of the style. The beak of V. oxyceras, 
with its stigmatic tube always pointing forward, is very different, having 
a diameter nearly as large as the head of the style (figs. 8, 9). 

The seeds of V. adunca and V. oxyceras differ also, in both shape and 
in size. Although the seeds of V. adunca vary quite widely in size, they 
are never as small as the seeds of V. oxyceras. But more important than 
size, the seeds of these two taxa are of different shape. The more rounded 
seed of V. oxyceras is indicated by a length to width ratio of 1.85 to 1, 
while in the case of V. adunca (from six localities) the ratio is 1.98 to 1. 
With these differences in mind, the maintenance of the two species seems 
justified. 


Representative specimens. CALIFORNIA. High mountain near Donner Pass, 
1865, Torrey 34 (UC), type of V. canina var. oxyceras Wats. ALPINE County: Lake 
Alpine, Allen 542 (UC); near Lake Alpine, 7,500 feet, Peirson 11567 (UC). AMADOR 
County: Silver Lake, 7,500 feet, Baker 5557 (NCH). ButTE County: Jonesville, 
1,500 m., Copeland 662 (UC); Jonesville, Copeland 1219 (NCH). Ext Dorapo 
County: Highway 50, 1 mile east of Strawberry, 6,500 feet, Robbins 1712 (UC); 
near Lyons Creek about 4 miles south of Wright Lake, 6,700 feet, Robbins 2020 
(CAS, UC), Robbins 2022 (NCH). Fresno County: Vidette Creek, 10,500 feet, 
1948, Dyer (CAS); Bubb’s Creek, at base of East Vidette, 1948, Chabaud (CAS) ; 
Second Recess, 9,000 feet, Raven 5697 (CAS). Inyo County: Cottonwood Lakes, 
11,000 feet, Alexander & Kellogg 3417 (UC). TuLARE County: Tyndall Creek, 1916, 
Campbell (CAS). LassEN County: Summit Lake, McCalla 646 (NCH); east fork 
of King Creek on Cinder Cone Trail, 7,000 feet, Jepson 4110 (JEPS). Mariposa 
County: Eagle Peak, Yosemite, 7,200 feet, Jepson 4372 (JEPS). Mopoc County: 
High Grade District, northern Warner Mountains, 7,000 feet, Smith 948 (JEPS). 
PrLacErR County: above Donner Lake, Copeland 1883 (UC); below Cisco, Heller 
12713 (UC); Summit Valley, Howell 18577 (CAS). PLumas County: Little Grass 
Valley, Baker 9966 (NCH); 1% miles west of Johnsville, 5,300 feet, Cantelow 4560 
(CAS). SHasta County: 22 miles southeast of McCloud, Cantelow (NCH). SrerRA 
County: Webber Lake, 8,000 feet, Baker (NCH). Sisktvou County: Marble Moun- 
tain, Chandler 1585 (NCH). Trenama County: near Morgan, 5,500 feet, Hall & 
Babcock 4326 (UC). TULARE County: Camp 170, nine miles north of Mt. Silliman, 
Brewer 2807 (UC); South Fork Kaweah River, 8,500 feet, Ferris & Lorraine 10850 
(UC); Center Basin, 11,200 feet, 1948, Howell 25057 (CAS); Second Lake, Center 
Basin, 11,400 feet, Munz 12565 (NCH). TuoOLUMNE County: Kennedy Lake, Hoover 
1458 (UC). 


Vr1oLa Bakert Greene subsp. shastensis subsp. nov. A subspecie Bakeri 
capsulis pubescentibus et paginis sepalorum pilos paucos breves adpressos 
gerentibus discedit. 

This subspecies differs from V. Bakeri subsp. Bakeri (section Chamae- 
melanium Ging.) in its pubesent capsules and in the presence of a few 
short appressed hairs on the faces of the sepals. 

During the flowering stage, before the appearance of the capsules, 
subsp. shastensis may be distinguished from subsp. Bakeri by a careful 
examination of the sepals. In subsp. Bakeri, the faces of the sepals are 


204 MADRONO [Vol. 15 


entirely glabrous and without any short, appressed hairs such as those 
found on the faces of the sepals in subsp. skastensis. In other respects, as 
far as I have observed, these taxa are identical. 

Type. Postpile Camp, altitude 6000 feet, western Tehama County, Cali- 
fornia, July 1, 1955, Baker 13045 (UC 1,199,915); topotype, July 10, 
1954, Baker 12961 (NCH). 

Since its discovery in western Tehama County in 1954, V. Bakeri subsp. 
shastensis has been found to occur from southern Oregon south in the 
western slopes of the Sierra Nevada of California to Amador County, as 
detailed below. 

Other collections. OREGON: Road to Dutchman’s Peak, May 30, 1938, Rowntree 
(NCH). CALIFORNIA. Trinity County: Scott Mountain, Cantelow 4576 (CAS), 
Baker 13139 (NCH), Wagnon & Barbe (NCH). Lassen County: “devastated area’’ 
earth slide, Highway 89, north side Mount Lassen, Baker 11975 (NCH). AmMapor 
County: Silver Lake, altitude 7,400 feet, Wagnon 1620 (NCH). 

One may ask, why should such inconspicuous characters justify the 
creation of a new subspecies? Our reply is that in the subsection Nuttal- 
lianae Becker, naked capsules have long been regarded as a diagnostic 
character separating the V. Nuttalli species complex from the V. pur- 
purea complex. However, recent studies in V. praemorsa of the Nuttalla 
complex have shown that two subspecies have some puberulence of the 
capsules. Nevertheless, the most widely distributed subspecies, V. prae- 
morsa subsp. major (Hook.) Baker, subsp. linguaefolia (Nutt.) Baker, 
and subsp. praemorsa have naked capsules. 

One also wonders how and when subsp. shastensis derived its charac- 
ters. Examination of V. praemorsa subsp. arida Baker and subsp. oregona 
Baker & Clausen reveals the fact that they both possess the same pattern 
of sepal pubescence as does V. Bakeri subsp. shastensis. In addition, these 
two subspecies of V. praemorsa have more or less pubescence on their cap- 
sules. Although there is no known overlap at the present time in the dis- 
tribution of V. Bakeri subsp. shastensis and these two Great Basin sub- 
species of V. praemorsa, it is known that an arm of the Great Basin Flora 
extends from southern Oregon southwest along the Klamath River into 
the Shasta and Scott valleys, and this lends credence to the belief that 
V. praemorsa subsp. arida will be found to occur somewhere in this area 
not far distant from where we collected V. Bakeri subsp. shastensis. 

From bud fixations of subsp. shastensis made in the Scott Mountains, 
(Baker 13133, 13139) Dr. Jens Clausen and Dr. Malcolm Nobs of the 
Carnegie Institution of Washington at Stanford obtained a chromosome 
count of 2n = 48, showing this taxon to be an octoploid. Viola Bakeri 
subsp. Bakeri and V. praemorsa subsp. arida and subsp. oregona are also 
octoploid, 2n = 48. Therefore, it may be assumed that gene interchange 
could have taken place between these entities at some past time, thus 
possibly accounting for the pubescence of the sepals and capsules of V. 
Bakeri subsp. shastensis. 


Santa Rosa Junior College 
Santa Rosa, California 


i) 
(=) 
GL 


1960] KRUCKEBERG: SILENE 


CHROMOSOME NUMBERS IN SILENE 
(CARYOPHYLLACEAE). II. 


A. R. KRUCKEBERG! 


The accumulation of cytotaxonomic data from members of the sub- 
family Silenoideae (Caryophyllaceae) continues, with this second report 
adding chromosome numbers of thirty-two species counted for the first 
time. Additional records are given here for twenty of the species included 
in the first paper of this series (Kruckeberg, 1954). In all, the present 
report comprises chromosome counts for one hundred and fifty-six collec- 
tions from continental North America, Hawaii, Europe, and Asia. The 
chromosome numbers are listed in Table 1; figures 1-28 are camera lucida 
drawings of chromosome complements for species hitherto uncounted. 

Records of chromosome numbers of the Silenoideae are being accumu- 
lated for two reasons. Chromosome counts have intrinsic value as part of 
the total self-portrait of a taxon. Secondly, the ploidy level of any two 
populations qualifies the success with which those population samples 
can be used in hybridizations to determine species interfertilities (see 
Kruckeberg, 1955). 

All counts have been made from squashes of microsporocytes handled 
in the manner outlined in Kruckeberg (1954). Most of the material I 
have collected in native habitats of western United States. Collections 
from eastern United States and from localities outside continental North 
America were kindly furnished by other botanists. 


TABLE 1. DIPLoOID CHROMOSOME NUMBERS IN LYCHNIS AND SILENE 


2n Number 
Chromosome of 
WESTERN NorRTH AMERICAN SPECIES Number Collections 

Lychnis drummondi (Hook.) Wats. 48 5 
Silene aperta Greene 48 1 
S. bridgesii Rohrb. 48 1 
S. californica Durand. Tetraploid 48 2 
Hexaploid (2 1 

S.campanulata Wats. 48 1 
S.clokeyiH.& M. 96 1 
S. douglasit Hook. 48 22 
S. grayi Wats. 48 3 
S. hookeri Nutt. ex T.& G. Te 3 
S.invisaH.& M. 48 3 
S.laciniata Cav. subsp. major H.& M. Octoploid 96 4 
S. laciniata Cav. subsp. greggiz (Gray) H.& M. Tetraploid 48 10 
S.lemmonii Wats. 48 5 


1 Supported by funds from the State of Washington Initiative No. 171 and by the 
National Science Foundation, Grant G-1323. 


206 MADRONO [Vol. 15 
2n Number 

Chromosome of 

Number Collections 
S. marmorensis Kruck. 48 1 
S. menziesii Hook. Diploid 24 2 
Tetraploid 48 12 
S. montana Wats. 48 4 
S. nuda (Wats.) H.& M. 48 3 
S. nuda subsp. insectivora (Hend.) H.& M. 48 1 
S. occidentalis Wats. 48 1 
S. oregana Wats. 48 1 
S. parishi Wats. 48 3 
S. parryi (Wats.) H.& M. Tetraploid 48 9 
Octoploid 96 6 
S. petersoni Maguire 96 2 
S.repens Patrin subsp. australeH.& M. 24 2 
S. sargenti Wats. 48 3 
S. scaposa Robins. 48 1 
S. scoulerit Hook. 48 1 
S. spaldingi Wats. 48 i 
S. thurberi Wats. 48 1 
S. verecunda Wats. subsp. verecunda 48 1 
S. verecunda Wats. subsp. andersoni (Clokey) H. & M. 48 1 
S. verecunda Wats. subsp. platyota (Wats.) H.& M. 48 10 
S. williamsii Britt. 24 D 
S. wrightia Gray 96 1 

EASTERN NorTH AMERICAN SPECIES 
Silene antirrhina L. 24 1 
S. caroliniana Walt. 48 2 
S.caroliniana subsp. wherryi (Small) Clausen 48 1 
S. poly petala (Walt.) Fern. & Schub. 48 il 
S. regia Sims. 48 2 
S. rotundifolia Nutt. 48 1 
S. stellata (L.) Ait. sensu lat. 48 5 
S. subciliata Robins. 48 1 
S. virginica L. 48 1 
SPECIES OUTSIDE CONTINENTAL NorTH AMERICA 

Lychnis wilfordit Maxim. 24 if 
Petrocoptis pyrenaica Braun. 24 1 
Silene keiskei Miq. 24 2 
S. repens Patrin. var. latifolia Turcz. 48 2 
S. struthioloides Gray 24 1 
S. species from Nepal 24 2 


DISCUSSION 


Having determined chromosome numbers for all but five of the thirty- 
three perennial species of Silene native to western North America, I am 
emboldened to make some guarded generalizations. Though chromosome 
numbers of the remaining uncounted species as well as any additional 
records for species previously counted may prove to be further exceptions, 
it seems safe to state that the tetraploid level (2n=48) is by far the 
common one among the western species. The tetraploid level is main- 


1960] KRUCKEBERG: SILENE 207 


tained even more rigidly for the eastern North American species. Apart 
from S. ovata and S. nivea (Nutt.) Otth., for which chromosome num- 
bers are not yet known, the other seven eastern perennial species are 
uniformly tetraploid. The present data emphasize the observation made 
in an earlier paper (Kruckeberg, 1954)—namely, that North America is 
the home of the polyploids whereas the continental areas of Eurasia are 
inhabited predominantly by diploid species. At first glance, this geo- 
graphic difference in average ploidy level might suggest that the Eurasian 
species were ancestral to the North American ones. The Siberia-Aleutian 
Islands land bridge, a path well beaten by the biogeographers—if not by 
the biota themselves—would seem the likely route. Yet one species for 
which I have obtained counts on both the Asian and North American 
plants defies the usual west-to-east migration route. Szlene repens Patrin 
in Japan (var. /atifolia Turcz.) is tetraploid while the North American 
variety, australe Hitch. & Maguire, is diploid. Therefore, I would offer 
an alternative hypothesis: North American polyploid species could have 
originated on this continent from diploids as a common phenomenon 
whereas polyploidy might have been a rare event among the ancestors of 
the Eurasian Silenes. 

The present list of chromosome numbers permits tentative conclusions 
to be drawn about certain species. 


1. SILENE CALIFORNICA Dur. AND S. HooKERI Nutt. On morphological 
grounds it is easy to see a close relationship between S. californica and 
S. hookeri. The prostrate habit, the showy flowers (red versus pink), the 
characteristic geniculation of the pedicels in fruit, the large black seeds, 
and the partial sympatric distribution in the Coast Ranges and Siskiyou 
Mountains all suggest a close affinity. Up to now, though, I was con- 
vinced that the two species were isolated by a barrier of ploidy difference. 
S. hookeri is consistently hexaploid (2n=72) while most collections of 
S. califorinca have been tetraploid (2n=48). The discovery of a hexa- 
ploid S. californica population and the subsequent production of a fertile 
F', hybrid (to be discussed in a later paper) support my intuitive feeling 
that the two species are closely related. 


2. THE THREE SUBSPECIES OF SILENE LACINIATA Cav. According to 
Hitchcock and Maguire (1943), three regional facies of S. lacintata occur 
in the southwest. Silene laciniata subsp. laciniata is widespread in Mexico, 
S. laciniata subsp. major H. & M. is confined to coastal southern Cali- 
fornia, and S. laciniata subsp. greggii (Gray) H. & M. occurs in Arizona, 
New Mexico, Texas, and adjacent northern Mexico. Subspecies major is 
octoploid (2n=96) while subsp. greggii is tetraploid (2n=48). Much 
wider sampling of this polymorphic species will be necessary before it 
can be fully evaluated cytotaxonomically. Since subsp. major and subsp. 
greggu are isolated both spatially and genetically, it becomes a moot ques- 
tion as to whether subsp. greggii should be restored to its former level of 


208 MADRONO [Vol. 15 


species. The center of distribution of S. /aciniata is undoubtedly in the 
plateau and mountain regions of north-central Mexico. Subspecies greg gi 
is a mid-continent derivative from this center while subsp. major is the 
coastal segregate. Additional collections of S. laciniata greggt, made in 
Arizona, New Mexico, and Texas in the summer of 1959, proved also to 
be tetraploid. However, in the two weeks of searching in the Mexican 
states of Nuevo Leon and Chihuahua, I did not encounter any plants of 
S. laciniata, though the species is frequently reported from this section of 
México. 

3. DIPLOID AND TETRAPLOID SILENE MENZIESII Hook. Further sampling 
of this wide-ranging species reaffirms my earlier report (Kruckeberg, 
1954) of the predominance of the tetraploid level (2n=—48). As against 
twelve collections of tetraploids, only two diploid samples are recorded 
in this current listing. One of the diploids was collected in the Sierra 
Nevada of California and the other in central Idaho. It would appear 
that the distribution of diploids is random and their occurrence rare. It 
is possible that the tetraploids are autoploid in origin. The sporadic dis- 
tribution of the diploids, plus the not infrequent occurrence of quadri- 
valent pairing in certain tetraploids lends support to this hypothesis. 
Amphiploid origin is less likely, even though the Idaho diploids do occur 
sympatrically with another diploid species, S. repens Patrin subsp. 
australis H. & M. Apart from being rhizomatous and having general vege- 
tative similarity, North American S. repens is not too likely a parent for 
tetraploid S. menziesii. 


4. TETRAPLOID AND OCTOPLOID SILENE PARRYI (Warts.) H. & M. With 
the present data it is now possible to define rather clearly the limits of 
the two ploidy levels in S. parryi. The tetraploid forms (2n=48) occur 
exclusively east of the crest of the Cascade Range, and then eastward to 
the mountains of northeastern Washington (and presumably northern 
Idaho), terminating in the Rocky Mountains of Montana and Canada’. 
The octoploid segment (2n=96) of the species has a much more restricted 
distribution; as yet it has been found only along the Cascade Range and 
the Olympic Mountains of Washington. 


2A collection of a tetraploid parryi made on Snowdrift Mountain in southern 
Idaho (Kruckeberg 4520) suggests that the tetraploid form extends southward along 
the Continental Divide. 


EXPLANATION OF FIGURES 1-12 

Fics. 1-12. Chromosomes of Silene microsporocytes. Fic. 1. S. aperta, I1M 
(Kruckeberg 3407). Fic. 2. S. clokeyi, Diak. (Kruckeberg 3911). Fic. 3. S. grayt, 
Diak. (Kruckeberg 3772). Fic. 4. S. invisa, 1M (Kruckeberg 2897b). Fic. 5. S. lacini- 
ata greggit, Diak. (Kruckeberg 3878). Fic. 6. S. lemmonii, Diak. (Kruckeberg 3530). 
Fic. 7. S. montana, Diak. (Kruckeberg 3529). Fic. 8. S. occidentalis, 1 T (Kruckeberg 
3696). Fic. 9. S. petersonii, Diak. (Kruckeberg 3904). Fic. 10. S. repens australis, 
Diak. (Kruckeberg 4290). Fic. 11. S. scaposa, Diak. (Kruckeberg 4031). Fic. 12. S. 
spaldingii, Diak. (Daubenmire, s.n.). Diak., diakinesis; I M, first metaphase; II M, 
second metaphase; I T, first telophase. —_> 


1960] KRUCKEBERG: SILENE 209 


Fics. 1-12. Chromosomes of Silene microsporocytes. 


210 MADRONO [Vol. 15 


5. SILENE WILLIAMSII BriTT., S. SEELEYI Mort. & THOMPS., AND S. 
MENZIESII Hook. These three species are grouped together because of 
their close morphological kinship. Szlene williamszi of central Alaska and 
S. seeleyi are both diploid so far as is known. The fact that some collec- 
tions of S. menziesa are also diploid makes gene exchange among the 
three species potentially possible. Hitchcock and Maguire (1947, p. 48) 
speculate that exceptional specimens of S. menziesii from western Canada 
show ‘“‘contamination” with S. williams. I have made artificial hybrids 
in various combinations with the three diploids; some of the hybrids are 
reasonably fertile. Details of these and other interspecific hybridizations 
will be discussed elsewhere. 


GEOGRAPHIC DISTRIBUTION OF COLLECTIONS 


The origin of each collection that provided a chromosome count is 
given below. All collections are represented by specimens deposited in the 
University of Washington Herbarium. To those botanists who collected 
living plant material for this study, the author wishes to express his 


sincere appreciation. 
WESTERN NortH AMERICAN SPECIES 


LYCHNIS DRUMMONDII (Hcook.) Wats. 2n=48. Arizona. Apache County: 12 miles 
from Red Rock, Lukachukai Mountains, Gould and Phillips 4835. Coconino County: 
Head of Kaibab Trail, north rim of Grand Canyon, Kruckeberg 3901. Nervapa. Clark 
County: ridge along trail to Charleston Peak, Kruckeberg 3910. NEw Mexico. Catron 
County: between Bursom Forest Camp and Willow Creek, Mogollon Mountains, 
Kruckeberg 4658; 5 miles northeast of Collins Park on road to the plains of St. 
Augustine, Kruckeberg 4702. 

SILENE APERTA Greene. 2n=48. CALIFORNIA. Tulare County: 1 mile west of Hockett 
Meadows, on trail to Atwell Mill, Sequoia National Park, Kruckeberg 3407. 

SILENE BRIDGESII Rohrb. 2n=48. CaLirornIA. Tuolumne County: Hog Ranch, 
near Mather, Pray s.n. 

SILENE CALIFORNICA Durand. 2n=48. Tetraploid. CaLirornta. Tulare County: 
Echo Point, near Moro Rock, Sequoia National Park, Kruckeberg 3394. Mendocino 
County: 4 miles north of Laytonville on U.S. Highway 101, Kruckeberg 3918. 

SILENE CALIFORNICA Durand. 2n==72. Hexaploid. CALIFORNIA. Del Norte County: 
gravel bar along Smith River at Gasquet, Aruckeberg 3919. 

SILENE CAMPANULATA Wats. 2n=48. CALIFORNIA. Siskiyou County: 12 miles south- 
west of Etna on Somes Bar road, Hitchcock 20237. 

SILENE CLOKEYI H. & M. 2n=96. Nevapa. Clark County: along ridge, 2 miles 
from summit of Charleston Peak, Kruckeberg 3911. 

SILENE DOUGLASIT Hook. 2n=48. CALIFORNIA. Plumas County: rocky flats above 
Elwell Lake, Kruckeberg 2897 ; rocky flats between Sand Lake and Packer Lake, near 
Sierra Buttes, Kruckeberg 3697. Trinity County: granite talus at Emerald Lake, 
Trinity Alps, Kruckeberg 3743; gravelly alluvium near Portugese Camp between 
Emerald Lake and Morris Meadows, Trinity Alps, Kruckeberg 3744. OREGON. Benton 
County: grassy summit of Marys Peak, Kruckeberg 2918. Multnomah County: Larch 
Mountain, Mrs. Mackaness, s.n.; Bonneville, Hitchcock sn. Clackamas County: ridge 
0.5 miles east of Devils Peak, Kruckeberg 4006. Ipano. Idaho County: Indian Hill 
lookout above Selway River, Kruckeberg 4095; granitic slopes above Canteen Mead- 
ows, Crags Mountains, Kruckeberg 4137; southeastern granitic rim of Crags Moun- 
tains, Kruckeberg 4140. Valley-Custer county line: rock crevices, Cape Horn Moun- 
tain, Kruckeberg 4177. Custer County: 2 miles east of Toxaway Lake, Sawtooth 
Mountains, Kruckeberg 4198; talus about Wild Horse Lakes, Mount Hyndman area, 


1960} KRUCKEBERG: SILENE 211 


Ur ed en Nee gin? CY 
ww ae 4 S&S 
<2 - oe 
MR a a vore Ce te 
Ly, 2 14 a o> 
oF “2 $v Kee 
a 0 13 Pr Sw] oe € ic Ve; 
> wale O% 
= 0 
Po - o% 3,°S : 
est Lo OSs “2a48 
x ee) Atey go FS 
We - “ ng — Oss 20 
0 eo 19 
wer 
oo aoe o£“ * ar 
8 oo A s 4 
Le ae ee 
no vt | \ 
2% © pO a tk *, 
eo” owe we 


28 

Fics. 13-28. Chromosomes of Silene microsporocytes. Fic. 13. S. thurberi, Diak. 
(Kruckeberg 3863). Fic. 14. S. williamsii, I M (Donnelly Dome, Gjaerevoll, s.n.). 
Fic. 15. S. antirrhina, Diak. (Palmer, s.n.). Fic. 16. S. caroliniana, I M (Channell, 
s.n.). Fic. 17. S. caroliniana wherryi, IM (Epstein, s.n.). Fic. 18. S. polypetala, Diak. 
(Galle, s.n.). Fic. 19. S. regia, Diak. (G. W. Carver National Monument, Palmer, s.n.). 
Fic. 20. S. rotundifolia, Diak. (Sherman, s.n.). Fic. 21. S. subciliata, Diak. (Dormon, 
s.n.). Fic. 22. Lychnis wilfordiu, 11 M (Alpine Garden Society 1280). Fic. 23. Silene 
keiskei, 1M (Mount Ho-o, Ozawa, s.n.). Fic. 24. S. repens latifolia, Diak. (Roberson, 
s.n.). Fic. 25. S. struthiloides, Diak. (Bryan, s.n.). Fic. 26. Silene sp., Diak. (Stainton 
Sykes & Williams 8108). Fic. 27. Silene sp., 1 M (Stainton Sykes & Williams 8178). 
Fic. 28. S. wrightii, I M, Kruckeberg 4716. Diak., diakinesis; I M, first metaphase; 
II M, second metaphase. 


212 MADRONO [Vol. 15 


Kruckeberg 4252. WasHincToN. Chelan County: Basalt ridge above Liberty, Krucke- 
berg A209; Icicle Creek, Hitchcock 20190. Okanogan County: between Twisp and 
Omak on Loup-Loup highway, Kruckeberg 3274; summit of Loup-Loup highway, 
Kruckeberg 3276; 5 miles west of Wauconda on highway to Republic, Kruckeberg 
3279. Skamania County: 1 mile southeast of Timberline Camp on Timberline Trail, 
Mount Saint Helens, Kruckeberg 3992. Stevens County: summit of Chewelah ski lift 
on road to Chewelah Peak, Kruckeberg 4052; between ski lift summit and Chewelah 
Peak, on open ridges, Kruckeberg 4057. 

SILENE GRAYI Wats. 2n=48. CALirornia. Trinity County: between Deer Creek 
basin and Deer Lake, Trinity Alps, Kruckeberg 3759; between Deer Creek Pass and 
Stonewall Pass, Trinity Alps, Kruckeberg 3766; 3.5 miles above Trinity Alps resort on 
Red Mountain trail below Stonewall Pass, Kruckeberg 3772. 

SILENE HOOKERI Nutt. ex T. & G. 2n=72. OrEGON. Benton County: between Kings 
Valley and Wren, Kruckeberg 2697. Douglas County: 5 miles up Little River, south 
of Glide, Cohen sn. Josephine County: peridotite alluvium bordering Whiskey Creek, 
3 miles west of Obrien, Aruckeberg 3777. 

SILENE INVISA H. & M. 2n=48. CALiForniA. Plumas County: rocky flats above 
Elwell Lake, Lakes Basin area, Kruckeberg 2897b; rocky swales above Upper 
Tamarack Lake, Sierra Buttes area, Kruckeberg 3694. Trinity County: wooded slope 
between Deer Creek basin and Deer Lake, Trinity Alps, Aruckeberg 3759a. 

SILENE LACINIATA Cav. subsp. Mayor H. & M. 2n=96. Catirornia. Los Angeles 
County: west of Beverly Glen Canyon on Mulholland Drive, Santa Monica Moun- 
tains, Snow s.n.; Turnbull Canyon, Whittier, Raven s.n. San Diego County: 0.5 mile 
east of Potrero Store along State Highway 94, Walters s.n. Santa Barbara County: 
Bishop Ranch, Clarke s.n. 

SILENE LACINIATA Cav. subsp. GREGGII (Gray) H. & M. 2n=48. Arizona. Apache 
County: at crossing of west fork Little Colorado River, between Greer and junction 
with Highway 73, Kruckeberg 4608. Cochise County: along Rustler Park road, Chiri- 
cahua Mountains, Kruckeberg 3862; rocky banks of Cave Creek, at Herb Martyr 
Forest Camp, Chiricahua Mountains, Kruckeberg 3863; between Rustler Park and 
Barfoot Lookout, Aruckeberg 4643; Crest Trail to Fly Peak, south of Rustler Park, 
Chiricahua Mountains, Aruckeberg 4650. Graham County: rocky slopes, 4 miles 
below Hospital Flats, Graham Mountains, Kruckeberg 3878. NEw Mexico. Catron 
County: ten miles northeast of Collins Park on road to Datil, Kruckeberg 4703. Grant 
County: rocky slopes overlooking copper mines at Santa Rita, Kruckeberg 4721. 
Texas. Brewster County: between Boot Springs and pass overlooking Inner Basin, 
Big Bend National Park, Kruckeberg 4776. 

SILENE LEMMONII Wats. 2n=48. CaLirorniA. Tulare County: above Lodgepole 
Camp on road to corral, Sequoia National Park, AKruckeberg 3393. Tuolumne 
County: grade east of Yosemite Creek along Tioga Pass Road, Kruckeberg 3530. 
Sierra County: along State Highway 89 at Calpine Lookout road, Kruckeberg 3667. 
Plumas County: woods at junction of Susanville and Lake Almanor roads, State 
Highway 89, Kruckeberg 3702. San Bernardino County: woods along road from 
South Fork of Santa Ana River to Barton Flats, Kruckeberg 3846. 

SILENE MARMORENSIS Kruck. 2n=48. CALrrorNIA. Siskiyou County: one mile 
north of Somes Bar on road to Camp Three, Hitchcock 20221, type. 

SILENE MENZIESII Hook. 2n=24. Diploid. CaLtrorntA. Fresno County: Jackass 
Meadows Forest Camp, below Florence Lake, Kruckeberg 3436. IpaHo. Custer 
County: 2 miles above Wild Horse Creek Forest Camp, Kruckeberg 4265. 

SILENE MENZIESII Hook. 2n=48. Tetraploid. OREGoN. Wallowa County: Evergreen 
Forest Camp, upper Imnaha River, Kruckeberg 3120. CALirorntA. Trinity County: 
upper end of Morris Meadows, Stuart Fork of Trinity River, Trinity Alps, Krucke- 
berg 3746. Ipauo. Valley County: lower Bear Valley at Poker Meadows Campground, 
Kruckeberg 4192. Custer County: Wild Horse Creek Forest Camp, Kruckeberg 4292. 
Bear Lake County: on tailings of phosphate mine, west slope of Snowdrift Mountain, 
Kruckeberg 4527. Custer County: 10 miles west of Challis, on talus of upper 


1960] KRUCKEBERG: SILENE 213 


Daugherty Gulch, Aruckeberg 4542. Wasutncton. Chelan County: 1 mile east of 
U.S. Highway 10 on State Highway 15C to Lake Wenatchee, Aruckeberg 3259. 
Kittitas County: along Swauk Creek, % mile below Liberty Guard Station, Krucke- 
berg 3010; base of Iron Peak trail, North Fork Teanaway River, Kruckeberg 3289. 
Okanogan County: rocky alluvium of Methow River, 2 miles below Mazama, Krucke- 
berg 3273; Sweat Creek Forest Camp, 7 miles west of Republic, Kruckeberg 3283. 
Stevens County: along trail between Calispell Meadows and Calispell Peak, Krucke- 
berg 4081. 

SILENE MONTANA Wats. 2n=48. CALIFORNIA. Tuolumne County: along Tioga Pass 
road between Tamarack Flats and Yosemite Creek, Kruckeberg 3529. Shasta County: 
between Chaos Jumbles and Noble Pass, Lassen Volcanic National Park, Kruckeberg 
3713. OREGON. Klamath County: slopes above Rim Drive, 3 miles above Park Head- 
quarters, Crater Lake National Park, Kruckeberg 3780. Nevapa. Ormsby County: 
along U.S. Highway 50, east of Spooner Summit, 10 miles west cf Carson City, 
Kruckeberg 3652. 

SILENA NUDA (Wats.) H. & M. 2n=48. CaLtrornia. Sierra County: 1 mile east of 
Calpine on road to Beckwourth, Kruckeberg 3664. Plumas County: 1 mile west of 
Portola along U.S. Highway 40, Kruckeberg 3669; 3 miles northwest of Lake Almanor 
along State Highway 89, Kruckeberg 3700. 

SILENE NUDA (Wats.) H. & M. subsp. rnsecTivora (Hend.) H. & M. 2n=48. 
Orecon. Klamath County: Sprague River valley, 7.5 miles west of Bly along State 
Highway 66, Kruckeberg 4030 (type locality). 

SILENE OCCIDENTALIS Wats. 2n=48. CALIFORNIA. Sierra County: timbered rocky 
flats between Sand Lake and Packer Lake, Sierra Buttes area, Kruckeberg 3696. 

SILENE OREGANA Wats. 2n=48. IpAnHo. Valley County: Lodgepole pine flats 5 miles 
west of McCall on highway to New Meadows, Kruckeberg 4151. 

SILENE PARISHIT Wats. 2n=48. CALIFORNIA. San Bernardino County: Grout Bay, 
Big Bear Lake, Everett 8248, Kruckeberg 3831; 2 miles above road’s end, along trail 
to Mount San Gorgonio, Aruckeberg 3848. 

SILENE PARRYI (Wats.) H. & M. 2n=48. Tetraploid. IpAHo. Bear Lake County: 
open, west-facing slopes of Snowdrift Mountain, Kruckeberg 4520. WASHINGTON. 
Chelan-Kittitas County line: Mount Lilian, eastern Wenatchee Mountains, Krucke- 
berg 3229, 3232. Stevens County: between head of ski lift and summit, Chewelah 
Peak road, Kruckeberg 4056; 34 mile below lookout, Chewelah Peak, Kruckeberg 
4064; southwest-facing slopes of Calispell Peak, Kruckeberg 4072; open ridge 34 mile 
south of Calispell Peak, Kruckeberg 4079. Montana. Flathead County: east slopes 
of Mount Aeneas, Swan River Range, Kruckeberg 4306. Glacier National Park: just 
south of Logan Pass, on rocky ledges of “Hanging Gardens,” Kruckeberg 4340. 
CanapaA. Waterton Lakes National Park, Alberta: east-facing slope above upper Car- 
thew Lake, on trail between Cameron Lake and Waterton Lake, Kruckeberg 43606. 

SILENE PARRYI (Wats.) H. & M. 2n=96. Octoploid. WAsHincTon. Clallam County 
(Olympic National Park): at Idaho Shelter, Hurricane Ridge, Kruckeberg 2776, 
4045; along ridge to Mount Angeles, Kruckeberg 2792; along trail to Moose Lake 
from Obstruction Point, Kruckeberg 4048. Kittitas County: alluvial flats at Fish 
Lake, upper Cle Elum River, Aruckeberg 3221; at base of Iron Peak trail, North 
Fork of Teanaway River, Kruckeberg 3288; along upper reaches of Miller Peak trail, 
Kruckeberg 39067. 

SILENE PETERSONIIT Maguire. 2n=96. Utan. Garfield County: on slopes of red 
talus and clay, 5.5 miles east of U.S. Highway 89, on Red Canyon road to Bryce 
Canyon National Park, Kruckeberg 3904. Iron County: limestone clay along west- 
facing rim of Cedar Breaks, Kruckeberg 3908. 

SILENE REPENS Patrin. subsp. AUSTRALE H. & M. 2n=24. Diploid. IpaHo. Custer 
County: Boulder Creek, 5 miles above Wildhorse Creek Canyon, Kruckeberg 4286a; 
Boulder Creek basin, 0.5 mile below Boulder Lake, Kruckeberg 4290. 

SILENE SARGENTII Wats. 2n=48. CALIFORNIA. Fresno County: pass between upper 
Bear Creek Meadow and Rose and Marie Lakes, Kruckeberg 3459; at Marie Lake, 


214 MADRONO [Vol. 15 


Kruckeberg 3482. Shasta County: near summit of Lassen Peak, Lassen Volcanic 
National Park, Kruckeberg 3711.3 

SILENE SCAPOSA Robins. 2n=-48. OreEGon. Harney County: 1 mile north of Squaw 
Butte on dirt road to U.S. Highway 20 (Burns-Bend highway), Kruckeberg 4031. 

SILENE SCOULERI Hook. 2n=48. IpaHo. Benewah County: between Tensed and 
Potlatch on U.S. Highway 95, Kruckeberg 4085. 

SILENE SPALDINGII Wats. 2n=48. WASHINGTON. Garfield County: near Colton, 
Daubenmire s.n. 

SILENE THURBERI Wats. 2n=48. Arizona. Cochise County stony alluvium along 
Cave Creek at Herb Martyr Forest Camp, Chiricahua Mountains, Kruckeberg 3863. 

SILENE VERECUNDA Wats. subsp. VERECUNDA 2n=48. CALIFORNIA. San Francisco 
County: rocky soil at east end of summit ridge of Mount Davidson, Raven s.n. 

SILENE VERECUNDA Wats. subsp. ANDERSONIT (Clokey) H. & M. 2n=48. NeEvapa. 
Clark County: rocky alluvium of Kyle Creek canyon, 1 mile below Charleston Park, 
Charleston Mountains, Kruckeberg 3915. 

SILENE VERECUNDA Wats. subsp. PLATYOTA (Wats.) H.& M. 2n=48. CaLIrornia. 
Tulare County: Last Chance Meadows area, Sierra Nevada Range, Kruckeberg 3345 ; 
7 miles above California Hot Springs on road to Johnsondale, Kruckeberg 3392; 5 
miles west of Hockett Meadows, Sequoia National Park, Kruckeberg 3422a. Los 
Angeles County: north slope of Mount Waterman, San Gabriel Mountains: Krucke- 
berg 3821; Horse Flats, San Gabriel Mountains, Kruckeberg 3822; north-facing slopes 
of Blue Ridge, San Gabriel Mountains, Kruckeberg 3916. San Bernardino County: 
5 miles below Lake Arrowhead on State Highway 18; flats above Moon Ridge, south 
side of Bear Valley on road to south fork Santa Ana River, Kruckeberg 3840 
(eglandular form) ; near summit of Sugarloaf Ridge, 6 miles southeast of Bear Valley, 
Kruckeberg 3843 (eglandular form). Riverside County: 3.4 miles south and east of 
State Highway 74, Bautista Canyon, Vasek s.n. 

SILENE WILLIAMSI! Britt. 2n=24. ALAsKa. Slopes of Shaw Creek, Richardson 
Highway, Gjaerevoll 1444; heath, Donnelly Dome, Alaska Range, Gjaerevoll 1284. 

SILENE WRIGHTII Gray. 2n=96. NEw Mexico. Grant County: in crevices of mas- 
sive boulders at base of sheer rock cliffs, overlooking copper mines at Santa Rita, 
Kruckeberg 47106. 


EASTERN NorTH AMERICAN SPECIES 


SILENE ANTIRRHINA L. 2n=24. Missourr. Jasper County: near Webb City, 
Palmer s.n. 

SILENE CAROLINIANA Walt. 2n=48. NortH Carorina. Franklin County: 9 miles 
east of Wake Forest, granitic “flat-rock” area, Channell s.n. Orange County: Chapel 
Hill, Bell s.n. 

SILENE CAROLINIANA Walt. subsp. WHERRYI (Small) Clausen. 2n=48. Garden 
culture: Larchmont, New York, Epsein s.n. 

SILENE POLYPETALA (Walt.) Fern. and Schub. 2n=48. Gerorcia. Talbot County: 
north of county bridge, hillside above Flint River, Galle s.n. 

SILENE REGIA Sims. 2n=48. Missourr. Dade County: near South Greenfield, 
Palmer sm. Newton County: George Washington Carver National Monument, 
Palmer s.n. 

SILENE ROTUNDIFOLIA Nutt. 2n=48. TENNESSEE. Marion County: crevices of the 
“Chimneys,” gorge of Pocket Creek, Whitwell Pocket area, Sherman s.n. 

SILENE STELLATA (L.) Ait. [including S. scabrella (Nieuwl.) Palm. and Steyerm. | 
2n=48. InpiAna. Starke County: Jackson s.n. Missourt. Newton County: near Dia- 
mond, George Washington Carver National Monument, Palmer s.n. Kansas. Riley 
County: 40 miles east of Aurora, woods along Fancy Creek, Fraser. WEST VIRGINIA. 
Monongalia County: on Permian shale banks, vicinity of Morgantown, Constable & 
Core s.n. LOUISIANA. northwestern portion of state, Dormon s.n. 


3 Plants intermediate between S. sargentii and S. Suksdorfii Robins. 


1960] TORRES: ZINNIA IAS 


SILENE SUBCILIATA Robins. 2n=48. Louisiana. “Western Louisiana,” Dormon s.n. 
(garden culture). 

SILENE VIRGINICA L. 2n=48. NortTH Carotina. Wake County: 18 miles north of 
Raleigh, across the Neuse River, on State Highway 50, Smith s.n. 


SPECIES OUTSIDE CONTINENTAL NORTH AMERICA 


LYCHNIS WILForDII Maxim. 2n=24. Japan: Garden culture, MW. Ozawa s.n.; garden 
culture, Alpine Garden Society 1280. 

PETROCOPTIS PYRENAICA Braun. 2n==24. Europe. Garden culture, Museum of 
Natural History, Paris s.n. 

SILENE KEISKEI Miq. 2n=24. Japan: Mt. Ho-o, M. Ozawa s.n.; Mt. Kitadake, 
Osawa s.n. 

SILENE REPENS Patrin. var. LATIFOLIA Turcz. 2n=48. JAPAN: Garden culture, Mrs. 
L.N. Roberson s.n.; garden culture, Epstein s.n. 

SILENE STRUTHIOLOIDES Gray. 2n=24. Hawai. Hawaii: near the Kilauea Crater, 
Hawaii National Park, Bryan s.n. 

SILENE species. 2n=24. NEPAL: Tegar, north of Mustang, Sykes & Williams 8108 ; 
Larjung, south of Tukucha, Kali Gandaki Valley, Sykes & Williams 8178. 

Department of Botany, 
University of Washington, Seattle 


LITERATURE CITED 


Hitcucock, C. L., and B. Macutre. 1947. A revision of the North American species 
of Silene. Univ. Wash. Publ. Biol. 13:1-73. 
KRUCKEBERG, A. R. 1954. Chromosome numbers in Silene (Caryophyllaceae): I. 
Madronfo 12:238-246. 
. 1955. Interspecific hybridizations of Silene. Am. Jour. Bot. 42:373-378. 
———§—., 1960. A new Silene from northwestern California. Madrofio 15:172-177. 


A NEW SPECIES OF ZINNIA FROM MEXICO 
A. M. Torres 


During the course of a cytotaxonomic study of the genus Zinnia (Com- 
positae), plants started from seeds kindly provided by Dr. Jerzy Rzedow- 
ski of the Universidad Autonoma de San Luis Potosi, México, were cul- 
tivated in the greenhouses of Indiana University. One collection, when 
grown to maturity, proved to be a new species known thus far only from 
the area where the seeds were collected. 

Zinnia citrea sp. nov. Planta perennis, cespitosa, ad 2 dm. alta; cauli- 
bus viridibus, strigosis; follis oppositis, amplexicaulibus, uninervis, 
linearibus, ad 3.5 cm. longis, 0.8-1.9 mm. latis, sparse strigosis aut 
glabrescentibus, sparse glanduloso-punctatis; capitulis terminatibus in 
pedunculis 0.8—2.0 cm. longis, subhemisphaericis, 0.4 cm. latis 0.5 cm. 
altis; phyllariis oblongis, firme-gradatis, herbaceis, minuto-glanduliferis, 
apicibus obtusis, ciliatis; radiis ca. 7, chloreis oblongis, ad 0.8 cm. longis 
0.5 cm. latis, sine tubo, in dorso viridis nervis, apicibus 0.3 lobis; achaeniis 
radiorum oblanceolatis, ad 4.2 mm. longis, tuberculatis, nigrescentibus, 
sine aristis; floribus disci ca. 22, tubis 3.1 mm. longis, lobis 1.4 mm. 


216 MADRONO [Vol. 15 


longis, intus lobis lanuginosis, supra flavo-viridibus, infra viridibus; 
achaeniis disci oblanceolatis, compressis, ad 2.6 mm. longis, sparse 
ciliatis vel vere glabris, marginibus ciliatis, pappis 2 inaequalis aristatis; 
receptaculi paleis lanceolatis, scariosis, apicibus acutis minuto-dentatis, 
plus minusve viridibus; receptaculis convexis. 

Plants perennial, cespitose, about 2 dm. high; stems green, strigose; 
leaves opposite, sheathing, one-nerved, linear, about 3.5 cm. long, 0.8—-1.9 
mm. wide, sparsely strigose or becoming glabrous, attenuate, sparsely 
glandular-punctate; heads terminal on peduncles 0.8—2.0 cm. long, sub- 
hemispherical, 0.4 cm. wide, 0.5 cm. high; phyllaries oblong, strongly 
graduated, herbaceous, apices obtuse, ciliate, minutely glandular; rays 
about 7, lemon-colored, oblong, about 0.8 cm. long, 0.5 cm. wide, tube- 
less, green-nerved on the back, apices O—3 lobed; ray achenes oblance- 
olate, about 4.2 mm. long, tuberculate, becoming black, awnless; disk 
flowers about 22, the tube 3.1 mm. long, the lobes 1.4 mm. long, velvety 
on inner surface, yellow-green above, green below; achenes of the disk 
oblanceolate, compressed, 2.6 mm. long, sparsely ciliate or essentially 
glabrous, the margins ciliate, the pappus of 2 unequal awns; pales of the 
receptacle lanceolate, scarious, the apices acute, minutely dentate, more 
or less green; receptacle convex. 

Type. Mexico. Seed collected near Santo Domingo, municipality of 
Guadalcazar, San Luis Potosi, on deep alluvial soil with Prosopis. Elev. 
1200 m. Cultivated in greenhouse, Indiana University, 1959, Torres 139 
(IND). 

Zinnia citrea is very Closely allied morphologically, but entirely dis- 
tinct from Z. grandiflora Nutt. and Z. acerosa (DC.) Gray (considered 
by the writer to include Z. pumila Gray). Zinnia grandiflora occurs in 
the southwestern United States as far north as Kansas, and in the Mexican 
states of Sonora, Chihuahua, Coahuila and Durango. Zinnia acerosa is 
found in the southern parts of Arizona and New Mexico, western Texas, 
and the adjacent portions of México as far south as Durango, Zacatecas 
and San Luis Potosi. 

Some of the differences between the three species are indicated below. 


Z. citrea Z. acerosa Z. grandiflora 
Number of leaf nerves 1 1 2 
Ray color lemon white yellow 
Number of rays 5—/ 4-6 3-6 
Number of disk flowers 18-25 8-13 18-24 
n=10 nic 721 
Chromosome no. no==20 n= 19 247 
n= 20 


Compared with Z. citrea, the rays of Z. grandiflora are yellow, usually 
fewer and considerably larger; the disk is red or sometimes green, instead 
of yellowish; the leaves are wider and longer but quite variable; the 
somatic chromosome number is 42. Jackson (1959) has reported a 
gametic chromosome number is 21 and the somatic chromosome number 


1960 | WAGENER: PINUS 217 


is 42. Jackson (1959) has reported a gametic chromosome number of 24 
for Z. grandiflora. The rays of Z. acerosa are white (drying pale yellow), 
generally fewer, slightly larger and the disk is reddish. Populations hav- 
ing gametic chromosome numbers of 10, 19, and 20 have been found. 
Voucher herbarium specimens for the chromosome counts are deposited 
at Indiana University. 


Department of Botany 
Indiana University 
Bloomington, Indiana 


LITERATURE CITED 
Jackson, R.C. 1959. Documented chromosome numbers of plants. Madronfo 15:52. 


A COMMENT ON COLD SUSCEPTIBILITY OF PONDEROSA 
AND JEFFREY PINES 


WItiis W. WAGENER! 


Dr. Haller’s recent paper® on factors affecting the distribution of 
ponderosa and Jeffrey pines prompts a supplementary note concerning 
the comparative effects of low temperatures observed on the two species 
in northeastern California. In this region extensive mixed stands of the 
two occur, many of them above the 5,000-foot level, providing a con- 
venient comparison of their reactions to environmental conditions for the 
geographical races represented there. 

Dr. Haller considers that Pinus jeffrevi is more tolerant than P. pon- 
derosa of extremes of low temperature and aridity, but he concludes that 
the differential limiting effect of low temperature must be exerted in the 
seedling stage or on young trees because mature trees of P. ponderosa at 
its upper altitudinal limit appear vigorous and show no evidence of stunt- 
ing. My observations over the past 25 years, following periods of severe 
cold, fail to indicate any material difference between the two species in 
their ability to withstand extreme cold, either as young or mature trees. 

In January 1937, California experienced two very cold periods, par- 
ticularly east of the Sierra Nevada crest. The first of these was from 
January 8 to 10 and the second from January 20 to 25. The lowest tem- 
perature reported to the United States Weather Bureau for these periods 
in California was —45°F. at Boca, California, on January 20. 

Early in February a belt of pronounced damage to pines and other 
vegetation became noticeable along the east face of the Sierra Nevada 
for almost its entire length. At the north end it was narrow, from 25 to 


1 Forest Pathologist, Pacific Southwest Forest and Range Experiment Station, 
Forest Service, U.S. Department of Agriculture. 


2 Haller, John R. Factors affecting the distribution of ponderosa and Jeffrey pines 
in California. Madrono 15:65-71. 1959. 


218 MADRONO [Vol. 15 


200 yards wide, depending on slope, and occurred at an elevation of about 
4,800 feet. Toward the south the belt was wider and was located at a 
gradually increasing elevation, between 7,000 and 8,000 feet at the south 
end. 

From the Truckee River southward the damage to pines consisted 
largely of foliage browning, from which practically all the trees later 
recovered. On the slope above Long Valley Creek, Lassen County, how- 
ever, damage was much more pronounced, and many mature pines, as 
well as trees of younger ages, failed to recover. Mortality in the pines 
ran as high as 75 percent. 

In the topographic gap created by the Susan River the belt-like char- 
acter of the zone of damage was less evident but the major injury was 
found at approximately the same elevation as above Long Valley Creek. 
A strong temperature inversion apparently occurred at this level. On low 
benches along the Susan River a number of cases were observed where 
the lower foliage of young Jeffrey pines 20 to 30 feet in height was almost 
completely killed, whereas that of the upper crowns on the same trees 
was only slightly damaged. In addition, many of the trees showed brown- 
ing of the phloem tissues of the inner bark of the lower trunks and 
branches. Most of these trees died later. The immediate cause of death 
appeared to be the activities of bark insects attracted by the killed phloem. 
Whether the trees could have survived in the absence of the insects was 
not determined. 

In Jeffrey pine the spread between noticeable injury to foliage and the 
complete killing of foliage and branches or trunks appeared to be very 
much narrower than in ponderosa pine. Cases were noted in which foliage 
on ponderosa pines was almost completely killed, but nearly all buds and 
twigs remained alive and produced fairly good growth the following sea- 
son. This was not true for Jeffrey pines. In them, severe foliage injury 
was accompanied by severe bark injury, and no recovery followed. Where 
both species were present in the same location there was less mortality in 
the ponderosa than in the Jeffrey pines. 

Another series of abnormally low temperatures prevailed in California 
in January, 1949. Minima of —38°F. were registered at Boca, California, 
on the 25th and of —31°F. at Alturas and Bridgeport Dam on the 26th. 

Damage to pines on low ground east of the Sierra Nevada as a result 
of this series of cold waves during the month was much more general in 
occurrence than in 1937. Heavy browning of foliage of young ponderosa 
and Jeffrey pines on the borders of low flats or along stream bottoms was 
noted in interior basin drainages from Modoc County to the San Bernar- 
dino Mountains. Stringers of pines along stream courses extending into 
the Honey Lake and Carson valleys were heavily damaged, and many 
trees eighteen to twenty inches in diameter breast high later succumbed. 
These were mostly Jeffrey pines but some ponderosa pines were present 
also. Mortality in them appeared to be no greater in proportion than in 
the Jeffrey. Recovery in young trees appeared to depend on whether or 


1960 | DOCUMENTED CHROMOSOME NUMBERS 219 


not the inner bark tissues of the main stem had been injured. Killing of 
these tissues was found both in ponderosa and Jeffrey pines, with later 
mortality after the stems had been invaded by bark-feeding insects. 

Neither in 1937 nor in 1949 was enough difference in mortality from 
the abnormal cold noted between established trees of the two species to 
account for the tendency of Jeffrey pine to be confined to the higher ele- 
vations or colder situations. If an ecologically significant difference exists 
between the two with respect to cold tolerance it must be operative in the 
seedling stage, as Haller suggests, or through some other influence than 
differential mortality from cold. 


DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 
(See MaproNo 9:257-258. 1948) 


SPECIES NUMBER COUNTED BY COLLECTION LOCALITY 
RANUNCULACEAE 
Delphinium i298 B. L. Turner, Thompson & Hardin County, 
carolinianum Walter TEX’ Turner 96 Texas 
TEX 
virescens Var. 
macroceratilis (Rydb.) n= 8 B. L. Turner, Turner 4395 Bexar County, 
Ewan TEX TEX Texas 
SAXIFRAGACEAE 
Saxifraga n—19 K. I. Beamish, Beamish 7828 Mt. Seymour near 
ferruginea UBC UBC Vancouver, B.C., 
Graham Canada 
Beamish 9000 Caulfeilds, near 
UBC Vancouver, B.C., 
Canada 
integrifolia 1-19 K. I. Beamish, Beamish 7057 Thetis Lake, 
Hook. UBC UBC Vancouver Island, 
B.C., Canada 
Beamish, Elk Falls, Van- 
Vrugtman & couver Island, 
Sparling 8017 B.C., Canada 
UBC 
montanensis n—19 K. I. Beamish, Vrugtman, Princeton-Merritt 
Small UBC Beamish & Road, B.C., 
Kallio 9027 Canada 
UBC 
occidentalis n= 19 K.I. Beamish, (Beamish, Botanie Valley, 
Watson sensu lat. UBC Vrugtman & near Lytton, 
Sperrings B.C., Canada 
8224, UBC 


1 Symbols for institutions are those listed by Lanjouw and Stafleu. Index Herbari- 
orum, Part I. Second edition, 1954, Utrecht. 


220 


SPECIES 


NUMBER 


MADRONO 


COUNTED BY 


COLLECTION 


[Vol. 15 


LOCALITY 


Saxifraga 
tolmiei 


D&G: 


UMBELLIFERAE 
Pseudotaenidia 
montana Mackenz. 


GARRY ACEAE 
Garrya lindheimeri 
Torr. 


VERBENACEAE 
Verbena 
hastata L. 


bracteata 
Lag. & Rodr. 


stricta Vent. 


urticifolia L. 


SCROPHULARIACEAE 
Penstemon clutei 
A. Nels. 


COMPOSITAE 
Blennos perma nanum 
(Hook.) Blake 
var. robustum 
Howell 


Encelia frutescens 
Gray 


Haplopappus 
havardi Waterfall 


phylloce phallus DC. 
subsp. phyllo- 
cephalus 


pbhyllocephalus 


subsp. annuus (Rydb. 
Hall 


spinulosus 
(Pursh) DC. 
subsp. spinulosus 


spinulosus 
(Pursh) DC. 
subsp. spinulosus 


n=A15 
2n = 22 
n= 11 
nei 
itesecet| 
a7 
n= / 
n=8 
n=/7 
tale 
n= 4 
n=6 
n=6 
n= 4+1 
n = 4-+2 


K. I. Beamish, 
UBC 


R. L. Guthrie 
WVA 


B. L. Turner, 
TEX 


J.D. Poindexter, 
KANU 


R. C. Jackson, 
KANU 


J. D. Poindexter, 
KANU 


J. D. Poindexter, 
KANU 


R. C. Jackson, 
KANU 


R. Ornduff, 
UC 


R. C. Jackson, 
KANU 


R. C. Jackson, 
KANU 


R. C. Jackson, 
KANU 


R. C. Jackson, 
KANU 


R. C. Jackson, 
KANU 


R. C. Jackson, 
KANU 


Beamish 7831 
UBC 


Guthrie 
s.n. UC 


Turner 3967 
TEX 


Poindexter 33 
KANU 


Poindexter 18 
KANU 

Poindexter 37 
KANU 


Poindexter 9 
KANU 


Ren 256s Ve 
Jackson 2683-1 
KANU 


Ornduff 4963 
UC 

Jackson 2684 
KANU 


Jackson 2717 
KANU 


Jackson 2610 
KANU 


Ungar 729 
KANU 


Jackson 2455-1 
KANU 


Jackson 2455-14 
KANU 


Mt. Seymour near 
Vancouver, B.C., 
Canada 


Greenbrier County. 
W. Va. 


Austin, 
Travis County, 
Texas 


Douglas County, 
Kansas 


Trego County, 
Kansas 


Douglas County, 
Kansas 


Douglas County, 
Kansas 


Coconino 
County, 
Arizona 


Pt. Reyes, 
Marin County, 
California 


Coconino 
County, 
Arizona 


Lea County, 
New Mexico 


Cameron 
County, 
Texas 


Stafford 
County, 
Kansas 


Socorro 
County, 
New Mexico 


Socorro 
County, 
New Mexico 


1960] 


REVIEW 


SPECIES NUMBER COUNTED BY COLLECTION LOCALITY 
Machaeranthera 28 R. C. Jackson, (Jackson 2901 Durango, 
blephariphylla KANU KANU Mexico 
(Gray) Shinners 
tagetina Greene ni 4 R. C. Jackson, |Jackson 2600 Hidalgo 
KANU KANU County, 
New Mexico 
Porophyllum i ese 1 R. C. Jackson, |R.C.& S.W.  |Otero County, 
scoparium Gray KANU Jackson 2701 New Mexico 
KANU 
Psilostro phe cooperi seal (6) R. C. Jackson, |R.C. & S.W. Yavapai 
(Gray) Greene KANU Jackson 2693 County, 
KANU Arizona 
Sclerocar pus uniserialis M12 B. L. Turner, Turner et al. Jackson 
(Hook.) Benth. & TEX 3313 TEX County, 
Hook. f. Texas 
Viguzera 
adeno phylla Blake Ve C.B. Heiser, Stoutamire North of San 
IND 2813 IND Luis Potosi- 
Nuevo Leon 
state line, 
Mexico 
deltoidea var. Parishii n= 18 C. B. Heiser, Neher in 1958 |Near Palm 
(Greene) Vasey & Rose IND IND Springs, River- 
side County, 
California 
dentata var. nH ay B.L. Turner, Turner 4463B — |Austin, Travis 
brevipes (DC.) TEX TEX County, Texas 
Blake 
porteri (A. Gray) n=17 |C.B. Heiser, From seed DeKalb 
Blake IND (Duncan) County, 
Heiser 4561 Georgia 
IND 
stenoloba Blake iis =aley, C. B. Heiser, From seed Eddy County, 
IND Tucker 3131 New Mexico 
REVIEW 


Carex—Its Distribution and Importance in Utah. By Mont E. Lewis. Brigham 
Young University Science Bulletin, Biological Series 1(2):1-43. 1958. $1.00. 

“The purpose of this report is to bring available information concerning the Carex 
species in Utah up to date.’’ With these modest words Mr. Lewis of the United States 
Forest Service intermountain regional office in Ogden, Utah, introduces his excellent 
and original study of the identification, distribution, ecology, and grazing values of 


Utah carices. 


Since the only Utah flora, namely Tidestrom’s, is now over a third of a century 
old, a modern study such as this is most welcome. It is doubly welcome in that it 
comes from a representative of the federal organization which spends more man-hours 
interpreting the native plant cover of Utah than any other group. It is gratifying 
that a man primarily concerned with administration should take the time to produce 


a work such as this. 


Dee MADRONO [Vol. 15 


The report contains a diagrammatic summary of the mountain physiographic 
provinces of Utah and of their zonal belts of vegetation. Most interesting floristic, 
vegetational, and ecological problems are evident from the scattered occurrence of 
Pinus ponderosa, a local intrusion of Pinus contorta into the state, a varying amount 
of pinyon-juniper vegetation in different provinces, a constantly present but sup- 
posedly seral aspen-fir belt, and scattered alpine vegetation. Descriptions as well as 
explanations of most of these phenomena are still lacking in the botanical literature. 
For subsequent editions we hope Mr. Lewis will find time and opportunity to prepare 
altitude scales to this zonal vegetation diagram and to characterize the plant com- 
munities and floras concerned. 

Although the physiographic units used certainly make far more sense for describ- 
ing the distributions of plants within Utah than do county units, floristic units would 
be best of all. Do the physiographic divisions of Utah coincide with floristic divisons ? 
For those unfamiliar with floras using floristically defined areas to describe plant dis- 
tributions, reference may be made to the “Flora of the USSR” [cf. Stearn’s paper in 
the New Phytologist 46:61-87, 1947. Note that a new map appears in Volume 18 
(1952) of the Flora USSR] and to recent local floras such as those of Tadzhikstan, 
Kazakstan, Central Siberia, Leningrad and Murmansk regions, etc. Hylander’s new 
Scandinavian flora and the new Greenland flora employ similar schemes. 

Illustrations of named Carex species explain the terminology used in the keys. 
These keys do not exactly duplicate Mackenzie’s and are easier to use for identifica- 
tion to section and to species. They require ripe perigynia; so do most Carex keys. 
For this reason A. Neumann’s key to the carices of northwest Germanv in vegetative 
condition deserves mention (Mitteil. Floristischsoziologischen Arbeitsgemeinschaft 
3:44-77, 1952). Unfortunately, of the 126 taxa keyed by Neumann, only 6 are found 
among Lewis’ 102 taxa. The species descriptions are pertinent and comparable; dis- 
tinguishing features are frequently mentioned, and here the author uses his field 
experience to great advantage. 

Distribution statements are short but probably adequate, with the Utah areas 
given in as full detail as present knowledge permits. This present knowledge is so poor 
as to make a phytogeographer weep, but one result of Lewis’ work will be a rapid 
advance in our knowledge of where various carices do grow. The ecological data are 
excellent and far better than anything else available. Writers of extra-Utah floras can 
most profitably use these data for comparisons. Finally, all interested in range man- 
agement will treasure Lewis’ unique notes on forage values. 

The monograph closes with an excellent list of local references, a glossary, an index, 
and a table summarizing for all species their distribution by physiographic provinces, 
the mountain belt of vegetation in which they occur, their habitat within this belt, 
their abundance, and geographical distribution—Jack Mayor, Botany Department, 
University of California, Davis. 


NOTES AND NEWS 


Epwarp LEE GREENE CORRESPONDENCE—Persons interested in the botanical history 
of the western United States may find material of value in the correspondence of 
Edward Lee Greene now in the archives of the University of Notre Dame. This file 
includes letters to Greene over a period of forty or more years of his botanical career. 
These range from single letters commenting on specimens or requesting information 
to extensive correspondence from many of the notable figures in botany. This 
material is available to scholars able to visit the University of Notre Dame. Also 
microfilm copies of letters can be supplied at about four cents per picture and photo- 
stat copies at about forty cents per copy. 

The following list includes the names of most correspondents whose letters are 
in the Greene files at Notre Dame. It is not necessarily complete. Some of the 


1960] 


NOTES AND NEWS 


223 


material is not alphabetically filed and no catalogue is available at present. To 
indicate the approximate extent of the correspondence from individuals the following 


scale is used: 


ADQWS 


. one or two letters 

. approximately five 

. approximately ten 

. approximately twenty five 

. extensive, usually fifty or more 


These designations are based on estimates and not on actual counts, and they do 
not purport to indicate the intrinsic interest or importance of the Jetters. 


Abrams, L. R. 
Alderson, R. D. 
Allen, A. A. 
Allison, E. M. 
Ames, O. 
Anderson, C. L. 
Andres, H. 
Andrews, L. 
Arber, Agnes 
Arthur, J. C. 
Atkinson, G. F. 
Austin, R. M. 
Autran, E. 


Bailey, L. H. 
Bailey, W. W. 
Baillon, L. 
Baker, C. F. 
Baker, M. 
Balfour, I. B. 
Ball, John 
Bancroft, F.& R. 
Barnes, W. 
Barnhart, J. H. 
Bartlett, H. H. 
Batalin, A. 
Bates, J. M. 
Baxter, M. S. 
Beal, W. J. 
Bebb, M.S. 
Becker, W. 
Beguinot, A. 
Bell, A. A. 
Benton, L. E. 
Bernard, H. 
Bessey, E. A. 
Bessey, C. E. 
Best, G. 
Bethel, E. 
Bicknell, E. P. 
Bioletti, F. 
Bissell, C. H. 
Blake, A. 
Blankinship, J. W. 
Blochman, J. 
Blumer, J. C. 
Boardman, S. L. 
Bolander, H. N. 
Bonté, J. H. 


FOPFEPPOPRSE SE ErRyS 


DWePrOOSrrBISrOrr rrr r errr arorrerrrmesny 


Brainerd, E. 
Bramton, E. 
Brand, Dr. 


Brandegee, K. & T.S. 


Brinton, J. B. 
Briquet, J. 
Britten, J. 
Britton, N. L. 
Brown, C. E. 
Brown, J. S. 
Bruckett, G. 
Buchanan, W. D. 
Buckenau, F. 
Budworth, G. 
Bull, H. R. 


Burgess, E. S. & T. J. 


Burglehaus, F. H. 
Burkill, I. H. 
Burrall, H. D. 
Buscalioni, L. 
Bush, B. F. 
Butler, F. & G. 
Buysman, M. 


Campbell, D. H. 
Canby, W. M. 
Candolle de, Alp. 
Case, J. H. 
Chase, A. 
Choate, H. 
Clarke, F. L. 
Claybrooke, G. 
Cleveland, D. 


Cockerell, T. D. A. 


Cocks, R. S. 
Cole, L. B. 
Conard, H. S. 
Cooper, A. J. 
Copeland, E. B. 
Coulter, J. M. 
Coville, F. V. 
Crawford, J. 
Cross, E. R. 
Curtis, G. P. 
Cusick, Wm. C. 


Davidson, A. 
Davidson, G. 
Davidson, J. 


Drm DWW SUNOS SSS LOOS SESS SS SPLUPSPSSSPSSPS Sere PUPSee weed 


Davy, J. B. 
Day, D. F. 
Dayton, W. A. 
Deam, C. C. 
Deane, W. 
Delafield, M. 
Derby, G. 
Dewey, L. H. 
Dickson, J. M. 
Dodge, C. K. 
Donnelly, H. B. 
Dudley, W. K. 
Duggan, J. 
Dupret, H. 
Dyer, W. 


Barle, F, S. 
Eastwood, A. 
Eaton, A. A. 
Egerton, J. B. 
Eggert, H. 


Eggleston, W. W. 


Elliot, W. 
Ellis, J. B. 
Elmer, A. D. E. 
Engelmann, Geo. 


Evermann, B. W. 


Farr, E. M. 
Farwell, O. A. 
Fawcett, Wm. 
Fedde, F. 
Fernald, M. L. 
Fischer, T. 
Fitzpatrick, T. J. 
Fletcher, J. 
Friedlander, R. 
Fronanl, E. 


Gager, C. S. 
Ganong, W. 
Garrett, A. O. 
Gates, F. C. 
Gautier, G. 
Gerard, J. N. 
Gibbons, G. 
Gibson, A. 
Gormann, M. W. 
Grant, G. B. 


QF SPOS rrr yS> SWOPE BAPUWD SMW SS PPP OyY PrrrroUrrersuUruxwy 


224 


Gray, A. 


Greenman, J. M. 


Gregorson, D. 
Gregory, E. 
Grimes, E. J. 
Grose, I. F. 
Grout, A. J. 
Guillou, A. 


Haberer, J. V. 
Harger, E. B. 
Harper, R. M. 
Hasse, H. E. 
Hastings, G. 
Havard, V. 
Hedrick, U. P. 
Heimer, I. A. 
Heller, A. A. 
Henderson, L. 
Henry, J. K. 
Hensen, G. 
Hill, E. f. 


Hillegass, G. W. 


Hine, J.S. 
Hitchcock, A. S. 
Holden, E. S. 
Hollick, A. 
Holm, T. 
Holmes, J. A. 


Holzinger, J. W. 


House, H. D. 
Howe, M. A. 
Howell, T. 
Huth, E. 


Jaggar, A. L. 
Jennings, O. E. 
Jepson, W. 
Johnson, R. H. 
Jones, M. E. 
Jordan, D. S. 


Kearney, T. H. 
Keller, R. 
Kelsey, F. D. 
Kennedy, P. B. 
Kneipp, L. F. 
Knight, O. W. 


PEEP re SEPT S Se SET USS SSE ESS SSOPUSPS SS SPP ye SEEPS ryrnr 


MADRONO 


Knuth, R. 
Koehne, E. 
Krautter, L. 
Kreke, M. 
Kumlein, L. 
Kunze, O. 


Langlois, A. B. 
Leiberg, J. B. 
Lemmon, J. G. 
Limberger, W. B. 
Lloyd, F. E. 
Lunell, J. 

Lyon, W. D. 
Lyon, W. S. 


MacDougal, D. 
Macfarlane, J. M. 
Mackenzie, K. K. 
MacMillan, C. 
Macoun, John 
Maxwell, C. F. 
Metcalfe, O. B. 
Millspaugh, C. F. 
Mohr, C. 
Morong, T. 
Moseley, E. L. 
Moyer, L. R. 
Mueller, Barron F.von 
Mulford, A. J. 


Nelson, A. 
Nelson, E. 
Nieuwland, J. 


Ogden, H. V. 
Olney, S. C. 
Orcutt, C. R. 
Osterhoutt, G. E. 


Parish, S. B. 
Parry, CeC, 
Patterson, H. N. 
Paull, C. F. 
Pearson, G. A. 
Peekinpah, L. A. R. 
Pennell, F. W. 
Phelps, O. P. 


OP rPrmnnmOQ WWOF CFU SSS SPSSOOS MESS SPWOSFSESAYS PPYrrpPyY 


[Vol. 15 


Piper, C. V. 
Pollard, C. L. 
Pond, C. F. 
Porter, C. E. 
Porter, T. C. 
Price, C. G. 
Pringle, C. G. 


Reed, i, 
Robinson, B. L. 
Rose, J. W. 
Rosendahl, C. O. 
Rusby, H. H. 
Ruth, A. 
Rydberg, P. A. 


Sargeant, C. S. 
Sears. J. Ely 
Shea, CG. 1. 
Sheldon, E. P. 
Sherff, E. E. 
Siegfried, J. P. 
Small, J. K. 
Smith, C. P. 
Smith, J. D. 
Sonne, C. F. 
Stapf, O. 
Stone, Wilbur 
Suksdorf, W. N. 


Thornber, J. J. 
Tracy, J. P. 
Trelease, Wm. 
Turner, J. H. 


Underwood, L. M. 
Urban, J. 


Vasey, Geo. 


Watson, Lawrence 
Watson, Sereno 
Wellington, R. 
Weigand, K. M. 
Wilder, C. M. 
Wilson, N. G. 
Wooton, E. O. 
Wright, W. G. 


POPU rr mm DW WW PUOWS> OFSFRPSPOSerrerrn BWTMQ0CNF BHO 


The undersigned will be happy to supply further information or to try to locate 
correspondence from individuals not included in this list. RoBert P. McInTosu, 
Curator, Greene-Nieuwland Herbarium, University of Notre Dame, Notre Dame, 


Indiana. 


UROSPERMUM PICROIDES (L.) SCHMIDT IN BERKELEY. This cichoriacecus Medi- 
terranean plant has apparently become established on a part of the University of 
California campus which has not been intensively landscaped (Hall in 1915, Mason 
in 1943, Carter 4109 in 1960). Urospermum picroides may be readily recognized by 
its flat tuberculate achenes with long slender beaks, the ampule-like bases of which 
are enlarged to a diameter greater than that of the achenes—ANNETTA CARTER, 


Department of Botany, University of California, Berkeley. 


INFORMATION FOR CONTRIBUTORS 


- Manuscripts submitted for publication should not exceed an estimated 
20 pages when printed unless the author agree to bear the cost of the ad- 
ditional pages at the rate of $20 per page. Illustrative materials (includ- 
ing “typographically difficult” matter) in excess of 30 per cent for papers 
up to 10 pages and 20 per cent for longer papers are chargeable to the 
author. Subject to the approval of the Editorial Board, manuscripts may 
be published ahead of schedule, as additional pages to an issue, provided 
the author assume the complete cost of publication. . 

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

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


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


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume I, 1916-1929 . . . . . . . $5.00 
Volume IT, 1930-1934 . . ... . 5.00 
Volume ITI, 1935-1936 . . . . . . = 5.00 
Volume IV, 1937-1938 . .... . 5.00 
Volume V, 1939-1940 . . ... . 5.00 
Volume VI, 1941-1942 . .... . 5.00 
Volume VII, 1943-1944 . . . . . . 5.00 
Volume VIII, 1945-1946 .... . 5.00 
Volume IX, 1947-1948 . . . . . . 6.00 
Volume X, 1949-1950... . . . 7.00 
Volume XI, 1951-1952. . . . . . 8.00 
Volume XII, 1953-1954 . . . . . . 8.00 
Volume XIII, 1955-1956... . . 8.00 
Volume XIV, 1957-1958 ... . . 8.00 
Single numbers, 

Volumes land: 2. “... “3 2 ate 50 
Succeeding numbers, 

(except Volume XI, No.2) . . . . 1.00 

Volume XI,No.2 ...... . ~~ 1.50 


The subscription price of MapRONo is $6.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


io7TaGN g 


MADRONO 


VOLUME 15, NUMBER 8 OCTOBER, 1960 


Contents 


PAGE 


DISTRIBUTIONAL NOTES ON PLANTS OF THE WARM 
SPRINGS AREA, OREGON, 


Robert Ornduff and David H. French 225 
G. THomaAs Rossins (1916-1960), Rimo Bacigalupi 231 


STUDIES IN THE PERENNIAL GENTIANS: G. NEWBERRYI 
AND G. TIOGANA, Charles T. Mason 233 


CHROMOSOME COUNTS IN THE SECTION SIMIOLUS OF 
THE GENUS MIMULUS (SCROPHULARIACEAE). IV., 


Barid B, Mukherjee and Robert K. Vickery, Jr. 239 
FLOWERING RESPONSES IN PHACELIA SERICEA AND P. 
IDAHOENSIS, George W. Gillett 245 


ReEviEws: Roxana Stinchfield Ferris, [//ustrated Flora 
of the Pacific States. Washington, Oregon, and Cali- 
fornia. Volume IV — Bignoniaceae to Compositae 
(Helen K. Sharsmith); Jens Clausen and W. M. 
Hiesey, Experimental Studies on the Nature of 
Species. IV. Genetic Structure of Ecological Races 
(Henry J. Thompson). 249 


INDEX TO VOLUME XV 253 


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. COPELAND, Sacramento College, Sacramento, California. 
Joun F. Davipson, University of Nebraska, Lincoln. 

Ivan M. JounstTon, Arnold Arboretum, Jamaica Plain, Massachusetts. 
MILpreD E.. MATurAs, University of California, Los Angeles 24. 
Marion OwNBEY, State College of Washington, Pullman. 

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: Herbert G. Baker, Department of Botany, University of California, 
Berkeley, California. First Vice-president: Malcolm A. Nobs, Carnegie Institution of 
Washington, Stanford, California. Second Vice-president: Herbert F. Copeland, Sac- 
ramento City College, Sacramento, California. Recording Secretary: Mary L. Bower- 
man, Department of Botany, University of California, Berkeley, California. Corre- 
sponding Secretary: Francia Chisaki, Department of Botany, University of California, 
Berkeley, California. Treasurer: John Thomas, Dudley Herbarium, Stanford Univer- 
sity, Stanford, California. 


1960] ORNDUFF & FRENCH: WARM SPRINGS 225 


DISTRIBUTIONAL NOTES ON PLANTS OF 
THE WARM SPRINGS AREA, OREGON 


ROBERT ORNDUFF AND Davin H. FRENCH 


In an earlier paper (Ornduff and French, 1958) we reported a number 
of distributional novelties discovered during the course of identification of 
nearly 2000 vascular plant specimens from the Warm Springs Indian 
Reservation in Oregon. This collection of plants was made in connection 
with ethnobotanical studies of the Chinookan (French, 1956), Sahaptin, 
and Paiute Indians at Warm Springs. The reservation consists of approxi- 
mately 564,000 acres located in the southwest portion of Wasco County 
and the northwestern portion of Jefferson County in north-central Ore- 
gon. The summit of the Cascade range forms the western limit of the 
reservation, the Metolius River most of the southern boundary, and the 
Deschutes River in the central Oregon plateau the eastern boundary. 
The northern reservation boundary has been the subject of recent litiga- 
tion and is technically farther north than indicated on most maps and 
marked on Highway 26. Our references in this paper are to the latter 
point on the highway, which we understand is located about 6.5 miles 
south of the junction of Highway 26 with Highway 52 (in Township 6 
South, Range 10 East, Section 11, Willamette Base Line and Meridian). 

Few plants from higher altitudes are represented inasmuch as most of 
our collecting was confined to areas visited by Indians—the yellow pine 
forests, wooded valley bottoms, and the treeless sagebrush and bunch- 
grass region. About 600 species are known to occur on the reservation, of 
which at least 56 are introduced. Herein we record additional novelties 
of distribution beyond those recorded in our first paper. Previously 
accepted distributional ranges are taken from such sources as Abrams 
(1940-51), Peck (1941), and Hitchcock, Cronquist, Ownbey, and 
Thompson (1955-59). In some of these works the ranges of the species 
are often broadly phrased and risk being over-generous. We have decided 
to eliminate from our list most, but not all, of those species covered by 
broad or ambiguous statements. It should be noted that our own discus- 
sions do not necessarily cover the total ranges of the species; unless 
otherwise stated, the statement of range following each species refers 
only to its distribution within Oregon. 

We have examined specimens deposited in the herbaria at the Univer- 
sity of Oregon (ORE), Oregon State College (OSC), University of 
California, Berkeley (UC), Willamette University (WILLU), and the 
University of Washington (WTU). We extend our thanks to the staffs 
of these herbaria for their many courtesies during our visits. 

Our study of herbarium specimens confirmed our impression that the 
Warm Springs Reservation has seldom been visited by collectors. In some 
instances, however, we found in the above herbaria specimens from the 


Maprono, Vol. 15, No. 8, pp. 225-256. October 21, 1960. 


226 MADRONO [Vol. 15 


Warm Springs area which alter the ranges of the species as they are now 
known through publication. The most significant of these collections are 
noted in our list. 

Most of our specimens were identified by the first author, but the 
following specialists have also assisted: the late Carleton Ball (Salix), 
Francia Chisaki (Boraginaceae), Lincoln Constance (Umbelliferae), 
LeRoy Detling (Cardamine), Marion Ownbey (Allium), and the late 
Albert N. Stewart (Panicum). Arthur Cronquist and C. Leo Hitchcock 
have provided determinations in a large number of families. We are 
indebted to these botanists for their assistance. 

In the following list the second author’s collection numbers are used 
and, for the most part, the specimens exist as unicates in his herbarium at 
Reed College. Where noted, duplicates are at the New York Botanical 
Garden (NY) and the Herbarium of the University of California, 
Berkeley (UC). Because most of the collections were not made in dupli- 
cate and are thus not incorporated in larger herbaria, we feel justified in 
presenting these reports of new distributional data. 


NATIVE SPECIES 


PANICUM PACIFICUM Hitchc. & Chase. This grass is rarely collected 
in Oregon east of the Cascades. Our specimen, No. 908, is from beside 
the Warm Springs River near the Agency-Simnasho road, Wasco County, 
and differs little from the coastal P. occidentalis Scribn., which may not 
be distinct. Other relevant collections: Peck 15781 (WILLU) from 
Maupin, Wasco County, and Peck 17490 (WILLU) from Imnaha, Wal- 
lowa County. 

CAREX EURYCARPA Holm. Although frequently collected on the eastern 
slopes of the Cascades in Deschutes and Klamath counties, this sedge is 
rare to the northward in central Oregon. It has been collected from the 
east side of Mount Hood, Hood River County, Henderson 959 (ORE); 
from the Suttle Lake area, Jefferson County, Hitchcock & Martin 4883; 
and our specimen, No. 1494 (NY) is from the east bank of the Deschutes 
River, near the Highway 26 bridge, Jefferson County. 

ALLIUM MACRUM §S. Wats. Published records limit this onion to the 
Blue Mountain region of Oregon and Washington, but the species appears 
to be well-distributed in our area. We collected it on Miller Flat, Jeffer- 
son County, No. 597, and on Sidwalter Flat, Wasco County, No. 1763— 
both near Highway 26. Other Wasco County collections: near Kent, 
Baker 804 (OSC, WTU); near Shaniko, Cronquist 6935 (OSC, WTU), 
Gale 99 (WTU); eight miles south of Maupin, Peck 26247 (WILLU). 

ALLIUM NEviI S. Wats. According to Marion Ownbey (in litt.) this 
species is infrequently collected in Oregon. It grows north of the reserva- 
tion in Wasco County (cf. Peck 26216 (WILLU) and other collectors) ; 
we found it on Sidwalter Flat, Wasco County, No. 1811 (duplicate at 
Washington State University, Pullman). 


1960] ORNDUFF & FRENCH: WARM SPRINGS 227 


ALLIUM TOLMIEI Baker. Recorded east and southeast of the reserva- 
tion, this onion appears to be well-distributed in our region. Our collection 
is from one mile up Tenino Creek valley from the Agency, Jefferson 
County, Nos. 478A, 478B; also collected in this county by Steward 6379 
(OSC, WTU), Madras. 

SALIX DRUMMONDIANA Barratt var. SUBCOERULEA (Piper) Ball. In 
Oregon, this species is rarely collected. Peck (1941) lists it only from the 
Wallowa Mountains, but our specimens are from thickets above Shitike 
Creek, near the Agency, Jefferson County, Nos. 40, 1291A, 1291B; also 
collected in Deschutes County, WAited in 1907 (OSC) and Grant County, 
Henderson 5165 (ORE). 

ATRIPLEX CONFERTIFOLIA (Torr.) S. Wats. A member of the Great 
Basin flora which is sparingly represented on the reservation, this species 
is known from a considerable distance east of Warm Springs in Wheeler 
County. It appears to reach its northwestern limit on the reservation, 
where it grows with Sarcobatus vermiculatus (Hook.) Torr. on an alka- 
line flat in Wolfe Hollow, No. 1287. 

ACER CIRCINATUM Pursh. Though found mainly to the west of the 
Cascades, this maple is commoner on the eastern slopes than previously 
suspected. It is frequent near Beaver Creek (No. 353 came from the north 
boundary, Wasco County) and occurs along water courses traversing 
the more arid parts of the reservation. No. 205 was collected less than two 
miles west of the Agency, near Shitike Creek, Jefferson County. Other 
collections noted in herbaria are from scattered localities in Jefferson 
and Wasco counties. Warm Springs Indians say that they formerly 
obtained the tough wood locally and used it for fishing net hoops. 

ANGELICA CANBYI Coult. & Rose. This species has not been reported 
from the Warm Springs region, where we have collected it south of 
Simnasho, Nos. 717 and 1966 (UC); Indian Head Canyon, No. 1273; 
near Beaver Creek bridge on the Simnasho-Hehe Butte road, Nos. 1279, 
1423, 1917, 1954 (UC); and near Nena Creek, No. 1553—all in southern 
Wasco County. Other relevant specimens: Thompson 4951 (WTU), 
Tygh Hill, Wasco County; and Peck 18646, near summit, Ochoco Forest, 
Crook County. 

LomaTIuM cous (S. Wats.) Coult. & Rose. Apparently not previously 
collected as far west as Warm Springs, where it is locally abundant. 
Peck 26166 (WILLU), from eight miles south of Maupin, Wasco County, 
is near the reservation. Our Wasco County specimens: Nos. 473, 528, 
south side of Warm Springs River valley, near Agency-Simnasho road; 
Nos. 1219, 1869 (UC), Sidwalter Flat, near Highway 26. Jefferson 
County: No. 649, Miller Flat, north of Highway 26. Like other Indians 
of the Columbia Plateau, the Warm Springs Indians dig the starchy tubers 
for food. 

LOMATIUM LEPTOCARPUM (Torr. & Gray) Coult. & Rose. Published 
records locate the western limit of this umbellifer in the Blue Mountains. 
However, Cronquist 7425 (UC) came from Big Summit Prairie, Wheeler 


228 MADRONO [Vol. 15 


County, and Peck has collected the species in various parts of Wasco 
County. Our Nos. 526 and 1896 (UC) came from west of Simnasho, and 
Nos. 584A, 584B, 584C, and 1755 were collected about a mile northwest 
of Hehe Butte, all in Wasco County. 

LOMATIUM NEVADENSE (S. Wats.) Coult. & Rose var. NEVADENSE. 
Warm Springs represents the northwestern limit for this Great Basin 
species, which was collected southeast of the Agency area longhouse, 
Jefferson County, Nos. 516A, 516B, 516C, and 1900 (UC). 

PTERYXIA TEREBINTHINA (Hook.) Coult. & Rose var. TEREBINTHINA. 
Widely distributed in eastern Oregon, but not previously collected within 
many miles of Warm Springs. Both Nos. 1274 and 1897 (UC) came from 
Indian Head Canyon, near the Agency-Simnasho road, Wasco County; 
No. 1582 was collected from a rocky point north of the junction of the 
Whitewater and Metolius rivers, Jefferson County. 


CAMPANULA SCOULERI Hook. Although frequent in the woods of west- 
ern Oregon, this species is rarely reported east of the Cascades. There are 
a few northern Oregon collections from the region east of Santiam Pass 
and from the eastern slopes of Mount Hood. Our specimens were found 
near Highway 26 at the northern reservation boundary, Wasco County, 
No. 1342. Thus this species might be considered a regular inhabitant of 
the eastern slopes of the Cascades, at least in the northern portion of the 
State. 


SENECIO MACOUNII Greene (=S. fastigiatus Nutt.) While this typi- 
cally western Oregon species is very closely related to the eastern Oregon 
Senecio canus Hook., most specimens are clearly referable to one or the 
other species, and intermediates are lacking. A number of specimens from 
eastern Oregon have been referred by various collectors to S. macounit, 
although they more properly belong in S. canus. We have, however, found 
“good” S. macouni in the open pine woods at the Hehe celebration 
ground, Wasco County, Nos. 1463 and 1464 (both NY). 

STYLOCLINE FILAGINEA Gray. Generally attributed to extreme eastern 
and southeastern Oregon, this species has recently been collected in 
Jefferson County 20 miles northeast of Madras, Peck 26156 (WILLU) 
and from a single colony in the Agency area at Warm Springs, our No. 
1690 (NY). 

TETRADYMIA GLABRATA Gray. Another species reaching its northwest- 
ern range limit on the reservation. In Oregon, it is most frequently col- 
lected in the southeastern counties, but recently it has been found near 
Mitchell, Wheeler County, Cronquist 7259 (OSC). Our plant, No. 1250 
(NY), was growing southeast of Hehe Butte, Wasco County, about 75 
miles northwest of Mitchell. 


INTRODUCED SPECIES 


RUBUS LACINIATUS Willd. Sparingly established in Shitike Creek valley 
west of the Agency, Jefferson County (No. 980). Only one other Oregon 


1960] ORNDUFF & FRENCH: WARM SPRINGS 229 


specimen seen from east of the Cascade Mountains, Small 30 (ORE), 
from Link River, Klamath County. 

ANTHRISCUS SCANDICINA (Weber) Mansfield. This European species 
is well established in western Oregon, but only a single collection from 
east of the mountains was located in the herbaria: Hitchcock 20441 
(WTU), near the mouth of the John Day River, Sherman County. Our 
Nos. 827 and 942 came from a settled area west of the Agency. 

MYoSOTIS MICRANTHA Pall. ex Lehm. In our earlier paper (Ornduff 
and French, 1958, p. 220) we mistakenly referred our No. 898 to M. 
discolor Pers. This has proved to be another Old World species, M. 
micrantha, as has our (previously uncited) No. 1808 from Sidwalter 
Flat, Wasco County. 


DISCUSSION 


In general, the new stations reported for the various species in the 
above list and in our previous paper are rather well-distributed over the 
reservation. There are few areas in which the “‘extra-limital” species are 
aggregated, as might be expected in view of both the geological and 
vegetational continuity of the reservation with much of the rest of central 
Oregon. An exception to this rule, however, is a large aggregation on or 
near the reservation of species characteristic of the more mesic western 
portions of the state, which are seldom collected east of the Cascade range. 
As might be expected, these species are typically inhabitants of moist 
ground along streams or rivers, or less frequently they are woodland 
plants. These species are: Holcus lanatus L., an introduction found in 
Wasco County by us and in Deschutes County by others; Carex aperta 
Boott., found by us in Wasco County, and known also from Union 
County; Eleocharis obtusa (Willd.) Schultes, known from Hood River, 
Umatilla, and Union counties, and here reported from Jefferson County; 
Cardamine oligosperma Nutt., found outside the reservation in Wasco 
and Sherman County, as well as by us inside the reservation in Wasco 
County; Perideridia oregana (S. Wats.) Mathias, found by us in two 
localities on the reservation in Wasco County, and known also from 
various Klamath County collections; Trichostema oblongum Benth., 
infrequent in the counties south of the reservation and collected by us 
on the reservation in Wasco County; Artemisia douglasiana Bess., grow- 
ing along the Deschutes River in Jefferson County; and Guaphalium 
chilense Spreng., sporadic in Deschutes and Umatilla counties and found 
by us in Jefferson County as well. 

In addition to these characteristically western species which are seldom 
collected east of the Cascades, we have noted additional species which 
have apparently never been reported from east of these mountains. These 
are: Juncus effusus L. var. pacificus Fern. & Weig., locally abundant 
along Shitike Creek, west of the Agency, Jefferson County, Nos. 215, 
940, and 1509 (NY) as well as along Beaver Creek near the north reser- 
vation boundary, Wasco County, No. 1644; Achlys triphylla (Smith) 


230 MADRONO [Vol. 15 


DC., found well down on the eastern slopes of the Cascades at the north 
reservation boundary near Highway 26, No. 363, and near the upper 
Warm Springs River, No. 866, both Wasco County; Ribes sanguineum 
Pursh, various collections noted from Jefferson County sites, and also 
found in a number of localities along upper Beaver Creek where it 
parallels Highway 26, e.g., our Nos. 355, 627A, 627B; Angelica genu- 
flexa Nutt., found in scattered moist areas southeast of Mount Hood in 
Wasco County, as near Highway 26 one mile north of the north reserva- 
tion boundary, No. 1846 (UC), and also collected by other workers east 
of the Cascade summit in Klamath County; Gentiana sceptrum Griseb., 
collected in a damp meadow near Highway 26 several miles southeast of 
its junction with Highway 52, Wasco County, No. 1389; and Veronica 
serpyllifolia L. var. serpyllifolia from along Beaver Creek about 0.5 miles 
northwest of the Highway 26 intersection with the north reservation 
boundary, Wasco County, No. 1803. 

Most of the species discussed in the present and previous papers fall 
into two phytogeographical groups: (1) those which occur at Warm 
Springs in populations disjunct from the reported range of the species, 
and (2) those which appear to be beyond the margins of their previously 
reported ranges. In the first group are species such as Convolvulus poly- 
morphus, Phacelia thermalis, and Stylocline filaginea. Intensive collecting 
in the areas adjacent to the reservation will likely show that many of 
these apparently disjunct stations are in fact connected with the main 
range of the species by geographically intermediate populations. In the 
second group are species such as Pinus lambertiana, Allium macrum, 
Achlys triphylia, Angelica genuflexa, and Tetradymia glabrata, many of 
which appear to be at their northern, southern, eastern, or western limits. 
The majority of the species in this latter group are those typical of the 
more mesic western portions of the state which have evidently migrated 
into suitable sites in our region either through the Columbia gorge and 
thence southward along the eastern slopes of the Cascades, or via the 
low mountain passes in the Cascades. Undoubtedly most of the weedy 
species reported are relatively recent introductions which may or may not 
become permanent members of the naturalized flora of the reservation. 
It is likely that a few native species, such as Phacelia thermalis, have 
been recently introduced and will not become established. However, the 
majority of the species we have discussed are well-established and occur 
in large colonies and/or in a number of widely separated vigorous 
populations. A notable exception is Pinus lambertiana, represented on 
the reservation by a few isolated senescent individuals which are not 
reproducing. 

Department of Botany, 
University of California, Berkeley. 


Reed College, 
Portland, Oregon. 


1960] BACIGALUPI: ROBBINS 238 


LITERATURE CITED 


ABRAMS, LEROY. 1940-1951. Illustrated Flora of the Pacific States, vols. I-III. Stan- 
ford, California. Stanford University Press. 

FRENCH, Davin. 1956. An Exploration of Wasco Ethnoscience. Year Book of the 
American Philosophical Society, pp. 224-226. 

Hircucock, C. LEo, ARTHUR CRONQUIST, MARION OwnBeEy, and J. W. THOMPSON. 
1955-1959. Vascular Plants of the Pacific Northwest, vols. IV & V. Seattle, Uni- 
versity of Washington Press. 

ORNDUFF, RosBerT, and Davip FRENCH. 1958. Notable Plants of the Warm Springs 
Indian Reservation, Oregon. Leafl. West. Bot. 8(9) :217-220. 

Peck, Morton E. 1941. A Manual of the Higher Plants of Oregon. Portland, Binfords 
and Mort. 


G. THOMAS ROBBINS (1916-1960) 


Early on the morning of February 11th, 1960, a few days after his 
forty-fourth birthday, G. Thomas Robbins, Herbarium Botanist in the 
Jepson Herbarium of the University of California Botany Department, 
died quietly in his sleep. The resultant sense of shock and loss among his 
botanical colleagues, especially those in Berkeley, was both deep and 
lasting. 

Tom Robbins was born in San Francisco on February 6, 1916. While 


232 MADRONO [Vol. 15 


he was still a boy, his family moved to Windsor in Sonoma County, Cali- 
fornia, and in the local schools and those in Santa Rosa, Tom received 
his elementary and high school education. His early interest in matters 
botanical was stimulated by the guiding hand of Mr. Milo S. Baker, who 
was his Professor of Botany during the two years of Tom’s attendance at 
Santa Rosa Junior College. Later he attended the University of California 
at Berkeley, working meanwhile as a valuable student assistant in the 
University Herbarium, and graduating there in Botany in 1938 with a 
Bachelor of Arts degree. Later, he spent a profitable year at the Univer- 
sity of Colorado during which he wrote an excellent monograph of the 
North American species of Androsace, which was later published in the 
American Midland Naturalist (Vol. 32, pp. 137-163, 1944). This work 
earned him the Master of Arts degree in 1941. 

In 1946, Tom was appointed Associate Professor of Biology at the East 
Central State College at Ada, Oklahoma. Tom enjoyed his teaching activi- 
ties greatly, but increasing deafness, which had been plaguing him for 
several years, became so acute as to force his retirement from a teaching 
career at the end of the school year in 1949. 

Subsequently, for nearly three years, Tom was engaged as one of 
several botanical assistants aiding Dr. Herbert Mason, Director of the 
Herbarium of the University of California, in the preparation of ‘‘A Flora 
of the Marshes of California.”” Tom carried on some of the field work in 
connection with this project, contributed the final draft of the manu- 
scripts of the Gramineae and Cyperaceae as well as of some genera in 
which he had a special interest, and aided in the solution of many taxo- 
nomic, bibliographic, and nomenclatural problems. Early in 1952, Tom 
joined the staff of the Jepson Herbarium as Herbarium Botanist, a posi- 
tion which he filled most capably, contributing manuscript on selected 
genera treated in the still unpublished portions of Jepson’s ‘‘A Flora of 
California,” which position he held at the time of his lamentably untimely 
death. 

For some years, Tom gave unstintingly of his time and effort to fulfill 
the often thankless chores associated with his activities as Corresponding 
Secretary of the California Botanical Society. 

As in the case of so many of his other pursuits, Tom’s passion for 
excellence eventually resulted in his becoming a very fine photographer of 
close-up studies of native flowers, as the Kodachrome collection which he 
built up for the Jepson Herbarium well attests. Thus, as in his botanical 
endeavors, his life was enriched by a happy merging of vocational and 
avocational pursuits. Purely on the avocational level, Tom’s interest in 
and understanding of music brought meaningful enjoyment, often shared 
with his closest friends, to his leisure hours. 

Tom Robbins’ outstanding characteristics were his gentleness, charity 
in judgment, an unusual ability to keep his own counsel, a passion for 
accuracy, and a tendency to reserve his judgment, either botanical or 
otherwise, until all pertinent data had been fully assessed and “digested.” 


1960] MASON: GENTIANS 233 


To quote from a letter from Joseph Ewan, who was his professor while he 
attended the University of Colorado, ‘“‘his meticulous care, almost fanati- 
cal, in the handling of records, and extreme interest in assembling all the 
pertinent literature on a topic before committing himself by way of a 
botanical judgment” were among his most valuable assets. These charac- 
teristics are amply exemplified in his last published work, ‘Notes on the 
Genus Nemacladus” (Aliso, Vol. 4, pp. 139-147, 1958), in which two 
new species and new interpretations of already published taxa were 
published. 

Tom’s name is commemorated in Phacelia strictiflora Gray var. Rob- 
binsti Constance (Contr. Gray Herb. 168:20, 1949), which was based on 
one of Tom’s collections in Oklahoma. 

Besides his membership in the California Botanical Society, he was a 
member of Sigma Xi, the Society for the Study of Evolution, and of 
the International Association for Plant Taxonomy. 

Tom Robbins will long be missed by those whose good fortune it was 
to know him at all well—R1mo BacicALuPI, Jepson Herbarium, Depart- 
ment of Botany, University of California, Berkeley. 


STUDIES IN THE PERENNIAL GENTIANS: 
G. NEWBERRYI AND G. TIOGANA!? 


CHARLES T. MASON, Jr. 


Gentiana newberryi Gray is the name applied to a group of dwarf 
perennial gentians of Section Pneumonanthe, which occurs in the Sierra 
Nevada and Cascade Mountains of California and Oregon. The name is 
used in many manuals (Abrams, 1951; Jepson, 1925; Munz, 1959; Peck, 
1941) to embrace not a single species but two species and a number of 
intermediate forms. While the author was studying the western perennial 
gentians under a National Science Foundation grant, the problem came 
to light, and an attempt is here made to resolve the difficulty. 

Asa Gray described Gentiana newberryi from material collected in 
Oregon by Newberry, a member of the Williamson Pacific Railroad 
expedition of 1855. He first presented the new name in a hand-written 
description on the type sheet in the Gray Herbarium. Only the Newberry 
specimen was cited with this description. However, by the time the name 
was published (Gray, 1876), the circumscription was modified to include 
California material, and a Bolander collection from Mariposa County 
was added as a syntype. 

Three other names have been applied to this complex. In 1931 East- 
wood described G. copelandu, which she separated from G. newberryi on 
the “much broader leaves and dark purple flowers.” The name G. cope- 
landi, having been used previously by Greene (1904) and by Elmer 


1 Arizona Agricultural Experiment Station Technical Paper No. 568. 


234 MADRONO [Vol. 15 


(1915), was invalid, causing Eastwood to correct the name of this species 
to G. eximia (Eastwood, 1934). A third species, G. tzogana, was recog- 
nized in 1940 by Heller, who separated it from G. newberryi by its 
smaller size. 

Of the three validly published names, G. newberryi, G. eximia, and 
G. tiogana, only the first has been accepted by students of the flora of 
California and Oregon. The present author, after studying many field 
and herbarium specimens, is convinced that two species should be recog- 
nized. They are both perennials with a rosette of leaves and with flowering 
stalks arising from the axils of last year’s leaves. Each flowering stalk 
usually has a single flower, and the patterning on the flowers is the same; 
however, the two species can be separated by the following characters: 


Flowers 4.0-5.5 cm. long; corolla bright blue with 5 dark greenish stripes, the 
lobes 1.0-1.5 cm. long; plicae many-lobed with two long thin central lobes 
0.7-0.9 cm. long; leaves broadly spatulate to suborbicular . G.newberryi 

Flowers 2.5—3.5 cm. long; corolla white or very pale blue with 5 dark greenish- 
brown stripes, the lobes 0.7-0.8 cm. long; plicae 0.4 cm., two-lobed 
with an occasional third or fourth lobe; leaves lanceolate to narrowly 
ObOVAtE- ges os 2. al A bees! Gee a. Boe eG a 0cd7d 


GENTIANA NEWBERRYI Gray, Proc. Am. Acad. 11:84. 1876. Pneumo- 
nanthe newberryi (Gray) Greene, Leafl. Bot. Obs. 1:71. 1904. Dasyste- 
phana newberryi (Gray) Arth. Torreya 22:30. 1922. Gentiana copelandu 
Eastw. Proc. Calif. Acad. Sci. Series 4, 20:150. 1931; non Gentiana cope- 
landii Greene, 1904, or Elmer, 1915. Gentiana eximia Eastw. Leafl. West. 
Bot. 1:96. 1934. 

Low rhizomatous perennial with a rosette of broadly oblong-spatulate 
to suborbicular leaves up to 6 cm. long and 2 cm. wide; flowering stems 
1-5, decumbent to erect, 15 cm. long, from axils of last year’s leaves; 
flowers 4.0—5.5 cm. long and usually 1 per stem; calyx tube 0.8—1.5 cm. 
long, the lobes 0.7—1.1 cm. long, lanceolate to elliptical; corolla convolute 
in the bud, funnelform after anthesis, 4.0—5.5 cm. long, bright blue with 
5 dark purple stripes extending from the tips of the lobes to the base of 
the corolla, the lobes 1—1.5 cm. long, entire or erose, broadly rounded 
with yellow-green dots on the inner surface and extending down into 
corolla tube, the apices apiculate; plicae 0.7—0.9 cm. long, bifid with 
two long attenuate lobes and several secondary lateral projections; 
stamens maturing before the pistil, the anthers extrorse; style none; 
capsule ellipsoidal, 1.5 cm. long, the stipe 1.5 cm. long; seeds broadly 
winged all around. 

Type. Crater Pass, west side Cascade Mountains, lat. 44°, Oregon, 
Newberry sn. (GH). Type was seen. 

Gentiana newberryi is known only from the Three Sisters in the 
Cascade mountain area of central Oregon, and from the Mount Eddy 
region in Siskiyou and Trinity counties, California. An unexplained dis- 
tributional gap exists from northern California to central Oregon. A 
similar disjunct distribution is exhibited by Limnanthes douglasi R. Br. 


1960] MASON: GENTIANS 239 


X Gentiana newberryi 
@ G.newberryi x G.tiogana 
O Gentiana tiogana 


Fic. 1. Distribution of Gentiana newberryi, G. tiogana, and their hybrid in 
Oregon, California, and Nevada. 


var. douglasii, which has the northern limit of its California distribution 
in Humboldt County and is again found in the Umpqua Valley, Douglas 
County, Oregon (Mason, 1952). 


Representative specimens. OREGON. Deschutes County: Three Creeks Meadow, 
Brandt & Steward 6985 (ID, UTC), Ellis & C. Mason 1712; meadow near Three 
Creeks Lake, Whited 478 (WS), Ellis & C. Mason 1711; Fremont’s Crossing of 
Tumalo Creek, Whited 479a (WS). Lane County: 5 miles west of McKenzie Pass, 
Campbell 17497 (CAS); Hand Lake, 4 miles west of Lane-Deschutes county line, 
T.& C. Mason 1791. 

CALIFORNIA. Siskiyou County: Mount Eddy, Copeland 3878 (CAS), Eastwood 
2037 (type of G. copelandii Eastw., CAS.). Trinity County: Morris Meadow, Stuart 
Fork, Alexander & Kellogg 5525 (UC); edge of Bull Lake, Parker in 1947 (DS). 


GENTIANA TIOGANA Heller emend. C. T. Mason. G. tiogana Heller, 
Leafl. West. Bot. 2:221-—222. 1940. 
Low rhizomatous perennial with a rosette of obovate to spatulate leaves 
ca. 4 cm. long and 1 cm. wide; flowering stems 1 or 2, decumbent to 
erect, 5—7 cm. long, from the axils of last year’s leaves; flowers 2.5-3.5 


236 MADRONO [Vol. 15 


cm. long, usually 1 per stem; calyx tube 0.6—1.0 cm. long, the lobes 0.6— 
1.0 cm. long, lanceolate to elliptical; corolla convolute in the bud, funnel- 
form after anthesis, 2.5-3.5 cm. long, white or very light blue with 5 
greenish-brown stripes extending from the tips of the lobes to the base 
of the corolla, the lobes 0.7—0.8 cm. long, entire or erose, narrow with 
yellow-green dots on the inner surface and extending down into corolla 
tube, the apices apiculate; plicae 0.4 cm., with 2 large lobes and an occa- 
sional third or fourth smaller lateral lobe; stamens maturing before the 
pistil, the anthers extrorse; capsule ellipsoidal, 1.0—1.2 cm. long, the stipe 
1.0 cm. long; seeds broadly winged all around. 

Type. Shore of a lakelet outside Yosemite National Park, Tioga Pass, 
Mono County, California, Heller 15453 (WTU #79272. Isotypes, CAS, 
DS, NY, UC, WTU). The type specimen was not seen, but the infor- 
mation was transmitted by letter from C. L. Hitchcock. Isotypes from 
California Academy of Sciences, Dudley Herbarium, and University of 
California were seen. 

Heller’s labels agree on the collection number, the date, and that the 
collection was made outside Yosemite Park boundary. They vary to some 
degree on the other data presented. The county is listed as either Mari- 
posa or Mono. As the park boundary is also the county line, ‘‘outside the 
park” would be Mono County. The distance outside the park varies from 
a few yards to one-fourth mile, and the elevation ranges from 9900 to 
9940 feet. The labels also state, “‘south of Tioga Pass entrance”’; the park 
boundary at that particular point extends east and west so that south of 
the entrance would be inside, not outside, the park. Heller probably con- 
sidered the general direction of travel from Mono Lake to the Tioga Pass 
entrance through Leevining Canyon as west; therefore, to the left would 
be south. In the area to the east of the road at the Tioga Pass entrance is 
a meadow with a number of lakelets, and it undoubtedly is in this area 
that Heller made his collection. 

The specific epithet tzogana was applied by Heller to specimens which, 
because of their smaller size, he considered distinct from the Sierra 
Nevada material recognized as Gentiana newberryt. This small form has 
been collected several times from the higher elevations and may warrant 
varietal recognition, but this author does not consider it distinct enough 
to be separated as a species; consequently Heller’s epithet becomes the 
first applied to this group and must be used, and his description has been 
here emended to include the larger forms. 

Representative specimens. CALIrornIA. Inyo County: Big Pines Lake, Howell 
24123 (CAS); Mosquito Flat, Rock Creek, Halperin 605 (CAS), Ferris & Lorraine 
11086 (DS); Heart Lake, Rock Creek Basin, Peirson in 1933 (DS, UC), Peirson 
9483 (COLO); Kearsarge Pass Trail west of Independence, Alexander & Kellogg 
3291 (DS, UC); Brown Lake, Raven & Stebbins 254 (CAS, UC); Cottonwood 
Lakes, Alexander & Kellogg 3316 (DS, UC, UTC). Tulare County: Crabtree 
Meadows, H. M. & G. Hall 8442 (UC); Lost Canyon, Howell 17787 (CAS) ; 
Rock Creek, Howell 25515 (CAS, UC, UTC, WS) ; Army Pass, Howell 26045 (CAS) ; 
Sequoia-Mount Whitney trail, Sisson & Kobs in 1928 (COLO); Humphrey Basin, 
Moran 490 (DS). Fresno County: Hilgard Branch, Bear Creek, Raven 7872 (CAS) ; 


1960] MASON: GENTIANS 237 


Humphrey Basin west of Mount Humphreys, C. Sharsmith 3149 (UC). Madera 
County: Iron Creek, Raven 3809 (CAS). Mariposa County: Mount Hoffman, Rodin 
885 (UC). Mono County: Slate Creek, Hall Natural Area, Clausen 920 (DS, UC), 
C. Mason 1514; Dana Meadow, Tioga Pass, Rowntree in 1931 (CAS); ™% mile 
upstream from Camp Tioga along Slate Creek, C. Mason 1512; southwest end Little 
Virginia Lake, Hendrix 604 (UC) ; Soda Springs, Tuolumne Meadows, Eastwood 625 
(CAS) ; Bourland Meadows, Belshaw 81 (UC); Dana Plateau, northwest of Mount 
Dana, C. Sharsmith 2331 (UC); White Mountain, Mount Conness Range, H. Mason 
11339 (UC); Ten Lakes Basin, H. Sharsmith 1329 (UC); near Dog Lake, Howell 
20434 (CAS). Alpine County: meadow 3 miles west of Lake Alpine, C. Mason 1610. 
Eldorado County: “Benwood Meadows, Camp Echo, Heller 12264 (CAS, COLO, DS, 
UC); Dicks Lake, Lake Tahoe, Alexander & Kellogg 3508 (UC); Meyers Station, 
Clemens in 1920 (CAS). Butte County: Jonesville, Spring Creek, Copeland in 1931 
(UC). NEvaDA. Washoe County: 3 miles south Mount Rose, Hitchcock & Martin 
5542 (DS, UC, UTC, WS); Galena Creek, south base Mount Rose, Hitchcock & 
Martin 5522 (DS, UC, UTC) ; Mount Rose, Heller 9970 (CAS, DS, MONTU). 


Naturally and artificially produced hybrids among the gentians are 
well-known (Mason, 1959), and a number of herbarium specimens from 
northern California show evidence of hybridization and introgression 
between G. tiogana and G. newberryi. Three collections from Eldorado 
County (Wrights Lake, Johannsen 452, UC; % mile north of Wrights 
Lake, Robbins 1355, CAS, UC; and Echo Summit, Howell 22974, CAS) 
have the characteristics of G. tiogana except that the plicae tend to be 
long and narrow. 

Several collections from Nevada, Sierra, Plumas, Lassen, and Shasta 
counties in California have characteristics of both species. Specimens from 
Sage Hen Creek (H. Mason 14472, UC), and Independence Lake (Floyd 
in 1925, CAS, Alexander & Kellogg 5160, UC), of Nevada County, and 
Webber Lake, Sierra County (Dudley in 1894, DS) have stem length, 
leaf size but not shape, and plicae lobing which approach those of Gen- 
tiana newberryi. The flower size is intermediate, ranging from 3.5—4.3 cm. 
on the collections from Independence Lake, and 3.8—4 cm. on the other 
two collections. The plicae are of the larger type found in G. tiogana, and 
the green and white flower color of that species is specified on Mason’s 
collection. Possibly the larger size of the leaves and stems is the result of 
ecological rather than genetic factors. 

A sheet of specimens from Mount Elwell, Plumas County (Wicks 2889, 
UC), has flowers and leaves approaching those of G. newberryi. The 
flowers are 4—4.5 cm. long and the calyx lobes are large and elliptic. The 
plicae, although multilobed, are the large type of G. tiogana. The flower 
color is not specified, but the specimens appear dark as though there may 
be some blue factors present. One plant on the sheet is quite different from 
the others; it has the smaller flowers, smaller leaves, white color, and 
heavy plicae of G. tiogana. The presence of the two types in the same 
collection gives good evidence that both are present in the same area so 
that hybridization might occur between the two species. 

Several collections from northern Plumas County (Big Meadows, 
Coombs in 1912, CAS, UC, Austin 399, UC; Prattville, Coombs in 1906, 
CAS), southeastern Shasta County (Lassen’s Peak, Brewer 2175, UC; 


238 MADRONO [Vol. 15 


upper King’s Creek Meadow, Hoover 4612, UC), and southwestern 
Lassen County (Susanville, Safford s.n., UC; Mountain Meadows, Austin 
in 1879, UC; Hog Flat, Stebbins et al 3998, UC; Harvey Valley Spring, 
Whitney 1505, UC; a mile east of Westwood, Heller 15341, DS, UC) 
show evidence of introgression. Of particular interest are the two collec- 
tions from Big Meadow which have several flowers per stem. This mul- 
tiple flower condition is uncommon in either of these parent species, but 
it is seen on some collections by Lemmon which are affixed to the type 
sheet of G. newberryi. Undoubtedly these Lemmon collections were made 
from this northern California area, although the data on the label are 
vague and incomplete. The Brewer collection from Mount Lassen and 
Safford’s collection from Susanville have very small plants and flowers 
and show more of the characters of G. tiogana than G. newberryi. 

The two species under consideration are usually separated by alti- 
tudinal differences. Most collections of G. newberryi have come from 
areas between 4000 and 6500 feet in elevation, but one specimen from 
Bull Lake, Trinity County, California (Parker in 1947, DS) is listed as 
7380 feet. Gentiana tiogana frequently is found above timberline, and 
specimens from 12,000—13,000 feet are not uncommon. The lowest eleva- 
tion noted for a collection of G. tiogana is 7160 feet at Jackass Meadow, 
Fresno County (Peirson 12880, CAS). 

The specimens cited as having characteristics of both species have been 
collected from areas which, for the most part, are intermediate in eleva- 
tion between the usual requirements of the two parents. The collection 
of specimens from “a mile east of Westwood,” Heller 15341, is cited as 
elevation 5000 feet, and these plants are predominantly of the G. new- 
berryi type. On the other hand the collection from Lassen’s Peak, Brewer 
2175, was made at 8000 feet, and these specimens have the majority of 
their characters similar to G. tiogana. This overlap in the altitudinal dis- 
tribution of intermediate forms might also be regarded as evidence that 
G. newberryi and G. tiogana hybridize in northern California. 


Department of Botany 
University of Arizona, Tucson 


LITERATURE CITED 


ABRAMS, L. 1951. Illustrated flora of the Pacific States Vol. 3. Stanford Univ. Press. 
ARTHUR, J. C. 1922. Changes in phanerogamic names. Torreya 22:30. 
Eastwoop, A. 1931. New species of plants from western North America. Proc. Calif. 
Acad. Sci. Ser. 4, 20:150. 
. 1934. Gentiana copelandii Eastwood. Leafl. West. Bot. 1:96. 
Ermer, A. D. E. 1915. Two hundred twenty-six new species. Leafl. Philip. Bot. 
7:2671 (not seen). 
Gray, A. 1876. Miscellaneous botanical contributions. Proc. Am. Acad. Sci. 11:84. 
GREENE, E. L. 1904. North American species of Amarella. Leafl. Bot. Obs. 1:53. 
. 1904. The genus Pneumonanthe. Leafl. Bot. Obs. 1:71. 
He ter, A. A. 1940. California plants, mostly new. Leafl. West. Bot. 2:221. 
Jepson, W. L. 1925. Manual of the flowering plants of California. Assoc. Students, 
Berkeley. 


1960] MUKHERJEE & VICKERY: MIMULUS 239 


Mason, C. T. 1952. A systematic study of the genus Limnanthes. Univ. Calif. Publ. 
Bot. 25:455-512. 
. 1959. A hybrid among the perennial gentians. Brittonia 10:40—43. 
Mowz, P. A. 1959. A California flora. Univ. Calif. Press. Berkeley. 
Peck, M. E. 1941. A manual of the higher plants of Oregon. Binford and Mort, 
Portland. 


CHROMOSOME COUNTS IN THE SECTION 
SIMIOLUS OF THE GENUS MIMULUS 
(SCROPHULARIACEAE). IV. 


Barip B. MUKHERJEE AND ROBERT K. VICKERY, JR. 


This report! on the determination of chromosome numbers in the sec- 
tion Simiolus of the genus Mimulus is part of a long range investigation 
into the evolution of species in Mimulus (Vickery, 1951). The chromo- 
some numbers and configurations presented in this article indicate a lack 
of cytological differentiation between several of the currently accepted 
species (Pennell, 1951) of the section Simiolus. Also they reveal the pres- 
ence of aneuploidy in different populations of two other species, and, 
lastly, they fill an important gap in the previously indicated (Mukherjee 
and Vickery, 1959) polyploid series that extends from North to South 
America. 

Essentially the same method of bud fixation was employed as in the 
previous investigation (Mukherjee and Vickery, 1959), 1.e., fixation in 
two parts absolute ethanol to one part glacial acetic acid saturated with 
ferric acetate, followed by staining of the anthers in iron-aceto-carmine. 
Work now in progress indicates that there may be possible improvements 
in this schedule. Each chromosome number determination is based on 
counts from an average of approximately eight pollen mother cells. Cam- 
era lucida drawings were made for three or four figures for each count 
and, in addition, photomicrographs were taken of many of the configura- 
tions. Herbarium specimens of each culture have been or will be deposited 
in the Garrett Herbarium of the University of Utah (UT). 


1 This work was supported by the National Science Foundation. Part of it was 
included in the dissertation of the senior author submitted to the Faculty of the 
University of Utah in partial fulfillment of the requirements for the Ph.D. degree. 

The authors wish to thank the following people who made this study possible by 
so kindly gathering seeds or living plants for them: Lawrence Beane, Martin Brown, 
Annetta Carter, Norman V. Chamberlain, Jens Clausen, John L. Creech of the 
Plant Introduction and Exploration Division of the United States Department 
of Agriculture, William M. Hiesey, Richard W. Holm, Carl L. and Laura C. Hubbs, 
David D. Keck, Oliver Norwell, George T. Oberlander, the late Francis W. Pennell, 
Carl W. Sharsmith, G. Ledyard Stebbins, Jr., H. J. Venema, Delbert Wiens, and 
Ira L. Wiggins. 


240 MADRONO 
of, 
: o On b ey 
mis ‘s pe ott 
. ; ? t ~*% 
5014 5091 5382 
_ 
oe 
Dee ¢ 0 ? ie iy +e 
SPA LE ate 
5864 502 5690 
2- 
os ay 4 wd 
sr 09s oFoaws a w%e 
*%e! o a 8 
5339 5658 5044 
§ N ‘ 8 
4ot ( bY WA ¢ oe, 84, 
ee% § & ) 3°6ye 
5327 5752 6062 
‘~ 
Pear NY p 8 
02 Wed Sec, b p> 4 ‘ 
2° ar sou’ é ’ g a 
aces a ore 
Se, ee X¢ ¢ 6 
5373 5081 5042 


@uced, = < 840). 


[Vol. 15 


s 
5018 


0 1020 micra 


Fic. 1. Meiotic chromosomes of Mimulus: M. guttaus, 5014, 5091, 5382, 5836, 
5864; M. tilingz var. tilingii, 5012, 5690, 6117; M. laciniatus, 5339; M. laxus, 5658; 
M. nasutus, 5044, 5018, 5327; M. platycalyx, 5752; M. glabratus var. utahensis, 
6062; M. glabratus var. fremontii, 5063, 5373; M. tigrinus, 5081; M. luteus, 5042, 
5043. All cells are in or near second metaphase except 5382, 5836, 5012, 5690, 5658, 
5018, and 5752, which are in first metaphase. (Camera lucida drawings as repro- 


1960] MUKHERJEE & VICKERY: MIMULUS 241 


A total of thirty-three cultures was studied during the present investi- 
gation (see table 1). They include representatives of ten species and vari- 
eties of the section Simiolus: Mimulus guttatus DC., M. tilingu Regel 
var. tilingii, M. laciniatus Gray, M. nasutus Greene, M. laxus Pennell, 
M. platvcalyx Pennell, M. glabratus var. utahensis Pennell, M. glabratus 
var. fremonti (Benth.) Grant, M. tigrinus hort., and M. luteus L. 

All fifteen cultures of M. guttatus were found to have n=14 chromo- 
somes. The configurations were regular and similar to those previously 
observed for other cultures of M. guttatus (Vickery, 1955; Muhkerjee, 
Wiens, and Vickery, 1957; Mukherjee and Vickery, 1959) and, therefore, 
only a few of the camera lucida drawings of M. guttatus chromosomes 
were included in figure 1. The fifteen cultures examined represent much 
of the geographical range of M. guttatus (see table 1) and much of its 
morphological and physiological diversity as well. Morphologically the 
cultures differ from each other in the average height of the plants, shape 
of the leaves, amount and distribution of anthocyanin pigmentation, and 
in the size of the flowers. Physiologically they differ in growth rates, time 
and speed of flowering, and time and speed of maturing seeds. Cytologi- 
cally the only detectable difference observed among the fifteen cultures 
was the presence of marked chromosome stickiness under the present fixa- 
tion schedule in two of the annual races. Despite the wide range of mor- 
phological and physiological differences between the various cultures, 
they all exhibit apparently similar karyotypes. 

The chromosome numbers of six cultures of M. tiling var. tilingu 
from the Sierra Nevada (see table 1) were found to be n=14 as in M. 
guttatus. Mimulus tilingit var. tiling is related to M. guttatus on the 
basis of morphology (Hitchcock, Cronquist, Ownbey and Thompson, 
1959), but is separated from it by strong crossing barriers (Vickery, 
1956). The chromosome configurations of the six cultures were regular 
and similar to those of M. guttatus and to our first M. tilingit var. tilingit 
count (Vickery, 1955), but differed in number from our more recent re- 
port of n=15 for a Utah population of M. tilingi var. tilingi (Mukherjee 
and Vickery, 1959). The populations studied represent much of the mor- 
phological diversity present in M. tilingw var. tilingi in the Sierra Nevada 
(table 1). 

Mimulus laciniatus and M. laxus, species that are genetically closely 
related to M. guttatus (Vickery, 1956, and unpublished), were found to 
have n—14 chromosomes as does M. guttatus. The karyotypes of the 
three species are apparently indistinguishable. Mimulus laciniatus is mor- 
phologically strikingly different from M. guttatus, whereas M. laxus is 
closely similar. Probably M. laciniatus should be treated as a variety of 
M. guttatus, while M. laxus should be considered as synonymous with 
M. guttatus. 

The chromosomes of three cultures from widely scattered populations 
(see table 1) of M. nasutus were counted. Two of the cultures have n=13 
chromosomes and the third has n=14, which confirms the previous de- 


242 MADRONO [Vol. 15 


TABLE 1. CHROMOSOME CoUNTS IN MIMULWUs, SECTION SIMIOLUS 


n=14 M. guttatus DC. 
Barry Summit, Humboldt County, California, altitude 3,400 feet, Keck 6007 
(5005). 
Yosemite Junction (rocky creek), Tuolumne County, California, altitude 1300 
feet, Hiesey 560 (5006). 
Lee Vining Canyon, Mono County, California, altitude 8000 feet, Clausen 2039 
(5014). 
Kern River, Kern County, California, altitude 1000 feet, L. Bean, April 16, 1949 
(5085). 
Botanic Garden strain, Hortus Cluj, Romania (5091). 
Rio Santo Thomas, Baja California, Mexico, altitude ca. 20 feet, C. and L. 
Hubbs, spring 1950 (5382). 
San Dimas Canyon, Los Angeles County, California, altitude 1500 feet, R.K. 
Vickery, Jr., September 29, 1950 (5678). 
Hugh’s Canyon, Salt Lake County, Utah, altitude 6000 feet, VN. Chamberlain, 
spring, 1952 (5836). 
Skagg’s Springs, Sonoma County, California, altitude ca. 50 feet, R. Holm, 
spring, 1951 (5864). 
Old Mine, Big Cottonwood Canyon, Salt Lake County, Utah, altitude 7650 feet, 
Vickery 683 (5961). 
Neff Canyon, Salt Lake County, Utah, altitude 5500 feet, D. Wiens, September 
6, 1956 (5995). 
Moab, Grand County, Utah, altitude 4100 feet, Vickery 762 (6080). 
Cane’s Spring, San Juan County, Utah, altitude 5800 feet, Vickery 763 (6081). 
Ledgemere, Big Cottonwood Canyon, Salt Lake County, Utah, altitude 5100 
feet, Vickery §80 (6082). 
East Creek, Morgan County, Utah, altitude 5700 feet, Vickery 883 (6083). 
n=14 M. tiling Regel var. tilingii 
Slate Creek (near Carnegie Transplant Garden), Mono County, California, alti- 
tude 10,000 feet, Clausen 2075 (5012). (In flower at time 6120, 6121, 6122 
were collected in bud.) 
Budd Lake, Tuolumne County, California, altitude 10,250 feet, C.W. Sharsmith, 
September 13, 1950 (5690). 
Tributary to Slate Creek (near Carnegie Transplant Garden), Mono County, 
California, altitude 10,050 feet, Vickery 1379 (6117). 
Slate Creek (near Carnegie Transplant Garden), Mono County, California, alti- 
tude 10,000 feet, Vickery 1382 (6120). (Light green, large leaves.) 
Same locality—Vickery 1383 (6121), dark green, medium sized leaves. 
Same locality—Vickery 1384 (6122), dark green, small leaves. 
n=14 M.laciniatus Gray 
Lake Eleanor road, Tuolumne County, California, altitude 4200 feet, Vickery 
179 (5339). 
n=14 M. laxus Pennell 
Yreka, Siskiyou County, California, altitude 3000-3200 feet, Pennell 26163 
(5658). 
n=14 M. nasutus Greene 
Hasting’s Reservation, Monterey County, California, altitude 1500 feet, Steb- 
bins 701 (5044). 
n=13 M. nasutus Greene 
San Augustine Pass, Dona Ana County, New Mexico, altitude 4500 feet, 
O. Norwell, October 30, 1946 (5018). 
Wild Cat Creek, near Yosemite Junction, Tuolumne County, California, alti- 
tude 475 feet, Vickery 168 (5327). 


1960] MUKHERJEE & VICKERY: MIMULUS 243 


n=15 M. platycalyx Pennell 
Crystal Lakes Reservoir, San Mateo County, California, altitude 800 feet, 
G. Oberlander, April, 1951 (5752). 
n=15 M. glabratus var. utahensis Pennell 
Johnson Pass, Tooele County, Utah, altitude 5800 feet, D. Wiens, October, 
1956 (6062). 
n=30 M. glabratus var. fremonti (Benth.) Grant 
Whipple Mountains, San Bernardino County, California, collected April 21, 
1940. U.C. 667,449 (5063). 
Kakernot Springs, Alpine Creek, Brewster County, Texas, Cory 53186, May 18, 
1946 (5373). 
n=32 M. tigrinus hort. 
Garden seed from the “Carlos Thays” Botanic Garden, Buenos Aires, Argentina 
(5081). 
n=32 M. luteus L. 
Vicinity of Illapel, Coquimbo, Chile, altitude 6200 feet, Plant Introduction and 
Exploration Division (U.S.D.A.) no. 144,535 (5042). 
n=30+0,1,or2 M. luteus L. 
Vicinity of Illapel, Coquimbo, Chile, altitude 2000 feet, Plant Introduction and 
Exploration Division (U.S.D.A.) no. 144,536 (5043). 


termination by G. L. Stebbins, Jr. (personal communictation) for the 
same culture. The n=13 cultures exhibit partial crossing barriers with 
M. guttatus, whereas the n=14 culture crosses readily with M. guttatus 
(Vickery, 1956, unpublished). The cytologic and crossing results suggest 
to us that M. nasutus as presently described (Grant, 1924, and Pennell, 
1951) includes at least two different entities. The proper naming of these 
entities must await further investigation and a detailed study of the taxo- 
nomic literature and type specimens of M. nasutus and and its relatives. 

Culture 5752, which was found to have n=15 chromosomes, was iden- 
tified as M. platycalyx with some misgivings. Although the plants clearly 
exhibit Pennell’s main key character of “fruiting calyces being fully as 
wide as long,” yet the corolla throats are open and not filled by “‘a palate 
nearly closing orifice’ (Pennell, 1951). Furthermore, Pennell had de- 
scribed M. platycalyx as occurring in the ‘‘southern Sierra Nevada from 
Mariposa to Tulare County, California,” whereas the plants from which 
culture 5752 were grown came from the Crystal Lakes region of the outer 
Coast Ranges of California. However, even if this culture does not prop- 
erly belong to M. platycalyx, it does represent an entity that is distinct 
from M. guttatus on the basis of morphology, crossing behavior (Vickery, 
1956, in press), and cytology. Here again a sound taxonomic decision 
must await further critical study of this entity and the literature. 

The culture of MW. glabratus var. utahensis (6062) from the Stansbury 
Mountains near the Great Salt Lake had n=15 chromosomes as did cul- 
ture 5265 from the population at Bicknell, Wayne County, Utah (Muk- 
herjee, Wiens, and Vickery, 1957). In contrast, the population from the 
shore of Mono Lake at the western edge of the Great Basin has n=14 
chromosomes (Vickery, 1955). Perhaps M. glabratus var. utahensis also 


244 MADRONO [Vol. 15 


consists of two morphologically similar but cytologically different entities 
as does M. nasutus, although corroborative crossing data is not yet 
available. 

Two cultures of WM. glabratus var. fremontii from southern California 
and Texas were found to have n=—30 chromosomes, although some of the 
plants of culture 5063 were observed to have as few as n=26 chromo- 
somes. This chromosome number fills an important gap in the polyploid 
series connecting the Great Basin form, M. glabratus var. utahensis, 
n=—14 and n=—15, with the South American M. glabratus var. parviflorus 
(Lindl.) Grant, n=45, and its ally M. pilosiusculus HBK., n=46 (Muk- 
herjee and Vickery, 1959). Mimulus glabratus var. fremontii is approxi- 
mately intermediate in appearance between the other two varieties. It is 
5 to 15 centimeters in height, whereas M. glabratus var. utahensis varies 
from 10 to 50 centimeters, and M. glabratus var. parviflorus is nearly 
prostrate. The leaves and flowers of M. glabratus var. fremontii are small- 
er than those of M. glabratus var. utahensis but larger than those of M. 
glabratus var. parviflorus. A broader cytogenetic and taxonomic study of 
the Mimulus glabratus complex of species is now being undertaken. 

The culture of M. tigrinus from the Botanic Garden of Buenos Aires, 
Argentina, was found to have n=32 chromosomes. This count agrees with 
the previous reports of Brozek (1932), Sugiura (Darlington and Wylie, 
1955) and the authors (1959). 

Chromosome counts obtained for two different cultures of M. luteus 
tend to support the previously indicated relationship of the horticultural 
species, M. tigrinus, to this wild species (Mukherjee and Vickery, 1959). 
Culture 5042 has n=32 chromosomes on the basis of two plants studied 
whereas culture 5043 was variable on the basis of the five or six plants 
examined. Of the twenty-three cells studied eleven had n=30 chromo- 
somes, six had n=31 and six had n=32. The cause of the variability was 
not clear from the data obtained. Perhaps accessory chromosomes are 
involved. Tentatively, the chromosome number for this culture appears 
to be n=30-+0, 1 or 2. The two cultures of M. luteus are morphologi- 
cally similar, but are distinguishable on the basis of flower markings and 
the general growth habits of the plants. 

In conclusion, this survey of chromosome numbers in section Szmiolus 
has verified previously published counts for M. guttatus (n=14), M. til- 
ingit var. tilingiti (n=14, M. glabratus var. utahensis (n=15) and M. 
tigrinus (n=32). It has shown that two species that are genetically close- 
ly related to M. guttatus, M.laciniatus and M.laxus, have n=14 chromo- 
somes also. In contrast, the culture, tentatively assigned to M. platycalyx, 
which is morphologically closely related to M. guttatus but genetically 
partially separated from it, has n=15 chromosomes. Mimulus nasutus 
appears to consist of two entities, one with n=14 chromosomes that is 
genetically closely related to M. guttatus, and the other with n=—13 
chromosomes that is genetically partially isolated from M. guttatus and 
from the n=14 form of M. nasutus. Mimulus glabratus var. fremontu 


1960] - GILLETT: PHACELIA 245 


was found to have n=30 chromosomes, which neatly fills an important 
gap in the polyploid series of M. glabratus var. utahensis, n=15, to M. 
glabratus var. parviflorus, n=45. Lastly, one race of South American 
M. luteus has n=32 chromosomes, as had been previously reported for 
its horticultural derivatives, but the other race apparently has n=30-+0, 
Ole 2: 


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


LITERATURE CITED 


Brozek, A. 1932. Mendelian analysis of the “red- orange-yellow” group of flower 

colors in Mimulus cardinalis Hort. Preslia 11:1-10. 

DaruincTon, C.D. and A. P.Wytiez. 1955. Chromosome atlas of flowering plants. 

Allen and Unwin Ltd. London. 519 pp. 

GranT, A. L. 1924. A monograph of the genus Mimulus. Ann. Missouri Bot. Gard. 

11:99-388. 

Hitcucook, C.L., A. Cronguist, M. OwnsBey and J. W.THompson. 1959. Vascular 
plants of the Pacific Northwest. Univ. Wash. Press, Seattle. 516 pp. 
MUKHERJEE, B. B. and R. K. VICKERY, JR. 1959. Chromosome counts in section 

Simiolus of the genus Mimulus (Scrophulariaceae). III. Madrono 15:57-62. 
MUKHERJEE, B. B., D. Wiens, and R. K. VICKERY, JR. 1957. Chromosome counts 

in 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. Phila. 99:155-199. 

. 1951. Jn Illustrated Flora of the Pacfic States by LeRoy Abrams. Stan- 

ford Univ. Press. Vol. III, pp. 688-731. 

Vickery, R.K., Jr. 1951. Genetic difference between races and species of Mimulus. 

Carnegie Institution of Washington Year Book 50:118-119. 

1955. Chromosome counts in section Simiolus of the genus Mimulus 
(Scrophulariaceae). Madrono 13:107-110. 

. 1956. Data on interracial and interspecific hybridization in the section 
Simiolus of the genus Mimulus (Scrophulariaceae). Utah Acad. Proc. 32:45-64. 
1959. Barriers to gene exchange between the species of the Mimulus 
guttatus complex. Evolution 13:300-310. 


FLOWERING RESPONSES IN PHACELIA SERICEA 
AND P. IDAHOENSIS! 


GEORGE W. GILLETT 


In the study of variation in the Phacelia sericea complex [P. sericea 
(Graham) A. Gray subsp. sericea; P. idahoensis Henderson; and inter- 
mediates], experimental cultures of P. sericea subsp. sericea and of P. 
idahoensis could not be brought into flower under actual or simulated 
summer conditions. In these cultures, the daily photoperiod was extended 
by incandescent lights, when necessary, to between 16 and 20 hours. 
Later cultures were brought into flower, however, by simulating the. fall 
conditions of the natural environment to the extent of materially reducing 


1 Supported by grant G-3886, National Science Foundation. 


246 MADRONO [Vol. 15 


Fic. 1. Geographical distribution of Phacelia sericea and Phacelia idahoensis. Races 
employed in experimental cultures: P. idahoensis (1), Valley Co., Idaho. P. sericea 
subsp. sericea: (G), Glacier Co., Montana; (GP), Routt Co., Colorado. Base map 
courtesy of University of Chicago Press. 


the length of the photoperiod and, at the same time, providing a cool 
temperature regime (Table 1). The data obtained from these cultures 
could be of value to those interested in growing alpine perennials experi- 
mentally, inasmuch as they show that certain alpines can be manipulated 
into flowering in a relatively short time. 

Both Phacelia sericea and P. idahoensis are spring-flowering perennials. 


1960] GILLETT: PHACELIA 247 


Early growth in each species is marked by the formation of a leaf rosette 
at ground level, with a virgate inflorescence axis later arising from the 
center of the rosette. In respect to geographical distribution (fig. 1), P. 
sericea occurs in the Rocky Mountains, the northern Cascades, the 
Olympics, and the ranges of the northern Great Basin. It is common at 
higher elevations, occurring on exposed gravel banks and talus slopes 
from 4500 feet, in the Canadian Rockies, to 13,000 feet in southern Colo- 
rado. By contrast, P. idahoensts is restricted to central Idaho, where it is 
found between 2800 and 7000 feet on wet meadows, stream banks, and 
on bottomlands subject to seasonal flooding. 

Limited field studies made on colonies of Phacelia sericea subsp. sericea 
have produced some information about the flowering response in this 
cordilleran subspecies. In several colonies examined in Colorado, Wyom- 
ing, and Montana, there was a pronounced tendency for early summer 
flowering, and for the simultaneous expression of a given phase of flower- 
ing by both large and small plants within a given colony. Furthermore, 
widely-separated colonies that were examined on the same day exhibited 
similar stages of inflorescence development. These points were convinc- 
ingly emphasized by an extensive and unsuccessful search for meiotic 
flower buds in several flowering colonies in Glacier County, Montana. In 
this area, and in Park County, Wyoming, colonies separated by as much 
as 1000 feet of elevation were found in similar stages of flowering. 

The examination of collection data from over 800 herbarium sheets of 
Phacelia sericea and P. idahoensis confirmed the above field observations. 
In both species, there is a very strong tendency to early summer flowering, 
and general uniformity in the stage of inflorescence development in 
specimens of a given collection. The sum total of this evidence would, 
therefore, suggest that flowering in these species is “triggered” by a 
broadly imposed environmental stimulus or stimuli, and it would tend to 
rule out the possibility that these are day-neutral species. 

Three races of these species were included in the present study. They 
were grown in experimental cultures (see Table 1) from seed collected 
in the following localities: 

Phacelia sericea subsp. sericea 
Gore Pass. Newly-graded road shoulder, highway 84, Routt County, 
Colorado, 9.9 miles west of Gore Pass, elevation ca. 8000 feet, 
Gillett 1145 (“GP” in fig. 1). 
Glacier. Gravel road shoulder, 0.8 miles west of Many Glacier Entrance 
Station, Glacier National Park, Glacier County, Montana, eleva- 
tion 4800 feet, Harry Robinson s.n, (“G” in fig. 1). 
Phacelia idahoensis 
Moist bottomlands 2.9 miles south of Donnelly, Valley County, Idaho, 
elevation 4800 feet, James Hockaday s.n. (“1” in fig. 1). 
Voucher specimens of experimental plants have been deposited at the 
herbaria of Michigan State University and the University of California, 
Berkeley. 


248 MADRONO [Vol. 15 


TABLE 1. FLOWERING RESPONSES IN PHACELIA IDAHOENSIS AND P. SERICEA 


I. Plants given long-day cycles only. 


A. Approximately 155 long-day cycles between October and March, 18-20 hour 
photoperiods. Temperatures between 10° and 25°C. 


No. Plants Flowered 
P.idahoensis 9 0 
P. sericea (Gore Pass) 8 0 
P. sericea (Glacier) 5 0 


B. Approximately 180 long-day cycles between April and September; with 
14—16-hour photoperiods. Temperatures between 15° and 30°C. 


P. idahoensis 20 ) 
P. sericea (Gore Pass) 10 ) 
P. sericea (Glacier) 10 0 


C. Approximately 250 long-day cycles between February and October; with 
14-16-hour photoperiods. Temperature between —3° and 35°C. 


P.idahoensis 8 0) 
P. sericea (Glacier) 4 @) 
II. Plants given long-day cycles (14-16 hr. photoperiods) followed by short-day 


cycles 8 
A. Approximately 110 long-day cycles (temp. regime of I.-C ) followed by 23 
short-day” cycles terminated by 22 cycles of 16-hour photoperiods under 
14°C. days and 7°C. nights, the 155-days-old plants then placed in a green- 
house (mid-July), with no supplementary light. 
Days to flowering after 


No. Plants Flowered short-day treatment 
P.idahoensis 8 8 16 
P. sericea (Glacier) 4 2 12 


B. Approximately 163 long-day cycles (temp. regime of I.-C) followed by 65 
short-day cycles, the 228-days-old plants then placed in a greenhouse (mid- 
September), with no supplementary light. 

P.idahoensts 3) 3 19 
P. sericea (Glacier) 2 1 16 

C. Approximately 169 long-day cycles (temp. regime I.-C) followed by 59 
short-day cycles, the 228-days-old plants then placed in a greenhouse (mid- 
September), with no supplementary light. 

P. idahoensts 3) 5 23 
P. sericea (Glacier) 2 i) 23 

4’ short-day cycles given inside a walk-in refrigerator with incandescent lights 

under temperature regime of 14°C. days and 7°C. nights. 

b 10-hour photoperiods. 

¢8-hour photoperiods. 


Culture techniques included germinating seeds in moist vermiculite, 
then transplanting young (ca. two weeks old) seedlings to four-inch pots 
holding equal parts of sterilized, screened river sand and peat moss. The 
plants were fed a nutrient solution (2 oz. commercial fertilizer per gallon 
of tap water) once a week through a siphon connection to the watering 
hose. 


1960] REVIEWS 249 


The light and temperature regimes provided for these cultures are 
given in Table 1. The results obtained indicate that Phacelia sericea 
subsp. sericea and P. idahoensis are neither day neutral nor “nominal”’ 
long-day plants. The positive flowering responses obtained in all cultures 
given the combination of cool temperatures and short photoperiods would 
suggest that these are obligate short-day species; although the possibility 
of their being conditioned for flowering by low temperatures, independent 
of day length, remains very strong. 

In a final experiment, seven plants of Phacelia idahoensis that had 
been held to long photoperiods of from 14 to 16 hours for 264 days were 
placed in an open cold frame and exposed to the late fall light and tem- 
perature regime of central Michigan. These conditions included temper- 
atures ranging from +10° to —10°C. After 37 days of ‘‘outside”’ weather, 
these plants, the pots frozen solid, were removed to the greenhouse. Six 
of the seven plants produced inflorescences and flowered within a month. 
These inflorescences were formed in a greenhouse where unaltered De- 
cember lighting conditions prevailed, demonstrating (as indicated in II-C 
of Table 1) that this species does not require, subsequent to induction, a 
long-day regime for flowering. It would seem practicable, therefore, to 
culture this species by a schedule that would include spring germination, 
and a fall induction period under a cool temperature regime, with the 
prevailing light of approximately 40° north latitude. It is probable that 
these suggestions also apply to P. sericea subsp. sericea inasmuch as its 
flowering response is similar. A close relationship is indicated by the 
genetic compatibility between the two taxa, the F, hybrids being highly 
fertile and also flowering after short-day induction treatments. 

In addition to providing knowledge for the successful culture of these 
and probably other species of alpine perennials, these experiments leave 
a pointed suggestion for plant geographers, namely, that Phacelia sericea 
subsp. sericea, a northern alpine perennial, has, in terms of photoperiod 
requirements, southerly rather than arctic affinities. This would not be 
surprising in view of the fact that the great bulk of Phacelia species are 
found south of 40° north latitude, while only two species occur farther 
north than P. sericea subsp. sericea. 


Department of Botany and Plant Pathology 
Michigan State University, East Lansing 


REVIEWS 


Illustrated Flora of the Pacific States. Washington, Oregon, and California. 
Volume IV—Bignoniaceae to Compositae. By ROXANA STINCHFIELD FERRIS. xiii + 732 
pages, 1124 figures, and an appendix (for all four volumes) with key to families, 
index to common names, and index to scientific names. Stanford University Press, 
Stanford, California. 1960. $17.50. 

In 1923, with the publication of Volume I, Ophioglossaceae to Aristolochiaceae, 
of the “Illustrated Flora of the Pacific States,” Leroy Abrams launched his life’s 
work—-an illustrated, descriptive flora of all vascular plants known to grow wild in 
the three Pacific states—Washington, Oregon, and California. Now, thirty-seven 


250 MADRONO [Vol. 15 


years later, with the publication on January 22, 1960, of Volume IV, Bignoniaceae 
to Compositae, by Roxana Stinchfield Ferris, the monumentally conceived project 
is completed and a milestone in the floristic botany of the western states is achieved. 
Although the task outlasted the life-time and efforts of Professor Abrams, who died 
in 1956 after about eight years of failing health, he was very fortunate in having the 
collaboration and devoted assistance of Mrs. Ferris almost from the beginning of 
the project, and progress on the flora continued uninterruptedly through the years. 
Mrs. Ferris gave increasing effort to Volume II, Polygonaceae to Krameriaceae, pub- 
lished in 1944, and the task fell upon her of finishing Volume III, Geraniaceae to 
Scrophulariaceae, published in 1951. She had the entire responsibility for Volume IV. 
That the finished work ably carries out the objectives stated in Volume I—“to furnish 
an authentic reference book that will be of greatest service not only to the trained 
botanist but to everyone interested in the native plant life of the Pacific States.”— 
is tribute not only to Professor Abrams’ vision and high level of scholarship but also 
to Mrs. Ferris’ devoted and able efforts toward full realization of his goals. 


The “Illustrated Flora of the Pacific States” was designed to duplicate for western 
botany what Britton and Brown, in their “An Illustrated Flora of the Northern 
United States, Canada and the British Possessions,” had done for eastern botany. 
These are the only two great North American floristic works in which all species 
are illustrated and in which the illustrations share along with keys and descriptions 
the task of species identification. Abrams not only followed the general pattern set by 
Britton and Brown, but, at first, also followed them in their use of the “American 
Code” for nomenclature. With the adoption of the International Code after the 
international congresses of 1930 and 1935, however, the two “codes” were brought 
almost into accord, and in the 1940 reprinting of Volume I of his flora, Abrams 
made such generic changes as were necessary to bring his nomenclature into con- 
formity with the International Rules. 

In all four volumes specialists were invited to contribute the text for certain 
plant groups, and in Volume IV Mrs. Ferris had the able assistance of ten such col- 
laborators. In addition she was fortunate in being able to base her treatment of the 
Compositae as a whole upon extensive manuscript notes from the late Dr. Sidney 
Fay Blake, who had originally planned to contribute the entire text for this family. 
Mrs. Ferris wrote the major part of the volume herself, however, and, in many 
groups such as Baeria, Plantago, and Galium, to name a few, she made significant 
contributions toward clarification of our understanding of the species. 


The illustrations deserve especial consideration inasmuch as they are an integral 
part of the plan of the entire work. In Volume IV, following the general plan of 
preceding volumes, the illustration for each species consists of a group of carefully 
arranged line drawings occupying a rectangle about 2 x3 inches. There may be two 
or three or as many as eight separate drawings artistically combined within the given 
rectangle in such a way as to give both overall aspect or habit of the plant and 
significant structural details, a combination necessitating much ingenuity and skill. 
These illustrations of species are then grouped six to a page, or in some cases two or 
four illustrations may occupy a third or two-thirds of a page. It is stated in the 
preface that the illustrations, except for the structural features, are half size unless 
otherwise marked. Legends are confined to binomials, but the pertinent information 
in the carefully worded text is close at hand, the illustrations and descriptions thus 
complementing each other and obviating the need for detailed legends. Most of the 
drawings in Volume IV were made by Jeanne Russell Janish, who was also the 
artist for many of the illustrations in volumes II and III. Many of the drawings of 
the Compositae in Volume IV, however, were made by Doris Holm Blake while her 
husband, Sidney Fay Blake, was working upon what he then hoped would culminate 
in his full text for the Compositae. The drawings for Agoseris and Helianthus were 
made by other artists under supervision of the specialists who contributed the texts 
for these genera. Except for a few cases where detail is obscured by the illustrations 
being overly black, reproduction is excellent. 


1960] REVIEWS zat 


Throughout the volume the keys are skillfully and evenly handled despite the 
number of different contributors. There are some unfortunate instances of the use of 
a negative rather than a truly opposing phrase in the second branch of a key 
dichotomy, but understandably these instances occur particularly in the “difficult” 
groups. One of the major tasks in preparation of Volume IV was the assembling 
of an appendix for all four volumes. This contains 1) a key to the families, 2) an 
index of common names, and 3) an index to scientific names. The family key gives 
not only the family numbers but also the volume and page on which each family 
is found, a very necessary aid in a work of this magnitude. The index to common 
names (there is a common name for every species treated in the four volumes) has 
the family names printed in small capitals and the genera in Roman type. The index 
to scientific names is much longer and more complicated than that to common names, 
having approximately 17,500 entries occupying 79 pages. It has the names of families 
and tribes printed in small capitals, the genera, species, subspecies, and varieties 
in Roman type, and the synonyms in italics, all appropriately indented. Because some 
groups have a great number of species as well as many generic and specific synonyms, 
the genera in the index to scientific names are not always easy to locate. Possibly 
greater indentation or perhaps the use of boldface type for generic names would 
have made them stand out more, although to do this would have necessitated a 
departure from the style of the previous volumes. 

The Stanford University Press has achieved another outstanding accomplishment 
in typography, printing, and binding, and the volume contains a minimum of typo- 
graphical and other mechanical errors. 

Dr. Bacigalupi, curator of the Jepson Herbarium, has given me permission to 
quote from the unpublished field notebook of Willis Linn Jepson, whose entry for 
February 3, 1910, reads: “I am just receiving the first reviews of my Flora of 
California, Pts. 1 and 2. The critics mostly or even entirely confine themselves to 
verbal slips, not touching general principles. It is, to be sure, disconcerting enough 
to have such errors, but after all the main thing is this: ‘Has the book got matter 
in it? Has it got stuff in it? Is it meaty? Not is it fazltless. A faultless book is 
impossible. It is inevitable in the nature of the human mind that such slips will be 
made, mistakes and blunders. But is the job a big one, is it really worthwhile? So 
satisfied am IJ in the affirmative that it is a big task, to be done in a big way, without 
too much considering the danger of possible minor errors, that I go on, to finish up 
my job, just as other big jobs have been finished aforetime.’ ” 

All will agree that Mrs. Ferris’ job has got matter, stuff, and meat in it, that it 
was a big task, done in a big way.— HELEN K. SHARSMITH, Department of Botany, 
University of California, Berkeley. 


Experimental Studies on the Nature of Species. lV. Genetic Structure of Ecological 
Races. By JENS CLAUSEN and W. M. HieseEy. Carnegie Institution of Washington 
Publication 615. Washington, D.C. Octavo, vii + 312 pp., 33 figs. 1958. Paper $4.25, 
cloth $4.75. 

This is the fourth in the series of scholarly monographs based on the studies of 
plant evolution conducted by these authors over the past three decades. This newest 
volume expands the earlier work on the evolutionary importance of ecological races 
by considering in detail the genetics of the altitudinal races of Potentilla glandulosa 
and then reviewing examples from the literature on the genetic structure of 
ecological races. 

The volume is organized into five chapters and although these are skillfully inter- 
related they are sufficiently distinct and different to require individual comment. 
Chapter I, Ecological Races of P. glandulosa, introduces the general topic of the 
volume by presenting what might be called the systematics of P. glandulosa as it 
occurs along the altitudinal transect across central California. This adroitly prepared 
chapter makes it possible to read the work without reviewing the previous publica- 
tions by these authors on P. glandulosa. Chapter II, Genetics of Ecological Races, 


252 MADRONO [Vol. 15 


comprises nearly one third of the entire work. The first portion describes the crosses 
made between selected plants of the various climatic races of P. glandulosa by giving 
in detail the characteristics of the parental plants and the segregation of these charac- 
teristics in the Fj, F2 and Fs generations. Of particular interest here is the use of 
punched cards for recording and analyzing the data on 14 different characteristics of 
each individual plant in these crosses. The inherent nature of the punched card sys- 
tem gave an index number series for each character and a summation of these gave 
an index value for each plant that proved useful in the general comparison of parents 
and their progenies. Frequency distribution of parent index values and hybrid index 
values are presented and give a picture of the spectrum seen in the segregation of 
the Fs from the crosses between the contrasting ecological races. The second section 
of Chapter II presents analyses of the segregation ratios by proposing gene systems 
that could account for the complex ratios observed. These analyses are detailed to 
the point of proposing for each characteristic the number of loci involved, the number 
of alleles at each locus, and the action and interactions of the various genes. The 
significance of these proposed gene systems lies not in their accuracy as to details 
but rather lies in the fact that viewed collectively they demonstrate that the differ- 
ences between the ecological races are controlled by units of segregation and recom- 
bination that can be described in terms of classical genetics. These points are clearly 
expressed in the concluding chapter. 

Chapter III, Response Patterns at Contrasting Altitudes, analyzes the responses 
of cloned individuals of an Fo between two ecological races to the different environ- 
ments of the transplant stations. Studying such an Fy» under different natural 
environments leads the authors to estimate the evolutionary potential of segregat- 
ing populations. They conclude this important chapter with the following: “The 
present races are the products of long-time selection, and have attained an equilibrium 
with their environments. Natural selection will therefore tend to favor the original 
racial combination as long as the over-all genetic structure and the habitats remain 
the same, although a certain amount of introgression may take place. Over long 
periods genes may gradually migrate across long distances from the original point of 
contact and may finally appear in combination where they have selective value.” 

Chapter IV, Systems of Genes Controlling Characters and their Significance in 
Environmental Adaptation and Evolution, is the longest chapter comprising over 
one third of the text material. It differs markedly from the previous three chapters 
in that it does not present new data but rather reviews a considerable segment of 
the genetic literature dealing with gene systems. The relevance of these reviews to 
the previous chapters lies in the fact that the gene systems discussed are of the same 
general sort as the gene system for P. glandulosa. Chapter V, Concepts of the Genetic 
Structure of Ecological Races, develops a general concept of the genetic structure and 
evolutionary importance of ecological races. 

In the tradition of the previous volumes a vast amount of the original data are 
presented in a tabular form and the same precise, clear writing and excellent illustra- 
tion are evident. 

The strength of this book clearly lies in the work on P. glandulosa. Hybridization 
studies and the observation of the responses of cloned individuals to different natural 
environments are two of the powerful tools of evolutionists. In combining both of 
these approaches in the study of P. glandulosa the authors present a new dimension 
of information about natural populations. At this moment we cannot predict the 
amount of influence this publication will have on our understanding of evolution. 
We can be sure, however, that it will remain the classic work of its kind for many 
years because the time, facilities, and skills necessary for this type of study are 
available to few botanists —HENnry J. THompson, Department of Botany, University 
of California, Los Angeles. 


1960 ] INDEX 253 


INDEX TO VOLUME XV 


Classified entries: Chromosome numbers, Reviews. New scientific names are in 
boldface type. Un-annotated taxa in floral lists are omitted from Index. 


Acer circinatum, 227 

Alliaria officinalis, 96 

Allium: macrum, 226; nevii, 226; tolmiei, 
227 

Angelica: canbyi, 227; genuflexa, 230 

Anthriscus scandicina, 229 

Arceuthobium: americanum, 130; cam- 
pylopodum, 132; douglasii, 137; dwarf 
mistletoes, in California, 129 

Arnott, H. J., Vivipary in Cordyline 
australis Hook., 71 

Aster: New combinations in, 128; occi- 
dentalis var. delectabilis, 128, var 
parishii, 128 

Atriplex confertifolia, 227 


Bacigalupi, R.: G. Thomas Robbins, 231 

Baker, M. S., Studies in western violets, 
IX, Sections Nomimium and Chamae- 
melanium, 199 

Beckmannia syzigachne, 64 

Benson, L., Typification of Prosopis 
odorata Torr. & Frem. 53 

Bupleurum rotundifolium, 64 


Campanula scouleri, 228 

Cardamine oligosperma, 229 

Cardiocarpon late-alatum, 180, fig. 182 

Cardiospermum halicacabum, 64 

Carex: aperta, 229; eurycarpa, 226 

Carter, A., Urospermum picroides (L.) 
Schmidt in Berkeley, 224 

Ceanothus: cordulatus, 80; integerrimus, 
80; seeds and seedlings on burns, 79 

Choreocolax: margaritoides, 33; poly- 
siphoniae, 33; suhriae, 33 

Chromosome numbers: Agastache urtici- 
folia, 51; Amsonia brevifolia, 50, var. 
tomentosa, 50; Astragalus bolanderi, 
82, crotalariae, 82, leucopsis, 82, 
oocarpus, 82, pomonensis, 82; Blenno- 
sperma nanum var. robustum, 220; 
Cathartolinum digynum, 49; Chaenac- 
tis douglasii, 52; Chamaesaracha nana, 
51; Chrysopsis foliosa, 52; Coldenia 
canescens, 50, palmeri, 50, plicata, 
50, purpusii, 50; Cryptantha 
pterocarya, 50; Datisca glomerata, 
82; Datura meteloides, 83; Delphin- 
ium carolinianum, 219, polycladon, 
82, virescens var. macroceratilis, 219; 
Dithyrea californica, 49; Encelia fru- 
tescens, 220; Epilobium obcordatum, 
82; Erysimum capitatum var. beali- 
anum, 49; Eschscholtzia mexicana, 82; 
Fraxinus velutina, 161, var. coriacea, 
161; Fritillaria biflora, 82; Garrya 
lindheimeri, 220; Gayophytum helleri 


var. glabrum, 91, humile, 91, lasiosper- 
mum, 91, nuttallii, 91, var. abramsii, 
91, racemosum, 82, var. erosulatum, 
91, ramosissimum, 92; Glandularia 
canadensis, 51, gooddingii, 50, perackii, 
50; Haplopappus ciliatus, 52, divari- 
catus, 52, hartwegii, 52, havardii, 220, 
phyllocephalus subsp. annuus, 220, 
subsp. phyllocephalus, 220, pleuriflorus, 
52, spinulosus, 220; Helianthus 
ciliaris, 57, crenatus, 57; Heliotropium 
parviflorum, 50; Hesperocallis un- 
dulata, 49; Isomeris arborea, 82; 
Lupinus superbus var. elongatus, 82; 
Lychnis drummondii, 205, wilfordii, 
206; Lyrocarpa palmeri, 49; Machaer- 
anthera blephariphylla, 221, gymno- 
cephala, 52, tagetina, 221, tanacetifolia, 
52; Marah macrocarpus, 52; Mimulus 
glabratus var. fremontii, 243, var. 
parviflorus, 59, var. utahensis, 243, 
glaucescens, 59, guttatus, 59, 242, lacini- 
atus, 242, laxus, 242, luteus, 61, 243, 
nasutus, 242, pilosiusculus, 59, platy- 
calyx, 243, tigrinus, 59, 243, tilingii 
var. corallinus, 59, var. tilingii, 59, 242; 
Monardella cinerea, 51, lanceolata, 51, 
linoides, 51, odoratissima, 82 ; Myosotis 
versicolor, 50, aristatus subsp. mon- 
tanus, 140, cupulatus, 140, minimus, 
139, var. filiformis, 140, subsp. apus, 
140, sessilis, 140, subsp. alopecurioides, 
140; Neptunia dimorphantha, 185, gra- 
cilis, 185, lutea, 185, monosperma, 185, 
plena, 185, prostrata, 185, pubescens 
var. floridana, 185, var. lindheimeri, 
185, triquetra, 185; Nolina parryi, 49; 
Oenothera californica, 82; Orthocarpus 
attenuatus, 51, linearilobus, 51, pur- 
purascens, 51; Pedicularis  groen- 
landica, 83; Penstemon clutei, 220; 
Petrocoptis pyrenaica, 206, 215; Plan- 
tago insularis, 83; Porophyllum 
scoparium, 221; Proboscidea jussieui, 
83; Pseudotaenidia montana, 220; 
Psilostrophe cooperi, 221; Ranunculus 
californicus, 82, occidentalis var. 
eisenii, 49; Ratibida columnaris, 52, 
peduncularis var. picta, 52, tagetes, 
52; Saxifraga ferruginea, 219, integri- 
folia, 219, montanensis, 219, occiden- 
talis, 219, tolmiei, 220; Sclerocarpus 
uniserialis, 221; Senecio fremontii var. 
occidentalis, 83, longilobus, 52; Silene 
antirrhina, 206, aperta, 205, bridgesii, 
173, 205, californica, 205, campanu- 
lata, 173, 205, caroliniana, 206, subsp. 
wherryi, 206, clokeyi, 205, douglasii, 


254 MADRONO 


205, grayi, 205, hookeri, 205, invisa, 
205, keiskei, 206, laciniata subsp. 
greggii, 205, subsp. major, 205, lem- 
monii, 173, 205, marmorensis, 173, 176, 
206, menziesii, 206, montana, 206, 
nuda, 206, subsp. insectivora, 206, oc- 
cidentalis, 206, oregana, 206, parishii, 
206, parryi, 206, petersonii, 206 poly- 
petala, 206, regia, 206 repens subsp. 
australis, 206, var. latifolia, 206, ro- 
tundifolia, 206, sargentii, 206, scaposa, 
206, scouleri, 206, spaldingii, 206, 
stellata, 206, struthioloides, 206, sub- 
ciliata, 206, thurberi, 206, verecunda 
subsp. andersonii, 206, subsp. platyota, 
206, subsp. verecunda, 206, virginica, 
206, williamsii, 206, wrightii, 206; 
Solanum xantii, 51; Streptanthus in- 
flatus, 49; Thelypodium lasiophyllum, 
82; Verbena bracteata, 51, 220, hastata, 
220, prostrata, 51, stricta, 220, urtici- 
folia, 220; Viguiera adenophylla, 221, 
deltoidea var. parishii, 221, dentata 
var. brevipes, 221, porteri, 221, steno- 
loba, 221; Viola bakeri subsp. bakeri, 
204, subsp. shastensis, 204, praemorsa 
subsp. arida, 204, subsp. oregona, 204; 
Zigadenus brevibracteatus, 49; Zinnia 
acerosa, 216, citrea, 216, grandiflora, 
52, 216 

Chromosome numbers of plants, Docu- 
mented, 49, 82, 219 

Clarkson, Q. D., Field studies of natural 
hybridization in the Oregon species of 
Iris L. subsec. Californicae Diels, 115 

Copeland, H. F.: The reproductive struc- 
tures of Fraxinus velutina (Oleaceae), 
161; The reproductive structures of 
Schinus molle (Anacardiaceae), 14 

Cordyline: australis, 71; Vivipary in, 71 

Crampton, B., The grass genera Orcuttia 
and Neostapfia; a study in habitat and 
morphological specialization, 97 

Cucurbita: interspecific cross in, 4; lun- 
delliana, 4, fig. 6, fig. 8, fig. 10; 
moschata, 4, fig. 6, fig. 8, fig. 10 

Cupressus: abramsiana, 1; goveniana, 
1; macrocarpa, 3; pygmaea, 1; sar- 
gentil, 1; in pygmy forests of Mendo- 
cino County, 1 

Dasystephana newberryi, 234 

Datura meteloides, 87, fig. 85 

Davidson, J. F., Cleared Cardiocarpon 
late-alatum Lesq. Cordaitean seeds 
from Michigan, 180 

Dixon, D. G., Chromosome counts in the 
genus Gayophytum, 90 


Echinodorus berteroi, 64 


Fan, K.-C., and G. F. Papenfuss, Red 
algal parasites occurring on members 
of the Gelidiales, 33 


[Vol. 15 


Fearing, O. S., and B. L. Turner, The 
basic chromosome number of the genus 
Neptunia, 184 

Ferris, R. S., 
Aster, 128 

Flint, F. F., Development of the maga- 
gametophyte in Liquidambar styraci- 
flua L., 25 

Fraxinus velutina, 161, fig. 163, fig. 165, 
fig. 167; The reproductive structures 
of, 161 

French, D. H., and R. Ornduff, Distribu- 
tional notes on plants of the Warm 
Springs area, Oregon, 225 

Fritillaria biflora, 83, fig. 85 


New combinations in 


Gayophytum: Chromosome counts in, 
90; helleri var. glabrum, 92 

Gelidiales, Red algal parasites of, 33 

Gelidiocolax: mammillata, 34, figs. 35- 
37; margaritoides, 34; microsphaerica, 
33; suhriae, 33 

Gentiana: copelandii, 234; eximia, 234; 
newberryi, 234; newberryi and G. 
tiogana, Studies in the _ perennial 
gentians, 233; tiogana, 235 

Gillett, G. W.: Flowering responses in 
Phacelia sericea and P. idahoensis, 245 

Gray, J., Review, Pollen and _ spore 
morphology/plant taxonomy, 62 

Greene, Edward Lee, Correspondence, 
Zee, 


Haller, J. R., Factors affecting the dis- 
tribution of Ponderosa and Jeffrey 
pines in California, 65 

Helianthus: ciliaris, 54; crenatus, 55; 
heiseri, 54; Two new species of, from 
New Mexico, 54 

Hoffman, Freed, 178 


Iris: bracteata, 120; bracteata x chry- 


sophylla, 121; chrysophylla, 116; 
douglasiana, 119; douglasiana xX in- 
nominata, 120; innominata, 119; 


Field studies of natural hybridization 
in the Oregon species of, subsection 
Californicae Diels, 115; tenax, 116, 
subsp. bracteata, 121, subsp. chry- 
sophylla, 121, subsp. douglasiana, 122, 
subsp. innominata, 122, subsp. tenax, 
121, subsp. thompsonii, 122; subsec- 
tion Californicae, hybridization in, 
115; thompsonii, 120 
Isomeris arborea, 84, fig. 85 


Jackson, R. C., Two new species of 
Helianthus from New Mexico, 54 

Jeffrey and Ponderosa pines: A com- 
ment on cold susceptibility of, 217; 
Distribution of, in California, 65 

Die Nadelge- 


Kasapligil, B., Review, 


holze, 30 


1960] 


Kruckeberg, A. R.: A new Silene from 
northwestern California, 172 ; Chromo- 
some numbers in Silene (Caryophyl- 
laceae), II, 205 

Kuijt, J.; Occurrence of Pilostyles thur- 
beri in California, 63; The distribu- 
tion of dwarf mistletoes, Arceutho- 
bium, in California, 129 


Liquidambar styraciflua, 25, figs. 26-27; 
megagametophyte development, 25 

Lomatium: cous, 227; leptocarpum, 227; 
nevadense, 228 

Lupinus superbus var. elongatus, 86 

Lychnis wilfordii, fig. 211 


Major, J., Review, Carex, its distribu- 
tion and importance in Utah, 221 

Mason, C. T., Jr.: Notes on the flora of 
Arizona, 64; Studies in the perennial 
gentians, G. newberryi and tiogana, 233 

Mathias, M. E., and Raven, P. H., Sani- 
cula deserticola, an endemic of Baja 
California, 193 

Maule, S. M., Xerophyllum _ tenax, 
squawgrass on Mount Rainier, 39 

McIntosh, R. P., Edward Lee Greene 
correspondence, 222 

McMillan, C., Survival of transplanted 
Cupressus in the pygmy forests of 
Mendocino County, California, 1 

McVaugh, R., Review, Spring flora of 
the Dallas-Fort Worth area, 94 

Mertensia: ciliata, 123, Variation pat- 
terns in, 123; fusiformis, 123 

Meyer, F. G., A new species of Valeriana 
from Brazil, 197 

Mimulus: Chromosome counts in sec- 
tion Simiolus of, 57, 239; glabratus 
var. fremontii, 243, var. parviflorus, 
58, var. utahensis, 243; glaucescens, 58; 
guttatus, 58, 242; laciniatus, 242; 
laxus, 242; luteus, 61, 243; nasutus, 
242; pilosiusculus, 58; platycalyx, 243; 
tigrinus, 58, 243; tilingii, 242, var. 
corallinus, 58, var. tilingii, 58 

Mirov, N. T., Review, The physiology 
of forest trees, 192 

Morrison, J. L., Freed Hoffman, 178 

Mukherjee, B. B., and R. K. Vickery, 
Jr.: Chromosome counts in the sec- 
tion Simiolus of the genus Mimulus 
(Scrophulariaceae), III, 57; IV, 239 

Myosotis micrantha, 229 

Myosurus: aristatus, fig. 146; cupulatus, 
fig. 141, fig. 146; minimus, fig. 144, 
subsp. apus, fig. 144, var. filiformis, 
fig. 141, fig. 145; Nuclear cytology, 
139; sessilis, figs. 141-142 


Neosecotium: 154; africanum, 156; 
macrosporum, 154, fig. 155; and Se- 
cotium, 152 


INDEX 299 


Neostapfia: 97; and Orcuttia, 97 

Neptunia, The basic chromosome num- 
ber of the genus, 184 

Nicotiana: palmeri, 148; trigonophylla, 
148; Variation in section Trigono- 
phyllae, 148 

Notes and News, 32, 63, 96, 128, 160, 
192, 222 


Oenothera californica var. californica, 87 

Orcuttia: and Neostapfia, habitat and 
morphological specialization, 97; cali- 
fornica var. inaequalis, 99, var. vis- 
cida, 103; greenei, 102; mucronata, 
107, figs. 106-107; pilosa, 99; tenuis, 
103 

Ornduff, R.: Alliaria officinalis in Ore- 
gon, 96; Review, Spring flowers of 
lower Columbia Valley, 94; and D. H. 
French, Distributional notes on plants 
of the Warm Springs area, Oregon, 225 


Panicum pacificum, 226 

Papenfuss, G. F., and Fan, K.-C., Red 
algal parasites occurring on members 
of the Gelidiales, 33 

Pelton, J. S., Variation patterns in four 
clones of Mertensia ciliata, 123 

Phacelia: idahoensis, 247; sericea, 247; 
sericea and idahoensis, Flowering re- 
sponses in, 245 

Picea: engelmannii, 111, and P. glauca, 
The taxonomic relationship between, 
111; glauca, 111, subsp. engelmannii, 
114, subsp. glauca, 114 

Pilostyles thurberi in California, 63 

Pinus: jeffreyi, 65, 217; ponderosa, 65, 
217 

Pneumonanthe newberryl, 234 

Ponderosa and Jeffrey pines: cold sus- 
ceptibility of, 217; Distribution of, 
in California, 65 

Prosopis: glandulosa, 53; juliflora var. 
glandulosa, 53, var. torreyana, 53, var. 
velutina, 53; odorata, 53, Typifica- 
tion of, 53; pubescens, 53 

Pterocladiophilaceae, 38 

Pterocladiophila hemisphaerica, 34, 
figs. 35-36 

Pteryxia terebinthina, 228 


Quick, C. R., Ceanothus seeds and seed- 
lings on burns, 79 


Raven, P. H., and Mathias, M. E., Sani- 
cula deserticola, an endemic of Baja 
California, 193 

Reviews: Clausen, J., and W. M. Hiesey, 
Experimental studies on the nature of 
species, IV, Genetic structure of eco- 
logical races, 251; Erdtman, Pollen 
and spore morphology/plant taxon- 
omy, 62; Ferris, R. S., Illustrated flora 


256 MADRONO 


of the Pacific States—Washington, 
Oregon, and California, Volume IV, 
Bignoniaceae to Compositae, 249; 
Foster, A. S., and Gifford, E. M., Jr., 
Comparative morphology of vascular 
plants, 158; Hill, Spring flowers of 
the lower Columbia Valley, 94; 
Howell, J. T., P. H. Raven, and P. 
Rubtzoff, A flora of San Francisco, 29; 
Kriissmann, G., Die Nadelgeholze, 30; 
Lewis, M. E. Carex, its distribution 
and importance in Utah, 221; Shin- 
ners, Spring flora of the Dallas-Fort 
Worth area, 94; Thimann, K. V., et 
al, editors, The physiology of forest 
trees, 192 

Robbins, G. Thomas, 231 

Rubus laciniatus, 228 


Salix drummondiana var. subcoerulea, 
227 

Sanicula: bipinnatifida, 194; deserticola, 
193, endemic of Baja California, 193 

Schinus molle, 14, fig. 15, fig. 17, fig. 20, 
The reproductive structures of, 14 

Secotium: aurantium, 73; gueinzii, 152, 
table 157; macrosporum, 153; and 
Neosecotium, Studies on secotiaceous 
fungi VII, 152; tenuipes, 73 

Senecio: fremontii var. occidentalis, 88, 
fig. 85; macounii, 228 

Setchelliogaster: 74; 
tenuipes, 74, fig. 77 

Sharsmith, H. K.: Review, Ilustrated 
flora of Pacific States, Vol. IV, 249 

Silene: antirrhina, fig 211; aperta, fig. 
209; bridgesii, 173; californica, 207; 
campanulata, 173, 210; caroliniana, 
fig. 211, subsp. wherryi, fig. 211; 
Chromosome numbers in, 205; clokey, 
fig. 209; douglasii, 210; grayi, fig. 209; 
hookeri, 207; invisa, fig. 209; keiskei, 
fig. 211; laciniata subsp. laciniata, 207, 
subsp. greggii, 207, fig. 209, subsp. 
major, 207; lemmonii, 173, fig. 209; 
marmorensis, 174, figs. 175—177; 
menziesii, 208; montana, fig. 209; 
New, from northwestern California, 
172; nuda, 213, subsp. insectivora, 213; 
occidentalis, 213, fig. 209; oregana, 
213; parishii, 213; parryi, 208; peter- 
sonii, fig. 209; polypetala, fig. 211; 
regia, fig. 211; repens subsp. australis, 
fig. 209, var. latifolia, fig. 211; rotundi- 
folia, fig. 211; sargentil, 213; scaposa, 
fig. 209; scouleri, 214; seeleyi, 210; 
spaldingii, fig. 209; struthioloides, fig. 
211; subciliata, fig. 211; thurberi, fig. 
211; verecunda subsp. andersonii, 214, 
subsp. platyota, 214, subsp. verecunda, 
214, subsp. virginica, 215; williamsii, 
210, fig. 211; wrightii, fig. 211 

Smith, A. H., and R. Singer: Studies on 


aurantium, 76; 


[Vol. 15 


secotiaceous fungi VI, Setchelliogaster 
Pouzar, 73; VII, Secotium and Neo- 
secotium, 152 

Snow, R., Chromosome numbers of Cali- 
fornia plants, with notes on some cases 
of cytological interest, 81 

Stone, D. E., Nuclear cytology of Cali- 
fornia mouse-tails (Myosurus), 139 

Stylocline filaginea, 228 

Syringocolax macroblepharis, 33 


Taylor, T. M. C., The taxonomic rela- 
tionship between Picea glauca 
(Moench) Voss and Picea engelmannii 
Parry, 111 

Tepfer, S., Review, Comparative morph- 
ology of vascular plants, 158 

Tetradymia glabrata, 228 

Thomas, J. H., Review, A flora of San 
Francisco, 29 

Thompson, H. J., Review, Experimental 
studies on nature of species, IV, 251 

Torres, A. M., A new species of Zinnia 
from Mexico, 215 

Turner, B. L. and O. S. Fearing, The 
basic chromosome number of the genus 
Neptunia (Leguminosae — Mimo- 
soideae,), 184 


Urospermum picroides, 224 


Valeriana: A new species of, from Brazil, 
197; glechomifolia, 197, fig. 198 

Vickery, R. K., Jr., and B. B. Mukher- 
jee: Chromosome counts in the sec- 
tion Simiolus of the genus Mimulus 
(Scrophulariaceae), III, 57; IV, 239 

Viola: adunca, 202; aliceae, 199, fig. 
201; bakeri, 203, subsp. shastensis, 
203; lobata subsp. psychodes, 200, 
fig. 201; oxyceras, 202, fig. 201; 
praemorsa, 204; psychodes, 200 


Wagener, W. W., A comment on cold 
susceptibility of Ponderosa and Jef- 
frey pines, 217 

Webber, J. M., Hybridization and in- 
stability of Yucca, 187 

Wells, P. V., Variation in section Trigono- 
phyllae of Nicotiana, 148 

Whitaker, T. W., An interspecific cross 
in Cucurbita (C. lundelliana Bailey 
x C. moschata Duch.) 4 

Xerophyllum tenax on Mount Rainier, 39 


Yucca: arizonica, 188, fig. 189, & glauca, 
190, fig. 189, & neomexicana, 190, fig. 
189; constricta, 188, fig. 189, x 
schidigera, 188, fig. 189; elata, 188; 
glauca, 190, fig. 188, x elata, 188; 
Hybridization and instability of, 187; 
neomexicana, 188; schidigera, 188 


Zinnia: A new species of, from Mexico, 
215; acerosa, 216; citrea, 215; grandi- 
flora, 216 


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. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


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


Volume I, 1916-1929" .. aw 0%. 2S, 1$5.00 
Volume II, 1930-1934 . .... . 5.00 
Volume IH), 1935-1936. = =. 4. 5.00 
Volume IV, 1937-1938... .. 4. | 5:00 
Volume V, 1939-1940 ..... . 5.00 
Volume VI, 1941-1942 . . ... . 5.00 
Volume VII, 1943-1944 . . . . . . 5.00 
Volume VIII, 1945-1946 . ... . 5.00 
Volume IX, 1947-1948 . . . . . . 6.00 
Volume X, 1949-1950 . .... . 7.00 
Volume XI, 1951-1952. . . . . . 8.00 
Volume XII, 1953-1954. . . . . . 8.00 
Volume XTIT, 1955-1956 . . .. . 8.00 
Volume XIV, 1957-1958 .... . 8.00 
Single numbers, 

Volumes: feand*2). 2. «i037 oo .50 
Succeeding numbers, 

(except Volume XI, No.2) . . . . 1.00 

Volume XI, No.2 2.3 452 bR oa eee 


The subscription price of MApDRONo is $6.00 per year. If your 
institution does not now subscribe to MADRONO, we would be 
grateful if you would make the necessary request. Since there 
is no extra charge for institutional subscriptions, an individual 
may hold membership in the California Botanical Society on the 
basis of his institution’s subscription. 


Address all orders to: 


FRANCIA CHISAKI, Corresponding Secretary 
Department of Botany 


University of California, Berkeley 4, California 


SMITHSONIAN INSTITUTION LIBRARIES 
IAEA 
3 9088 01111 8080