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Full text of "Proceedings of the California Academy of Sciences, 4th series"

PROCEEDINGS 



OF THE 



California Academy of Sciences 



FOURTH SERIES 



Vol. XXV 

ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 



••i^ 



Marine Biological Labo^tjlo/y 
FEB 2 31949 

WOODS HOLE, MASS. 



SAN FRANCISCO 

PUBLISHED BY THE ACADEMY 

1943-1949 



■•^\ 




— ^ ^ 



CttcCt^ ^xJ^^'-^^'T^J^ 



TO 

ALICE EASTWOOD 

IN COMMEMORATION OF FIFTY YEARS OF SERVICE AS 
CURATOR OF THE DEPARTMENT OF BOTANY OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

1892-1942 



THIS VOLUME IS DEDICATED IN RESPECT AND AFFECTION 



FOREWORD 

Imsx'OLUMEof the Proceedingsoi the Caliloi Ilia Academy 
of Sc iences honoring Miss Alice Eastwood ^vas initiated in 
i()42. on the occasion of her comj)lction ol lilty years ol 
service as Curator ol tlie Academy's Department of Botany. 
O^sing to u'artime problems of publication, the first num- 
ber of the projected volinne was not isstied until December. 
ic)43. The volume has been continued through the suc- 
ceeding years and is now ccjncluded ^vitll this number 
which. ha}:)pily. we are able to isstie on the occasion ol 
Miss Eastwood's ninetieth birthday, and in recognition ol 
Iier liity-seven years of service to the Academy. 

On her ninetieth birthday Miss Eastwood \vi\\. at her 

own rccjuest, retire from her post as Cin-ator of Botany. 

Ihc Academy's Cotuicil is protid to grant her the title of 

Curator Emeritus— a title which, however, seems singularly 

inappropriate to a gallant lady ^v ho ^vill be forever young. 

The biograpliy of Miss Eastwood that is here presented 
was prepared by Dr. F. M. MacFarland. President of the 
Academy from 19^^^ to i94(), in collaboration with Dr. R. 
C. Miller. Director of the Academy, and Mr. John lliomas 
Howell, Assistant Curator of Botany. The bibliography of 
Miss Eastwood's writings was prepared by Dr. MacFarland 
and Miss Veronica }. Sexton, the Academy's Executive 
Eibrai ian. The thankless task of indexing the \-olume ^vas 
performed by Dr. MacFarland alone. 

For all those A\'ho participated in its preparation and 
publication, this volume represents a labor of l(^\e. 

M. E. LOMRARDI. 

Presidciu ol the Acadcniv 



CONTENTS OF VOLUME XXV 

PAGES 

Biograpliical Skctcli of Alice Eastwood IX- XIV 

BibliogTaphy of the Writings of Alice Eastwood XV-XXIV 

Contributed Papers : 

No. 1. MuNZ, Philip A. A Revision of the Genus Fuchsia (Ona- 

graceae) (Plates 1-16) Published December 11, 1948 .... 1-138 

No. 2. Campbell, Douglas Houghton. Relations of the Temperate 
Floras of North and South America. Published June 1, 
1944 139-146 

No. 3. Barneby, R. C. Pugillus Astragalorum Alter (Plate 17). 

Published June 1, 1944 147-170 

No. 4. Smith, Gilbert M. Sublittoral Marine Algae of the ^lon- 

terey Peninsula. Published June 1, 1944 171-176 

No. 5. Wolf, Carl B. The Gander Oak, a New Hybrid Oak from 
San Diego County, California. (Plates 18-19). Published 
June 1, 1944 171-188 

No. 6. Wiggins, Ira L. The Genus Drymaria in, and adjacent to the 

Sonoran Desert. (Plates 20-22) . Published June 1, 1944 . . 189-214 

No. 7. Pierce, George J. Water and Plant Anatomy. Published 

June IJ 944 215-220 

No. 8. ]\Iason, Herbert L. A Pleistocene Flora from the McKittrick 
Asphalt Deposits of California. (Plates 23-24) . Published 
June 1, 1944 221-234 

No. 9. Ew^an, Joseph. The Perennial Southwestern Datura and the 
Validity of Matthew's Hypothesis in Plant Geography 
(6 text figures) . Published June 1, 1944 235-244 

No. 10. Benson, Lyman. A Revision of some Arizona Cactaceae 

(Plate 25). Published November 10, 1944 245-268 

No. 11. Babcock, Ernest B. Endemism in Crepis (4 text figures) 

Published November 10, 1944 269-290 

No. 12. Goodspeed, T. H. Nicotiana Arentsii — A New, Naturally 
Occurring Amphidiploid, Species. (Plates 26-27). Pub- 
lished November 10, 1944 291-306 

No. 13. Stebbins, G. L., Jr., Tobgy, H. A., and Harlan, Jack R. The 
Cytogenetics of Hybrids in Bromus II. Bromus Carinatus 
and Bromus Arizonicus. (13 text figures). Published No- 
vember 10, 1944 307-322 






PAGES 

No. 14. McMiNN, Howard E. The Importance of Field Hybrids in 
Determining Species in the Genus Ceanothus. (Plates 
28-36, 3 charts, 3 histograms). Published November 16, 
1944 323-356 

No. 15. Howell, John Thomas. A Revision of Phacelia Section Mil- 

titzin. Published December 27, 1944 357-376 

No. 16. St. John, Harold. Endemisni in the Hawaiian Flora, and 
a Revision of the Hawaiian Species of Gunnera (Hal- 
oragidaceae) Hawaiian Plant Studies 11. (Plates 37-46). 
Published November 15, 1946 377-420 

No. 17. Keck, David D. A Revision of the Artemisia vulgaris Com- 
plex ill North America. (19 text figures) Published No- 
vember 15, 1946 421-468 

No. 18. Setchell. William A. The Genus Ruppia L. (Plates 47-48) . 

Published November 15, 1946 469-478 

Index to Volume XXV 479-485 



BIOGRAPHICAL SKETCH OF 
ALICE EASTWOOD 

Ox .January' 19, 1851), in two localities, some two thousaiul miles apart, there 
were in progress two separate and apparently unrelated series of events that 
were destined to become, at a later date, of considerable mutual importance. 
In San Francisco, the California Academy' of Sciences, an organization then 
less than six years old, was struggling successfully through the precarious 
early years of its existence. In Toronto, Canada, Alice Eastwood Avas born. 

^lore than tliirty years were to elapse before the events referred to became 
signifieant for each otlier. Nevertheless, even before Alice Eastwood was born, 
the avenue that was to lead her to distinguished botanical career had been 
opened by the "scientific gentlemen" (as they described themselves in their 
fir.st constitution) who made up the California Academy of Sciences. As if 
with prophetic insight, Dr. Albert Kellogg, one of the founders of the Academy 
and its most prominent early botanist, had proposed in a quaintly worded 
resolution that was adopted at the meeting of August 1, 1853 : "Be it resolved, 
as the sense of this society, that we highly approve of the aid of females in 
every department of natural history, and that we earnestly invite their coop- 
eration." Thus, through Dr. Kellogg's resolution, as gallant in intent as it 
was inej)t in phrase, the California Academy of Sciences became very probably 
the first institution in the world to formally encourage the participation of 
women in scientific work, as it became later the first to admit women to senior 
curatorial positions. 

While Alice Eastwood was still a young girl, her family moved from Toronto 
to Denver, Colorado, where she obtained her high school education, graduating 
from the East Denver High School with its first class in 1879. Her marked 
ability and eager thirst for knowledge led her to be drafted as a substitute 
teacher before she had finished high school, and slie continued as a regular 
teacher for the next ten years. Despite the burden of teaching a great variety 
of more or less unrelated subjects, her innate love of Nature asserted itself 
and the plants, shrubs, and trees of Colorado appealed to her with irresistible 
charm. At every opportunity she explored the plains, valleys, and mountains, 
the rugged canyons, streams, and rivers of the state, and brought together a 
tangible record of her field studies in the Eastwood Herbarium, which later 
was to become the pride of the Denver Museum. Without access to a scientific 
library or contacts with other biologists, she became, herself, a self-taught 
botanist. Gray's IManual, covering the flora of the eastern states was her 
earliest guide to unraveling the family and in part the generic relationships 
of the Colorado plants, and she welcomed with delight the appearance of 
Coulter's ^Manual of Rocky Mountain Botany in 1885, and she soon found 
ample work to do in extending its application to the rich Colorado flora. 

In 1890, Miss Eastwood made a vacation trip to southern California, study- 
ing especially the plants of the San Diego region. Returning in the next year, 

[ IX 1 



X CAUFOBMA ACADEMY OF SCIENCES [Proc 4th Ser. 

she spent some niunths as an assistant in the li('i'l)ai-iiiiii (if the California 
Academyof Sciences. Asa result, she Avas induced to leave her Denver position 
in 1892. and to east her h»t with the Aca(h'iiiy. soon heeomiii^' joint ("ui-ator 
of Botany with Mrs. Katliei-iiie IJraiiih'izce. Thi'ci' ye;ii-s hitei'. upon t he retire- 
ment of ]Mrs. I'.randejree, she became Cui-ator and llrjid of the Dt'iiai-tinent of 
Botany wliich position she has fUhMl until the pi'escni. 

In the forty-seven years of its existence, the (jilifoinia Academy of Sci- 
ences had grown fnmi its simple beg'innings to he the leading center of research 
in the Natural Sciences of tiie Pacific Coast. Its heH)arium, built up by the 
pioneers. Allx-rt Kellogg. 11. X. P>olander. Harry M. Ilarkness, and E. L. 
(Jreen, and continued by T. S. and Katherine lirandegee and others, had 
become one of the largest in the West, and was in constant use by student 
workers. ]\Iiss Eastwood at once entered upon W( rk to inci-ease its value. Aside 
from her I'ontinc ciiralorial duties, she collected and studied plants and trees 
near and fai'. The Hay I'cgion was thoroughly explored and ti'ips were made 
to Mt. Shasta, to Del Norte County, to the Santa Lucia Mountains, and the 
interior valleys, returning with much new material for the herbariiun. 

Tn those pre-autoniobile days, travel away from the railroads was often 
difticult. liV stage, on hoi-seback, or afoot. Miss Eastwood made her Avay over 
mountain roads and trails with unflagging zeal. 

As a resnlt of her activity and that of those whom she inspired, the herl)ar- 
iiim steadily increased in size and scientific value. The great earth(|uake and 
fire of 1!H)(). however, destroyed much of San Francisco in a few hoin-s. and 
the nuisenni and collections of the Academy shared the same fate. Through 
Miss Eastwood's iiutiative and courage, though, some important things were 
saved — the I'eeords of the Academy from its beginning, and twelve liinidred 
and eleven invaluable botanical types, irreplaceable if lost. Miss Eastwood 
had long recognized their value and had segregated them from the thousands 
of other specimens, so that when the emergency came, she was able to find 
them without delay in the wrecked building and to transport them to a place 
of apparent safety — a I'efuge that nevertheless had to be changed twice before 
the approaching fiames. Tn doing this, she unhesitatingly abandoned her ow'n 
personal possessions, her favorite pocket lens alone being saved. In a letter 
to Science for May 25, lf)()(i, describing the (h'stiMidion of the Academy and 
collections, she wrote: "1 do not feel the loss to be nnne, but it is a great loss 
to the scientific woi'ld and an irreparable loss to California. My own destroyed 
work 1 do iu)t lament, foi' it was a joy to me while I did it, and 1 still can have 
the same joy in sfjuting it again. ... To me came the chance to care for what 
was saved from the ruins of the Academy, and with the hel]i of my devoted 
friends T was able to do it." 

The type s]")ecimens saved thi'ough ^liss Eastwood's courage formed the 
luicleus of the new collections. But owing to the total destruction of its prop- 
ei-ty and the interrujition of its income, it was several years before normal 
aetivitv could be resumed. During this lime, ]\Iiss Eastwood studied in the 




#0^ W9^ 



^%t 





ALICE EASTWOOD 

Ahiiiil iSSo, ;ii ilic Ijfi^imiiiif^ ol lic'i lioUiiiii ill (aicci 



Vol. XXV] ALICE EASTWOOD XIIT 

j^'i-o^at lierbaria of the Siuitli.soiiiaii Institution at Wasliinuton, I he Xt'w Voi-lx 
Botaiiical Garden, and Harvard I'niversity, and also spent a ninnber oi' 
months in the Kew Gardens, the British Museum, tlie University of Cam- 
bridge, and the Jardin des Plantes. 

With tlic new site for its bnihlini^s in (iokh'ii Gate Park assured and con- 
struction begun, tlie Academy's normal scientific activity was resinned. Miss 
Eastwood returned in June 1912, and at once vigorously began the replace- 
ment of the herbarium of the Department of Botany and tlie rebuilding of 
its library; collecting trips were made at every opportunity. Through the 
cooperation of Professor Charles S. Sargent of the Arnold Arboretum, she 
made a trip to Alaska in 1914, collecting and studying especially the willows 
of the Yukon region for his Sylva of North America . Later, studies were also 
made by her in southern California, Arizona, and New Mexico for Professor 
Sargent. She made other extensive trips to all parts of California, to Oregon, 
Nevada, Utah, Arizona, New Mexico, and Lower California, returning with 
abundant material for the rapidly growing herbarium. Additional collections 
made by others came from Mexico, Central America, South America, and the 
Pacific Islands. 

Especial attention has also been given to securing specimens of the exotic 
plants which have been introduced into California gardens and parks, a field 
often neglected in herbaria, but of great importance for com]iarative study. 

Miss Eastwood's publications as shown in the accompanying bibliography 
of over three hundred titles have been many and varied. The majority of them 
are, of course, technical botanical studies, but others are of more general 
interest inculcating an appreciation of the out-of-doors and the conservation 
of its ti-easures. In addition, the admirably edited "Leaflets of Western 
Botany" published by her and Mr. J. T. Howell, Assistant Curator of Botany, 
affords a prompt means of publication for shorter articles by western botanists. 

It is worthy of particular remark that, while Miss Eastwood had written 
more than one hundred papers by the time she was fifty years of age, and 
was considered sufficiently distinguished to be starred in the first edition of 
Americcni Men of Science (1906), two-thirds of her published writings have 
been issued since that time, and fifty-five of them since her eightieth birthday. 

Miss Eastwood has taken a prominent part in promoting the welfare of 
many scientific organizations interested in plant life. She has been the active 
head of the California Botanical Club (of San Francisco) since its second 
year in 1892. She has participated in the work of the California Floral Society, 
the San Francisco Garden Club, the California Spring Blossom and Wild 
Flower Association, the American Fuchsia Society, the California Horticul- 
tural Society, and is a member of the Science Club, the Alpine Club, the 
Tamalpais Conservation Club, and other organizations devoted to the out-of- 
doors and its conservation. She became a Member of the California Academy 
of Sciences in 1892, a Life Member in 1917, was elected a Fellow of the Acad- 
emy on February 25, 1928, was declared a Patron on December .{l, 1930, and 



XIV CAUFOSNIA ACADEMY or SCIENCES [Proc. 4th Ser. 

was iiiiaiiimously elected an Honorary ^Member of the Academy at its Annual 
Meeting on February 18, 1942. 

She has represented the Academy in many scientific meetinos at home and 
abroad. Among these latter are the Internatioiuil Congress held in Cambridge 
ill l!t.")(). in Holland in 1!>.'}5. and the Centennial Anniversary of the Jardin 
des riantes in the same j'ear. 

Xdtwitlistanding the ]ires.sure of her scientific activities. Miss Eastwood 
lias never been too busy to help others, to answer the countless questions that 
come to her daily, to extend willingly to others the richness of her knowledge. 
Her frank, direct approach to every problem and her modest, kindly spirit 
have won for her the esteem and love of countless loyal and devoted friends 
who wish for her many more years of productive activity in the field she loves. 



BIBLIOGRAPHY OF THE WRITINGS OF 
ALICE EASTWOOD 

181U. Thf luorning glory liusli, liKiiiund hpiophylla. West. Amer. Scientist, 7:119. (Feb.) 
LcKcocrinum montanum. West. Amer. Scientist, 7:141. (Mar.) 
The common shrubs of sonthwest Colorado. Zoe, 2 : 102-104. (July) 
The fertilization of geraniums. Zoe, 2:112. (July) 
The Mariposa lilies of Colorado. Zoe, 2:201-203. (Oct.) 
Additions to the flora of Colorado, I. Zoe, 2:226-333. (Oct.) 

1892. The loco weeds. Zoe, 3:53-.58. (April) 

Eeview on "Outlines of lessons in botany for the use of teachers, and mothers studying 

with their children" by Jane H. Newell. Zoe, 3 :78. (April) 
Notes on some species of the genus Oenothera. Zoe, 3 : 248-252. (Oct.) 

1893. Notes on the cliff dwellers. Zoe, 3 :375-376. (Jan.) 

General notes of a trip througli southeastern Utah. Zoe, 3 : 354-361. (Jan.) 

Notes on some Colorado plants. Zoe, 4:2-12. (April) 

Additions to the flora of Colorado, II. Zoe, 4:16-20. (April) 

A popular flora of Denver, Colorado. San Francisco, Zoe Publ. Co., May, 1893. 57 pp. 

List of plants collected in southeastern Utah, with notes and descriptions of new 

species. Zoe, 4:113-127. (July) 
Field notes at San Emidio. Zoe, 4:144-147. (July) 
Botanical notes. Zoe, 4:286-287. (Oct.) 
Gilia supcrha. Phacelia nudicaiijiti. Zoe, 4:296. (Oct.) 

1895. Additions and corrections, January 14, 1895. 3 pp. [To: A popular flora of Denver, 

Colorado. 1893] 
Two new species of Aquilegia from tlie Upper Sonoran Zone of Colorado and Utah. 

Proc. Calif. Acad. Sci., (2), 4:559-562, pis. 18-19. (Mar.) 
On heteromorphic organs of Sequoia sempervirens. Proc. Calif. Acad. Sci., (2), 5 :170- 

176, pis. 15-18. (May) 
Leucocrinum montanum. West Amer. Scientist, 9, 78:69. (June) 
Observations on the habits of Nemophila. Erythea, 3 : 151-153, pi. 3. (Nov.) 

1896. New localities for west American plants. Erythea, 4:32. (Feb.) 
Pelargonium anceps, Ait. Erythea, 4:34. (Feb.) 

New localities for two introduced plants. Erythea, 4:34-35. (Feb.) 

Ephedra viridis Coville. Erythea, 4:71. (April) 

Trillium sessile. Erythea, 4:71. (April) 

On Dr. D. Prain's Account of the genus Argemone. Erythea, 4 : 93-96. (.June) 

New stations for two introduced plants. Erythea, 4 :99. (June) 

Arbutus Memiesii in San Francisco County. Erythea, 4:99. (June) 

Report on a collection of plants from San Juan County, southeastern Utah. Proc. 

Calif. Acad. Sci., (2), 6:270-329, pis. 44-46. (Aug.) 
The alpine flora of Mt. Shasta. Erythea, 4:136-142. (Sept.) 
Geranium parviflorum Willd. Erythea, 4:145. (Sept.) 
Scolymus Hispanicus L. Erythea, 4:145. (Sept.) 
New stations for two plants. Erythea, 4:151. (Oct.) 
Abies bracteata. Erythea, 4:174-175. (Nov.) 
Centaurea calcitrapa L. Erythea, 4:175. (Nov.) 

1897. A guide for the analysis and description of flowering plants. Prepared for the Botani- 

cal Club, California Academy of Sciences. San Francisco, George Spaulding & Co. 
8 pp. 

[ XV ] 



XVI CALIFOKXIA ACADEMY or SCIENCES [Pkoc. 4th Ser. 

[Ht-vit-w ofj Volinne X of the "Hilva of North Aiiiorii-ii," l)y Sargent. Krythea, 5:43- 

44. (March) 
[Review of] "L.-ilmratory ]iractic(' for lieyiiincrs in liotnny." Erythea, Si.lfl-Cil. 

(April I 
Tlie coiiifcrac of the Santa T.m-ia Mountains. Krytln-M, 5:71-74. (June) 
Caiiforniaii licrli loic — J,'li(t nniu.s CalifoniicKs. I']iytlica, 5 :98. (Sept.) 
Studies in the herliaiium and 1h<' titd.l. I. I'rnc. ( 'alif. Acad. Sci., (3), Botany, 1 :71-88, 

pi.s. ()-7. (Nov. ) 

1898. Ferns of the Yoseniite and the neifrlil.oriny Sierras. Krythea. (i: 14-15. (Feb.) 
[Review of J The second fascicle of ''IIow ell's Flora of northwest America." Erythea, 

6:58-60. (May) 
Plant inhabitants of Nob Hill, San Francisco. Erythea, 6:61-67. (June) 
Notes on tin- flora of Marin County, California. Erythea, 6:72-75. (July) 
Is X( riiiiliiillinu frnar n septennial.' Erythea, 6 : 75-76. (July) 
I'l/iiihi iiiiiKir \j. in California. I''iytliea, 6:93. (Sept.) 
Studies in the heriiariiini and tlic Held, I 1 . i'roc. Calif. .Acad. Sci., (3), Botany, 1 :89- 

14(;, jds. S-l I. (Sept.) 
I'lants in Hower in November and December, 1897. Erythea, 6:114-115. (Nov.) 

1899. New localities for rare California plants. I'^rythea, 7:76-77. (Aug.) 
I'ariiti.sfiia califoniica Greene. Erythea, 7:84. (Sept.) 

Ai( ntiiiii iKiliitliciild. Krythea, 7:149. (Nov.) 

Introduced plants in Placer County, California. Erythea, 7 : 150. (Nov.) 

Sedge used in Indian basket making. Erythea, 7:15U. (Nov.) 

.Mr. Ilendrie's gifts to the California Academy of Sciences. Erythea, 7 :15U. (Nov.) 

Migratory plants in A l.i nieda County. Erythea, 7:175-176. (Dec.) 

.\n(itlici- int i(H|iii-ed plant. Krythea, 7:176. (Dec.) 

190(1. ... Krv and tidia. Rocky Mountain cilitidn. I'i-cparcd Ijv .Mice Kastwdiid. I'.oston, 
(;iiiii \- Cinnpany, | (-19001 1311 p. tigs. 1- I I. At head of title: P.ergen's Foundations 
of Jjiitany. At lie;i<l of cover-title: Bergen's Klenients of botany. 

Notes on ('ii/in s.^ii.s M acnahkma. Zee, 5:11-13. (.Tune) 

.\iliilli (/Iti i.iiiiild. Zoe, 5:28-S0. (July) 

On the occurrence of IiJi(i(Ki<liolas liedj/pnois All. ( Ifi ilii/nuiis /kiI i/iinir/iliid DC.) in 
North AuH'rica. Zoe, 5:35-36. (July) 

Finns Lmiihi rluina on Mt. St. Helena. Zoe, 5:36-37. (July) 

Arcloshijiliiiliis as a host plant for Phoradi ndvoii. Zoe, 5 :37. (July) 

ScorzoiK ra hispanicd 1^. Zoe, 5:37. (July i 

Some plants of Mendocino County new to the tlora of California. Zoe, 5:58-()(i. (Aug.) 

Rediscovery of Tliermopsis macrophylla IT. & A. Zoe, 5 : 7()-78. (Sept./Oct.) 

New sjiecies of California jdants. Zoe, 5:80-90. (Sept./Oct.) 

llMi 1 . . . . Kc\' iiikI tloia. Pacific Coast editi(ni. I'i('|iarrd iiy .Mice Fastwood. 1 ioslon, ( i inn 
& ('oni|iaiiv, liiol. |cl897, 1901]. l!»!l p. 4 ligs. At head of title: Bergen's Botany. 

. . . K(\ and tidia. Pacific Coast edition. i're])ared liy .Mice Eastwood. Boston, dinn & 
( (iiiilKiiiN , |iis:i7, PHil |. 199 p. 4 figs. At head of title: Bergen's Botany. liound 
witli: " l''oundations of botany," by .Fosepli )'. PxTgcii. Pioston, (iinn lV ('oiii|iany, 
I cl901J. 412 p. front., illns., plates. 

Asrirpias Kofolo. Zoe, 5:98-99. (Feb./Apr.) 

A study of Ertj.sinium (jrandiflorum Nutt. and the species aggregated undi r it. Zoe, 
5:100-103. (Feb./Apr.) 

Alriph .1- srniibarcdtnm R. Br. in Marin County. Zoe, 5: 13()-137. (T-'eb./Apr.) 

Iliihrnarid nidril inia (Jreene. Zoe, 5:137. (Feb./Apr.) 

Plants used for poisoning fish. Zoe, 5:136. (Feb./Apr.) 



Vol. XXV] WBITINGS OF ALICE EASTWOOD XVII 

Some sniJiU-flowered species of Neniupliila from the Pacific Coast. Jiulleliii of the 
Torrey Botanical Club, 28:137-160, pis. 15-20. (Mar.) 

An undescribed species of Paron/icliid from California, r.ulletin of tlic 'I'm rey Botani- 
cal Club, 28 :2S8-289. (May ) 

Notes on Californian species of Delphinium. UuUetin of the Torrey liotanical Club, 
28:667-674. (Dec.) 

1902. From Bedding to the snow-clad peaks of Trinity County ; also. List of trees and shrubs 

seen en route. Sierra Club Bulletin, 4:39-58, pis. LV-LVI. (Jan.) 
Some new species of California plants. Bulletin of the Torrey Botanical Club. 29 : 7.j- 

82, pis. 6-7. (Feb.) 
A descriptive list of the plants collected by Dr. F. E. Blaisdell at Nome City, Alaska. 

Botanical Gazette, XXXIII : 126-149, figs. 1-3, Feb. 21, 1902; 199-213, figs. 4-5, 

Mar. 22, 1902 ; 284-299, figs. 6-10, Apr. 18, 1902. 
Some new species of Pacific Coast Ribe.s. Proc. Calif. Acad. Sci., (3), Botany, 2:241- 

254, pis. 23-24. (Apr.) 
New species from the Sierra Nevada Mountains of California. Proc. Calif. Acad. Sci., 

(3), Botany, 2:285-293. (June) 
A flora of the South Fork of Kings River from Millwood to the head waters of Bubbs 

Creek. Sierra Club. Publ., No. 27. pp. 1-96, illus. (June) 
New species of Nemophila from the Pacific Coast. Bulletin of the Torrey Botanical 

Club, 29:471-474, pi. 21. (July) 
New western plants. Bulletin of the Torrey Botanical Club, 29:523-525. (Aug.) 

1903. New species of Oreocarya. Bulletin of the Torrey Botanical Club, 30 :238-246. (April ) 
New species of western plants. Bulletin of the Torrey Botanical Club, 30:483-502. 

(Sept.) 
Notes on Garrya with descriptions of new species and key. Botanical Gazette, 36:456- 
463. (Dec.) 

1904. Description of a light, compact arrangement of bedding and personal effects for a 

camping trip. Sierra Club Bulletin, 5:77-78. (Jan.) 
Hans Herman Behr. Science, 19:636. (April) 

Some new species of western Polemoniaceae. Botanical Gazette, 37:437-447. (June) 
A new GUia. Botanical Gazette, 38:71-72. (July) 

1905. Report of the Committee appointed to prepare and present an account of the life and 

services of Doctor Hans Herman Behr. Read before the California Academy of 
Sciences, March 21, 1904. San Francisco, The California Academy of Sciences, 
1905. 7 pp. front, (port.) F. Gutzkow, George Chismore, Alice Eastwood, Committee. 

New species of western plants. Bulletin of the Torrey Botanical Club, 32:193-218. 
(April) 

A handbook of the trees of California. Occasional Paper, No. 9, Calif. Acad. Sci. 86 
pp., 57 pis. (July) 

1906. New species of California plants. Botanical Gazette, 41 : 283-293, 2 figs. (April) 

The California Academy of Sciences. Science, N.S., 23 : 824-826. May 25, 1906. [Special 
article quoting Miss Eastwood's letters describing the destruction from the earth- 
quake and fire of 1906] 

The earthquake and the California Academy of Sciences. Torrey, 6:120-123. (June) 

1907. PeltipJiyUum pelfatum. ^luhlenbergia, 3 :78. (Aug.) 
EaHlardella muirii Gray. Muhlenbergia, 3:78. (Aug.) 
Truutvetteria grandis Nutt. Muhlenbergia, 3:78. (Aug.) 

Cypripedium fasciculaium in Santa Cruz County, California. .Mulilcnlicrgia, 3:97. 

(Sept.) 
Notes on California i)lants. ^Muhlenbergia, 3:131. (Dec.) 



XVIII CALIFOBXIA ACADEMY OF SCIEXCES [Proc. 4tu Sku. 

1908. Extensions of range. Lewisia redivivn ruisli. J^c\vi.si;i Icaiia (Porter i l^oliiiison. 

Muhlenbergia, 3 :147. (.I.iii. 

1909. Sonu' iiiiilesc-rilKMl species of Mcxiciii I'liancroganis. i'roc. Aincr. Aciul. Aits, Sci., 

44:603-608. (May) (Coiitrili. (ir.i.v ll(rl.:ii iiim, N.S. No. 36) 
Synopsis of tlie INfexican and Central Aiui'rican s]iecies of Cfisfilleja. Proc. Anier. 
Acad. Arts, Sci.. 44 : .')f;:'.-.";91 . CMay i (Contril.. (iray Ilcrl.ariinn. N.S. Xo. 'M'> > 

11*11. liartweg's Explorations in California, — 1; 11; 111; 1\'. M iililiiilicrKia, 7:2U-li4; '25- 
32; 37-43; 49-50. (Mar./May) 
[Letter] Die Zerstorung der Academie dcr Wissenscliaftfii in San Fraiuisco. Kihiig- 
lidu's Oyinnasinni zn Soran. Beilagc znni .lalircsheridit. No. lUli:lli-12. Translation 
of a letter to Dr. Angust Brand, dated May 14, 190(), 2421 Ridge Koad, Berkeley, 
concerning tlie destrnetion of the Academy's Herbarium in the eartlKpiake anil fire. 

lit 14. The iiiany.anitas of Mdinit Tama I] mis. ( 'alifoi iiia Out -of I loiirs. p. 7. ( A))|-il) 
The C(<i/iiilli IIS itf Mnuut Tamalpais. Califoiiiia ( )ut-()f- Doors, j). 4. (May) 
Natnre reclotliing Mount Tamalpais. California Out-of-Doors. ]>. 19. (Oct.) 
The rosaceous shrubs of ^fount Tainal])ais. The Tamaljiais Magazine, 4:6. (Nov.) 

l!M."). K'epiirt of the 1 )epai-t nn'iit of Botany for the year 1!I14. I'roc. Calif. Acad. Sci., (4), 
."'):2!t. (Mar. i 
How plants gidw on .Mount 'J'amalpais. California Out i)f-I>oors. ]>. 110. (Dee.) 

l!tl(i. K'ejioif of the Department of Botany for the year 1915. Proe. Calif. .\ca(l. Sci.. (4), 
5:208-209. (May) 

l!tl7. Ifeport of the Department of Botany for the year 1916. Proc. Calif. .Vcad. Sci.. (4) 
6:251-25;i (June) 

1918. Midwinter flowers on Mount Tamalpais. California Out-of-Doors. (Jan.) 
The shrnl)s covering Mount Tamalpais. California Out-of-Doors. (April i 

K'eport of tlie Department of Botany for tlie year 1917. Proc. Calif. Acad. Sci.. (4), 

7:;'.47-;i49. (June) 
Late summer on Mount Tamal]iais. California Out-of Doors. (Aug.) 

1919. Kaily spring at the Grand Canon near El Tovar. Plant World, 22:95-99. (April) 
Kejiort of the Department of Botany for the year 191S. Proc. Calif. Acad. Sci., (4), 

S :.•{;•,()-.'];? 2. (June) 

192n. Peport of the Dt'iiartmeiit of Botany for the year 1919. I'roc Calif. .Vend. Sci., (4i, 
9:379-380. (Aug.) 
Oaks on Mount Ta)n;il]iais. California Out-of-Doors. ji. 182. (Oct.) 

1921. Eai'ly s|ii-iiig on .Mcuiiit Tamaljiais. ( 'a lifoiiii a Out <if 1 )(ioi-s. p. 1 Sli. (Jan.) 
Tainal|iais wild tloweis. Trails, I :.j()-.")4. 

A plea for the protection of wild flowers. ( 'aliforiiia ( )ut ot' Doois. ( .\pi-il i 
Peport of the Deiiartmeiit of P.otany for the year 1920. I'roc. Calif. .\cad. Sci., (4), 
10:137-1:59. (.luly ) 

1922. h'eport of the Department of Botany for the yeai- 1!>21. I'roc. Calif, .\cail. Sci., (4), 

n :()25-(i29. (Aug.) 

1923. Winter flowers on Mt. Tamalpais. California Out-of-Doors. (.T.an. ) 
Trees of Mount Tamalpais region. Trails, 2:66-69. 

The origin of some lociility and trail names of Mount T.imalp.iis. C.ilitonii.i Out-of- 
Doors. ( .\piil I 
The fall flora of .Mcuint Tamalpais. California Ont-of-Doors. (Aug.) 
Peport of the Department of Botany for the year 1922. Proc. Calif. .\c;id. Sci., (4), 
11:674-676. (Aug.) 



Vol. XXV] WBITIXGS OF ALICE EASTWOOD XIX 

19124. Aix-hiliald Meuzies Jourii.il of tlie Vancouver expedition; extracts covering the visit 
to California, with an introduction and notes by Alice Eastwood. California His- 
torical Society Quarterly, 2 : 265-340. (Jan.) 
Report of the Department of Rotany for the year 192.'^. Proc. Calif. Acad. Rci., (4), 
12:1258-1260. (Oct.) 

1925. I Review ofj "The Giant Sequoia," by Rodney S. Ellsworth. Sierra Chih I'.ullctiii, 

XII: 204-205. 
The madrona. California ()ut-of-Doors. (Jan.) 
Sequoia versus Eucdli/pfK.s. California Out-of -Doors. (April ) 
Report of the Department of Botany for the year 1924. Proc. Calif. Acad. Sci., (4), 

13:467-468. (May) 
The rose family on Mt. Tamalpais. California Out-of-Doors. (July) 

1926. The aftergrowth of a niouiitaiii fire. California Out-of-Doors. (Jan.) 

Report of the Department of Hotany for the year 1925. Proc. Calif. Acad. Sci., (4), 
14:536-537. (April) 

1927. The madrona. Marin Messenger. (Feb.) 

Report of the Department of Botany for the year 1926. Proc. Calif. Acad. Sci., (4), 

15:528-530. (Mar.) 
New species of Ceanothus. Proc. Calif. Acad. Sci., (4), 16:361-363. (Ai)ri] ) 
X^otable plants on Mt. Tamalpais. California Out-of-Doors. (Sept.) 
ISIarin County in Noveml)er. Trails, 4:3. (Nov.) 
Toadstools and mushrooms. Trails, 4:2. (Dec.) 

1928. Trees of Mount Tamalpais region. California Out-of-Doors. (Feb.) 
Midwinter flowers on Mount Tamalpais. Trails, 5:2. (Feb.) 
Violets of Marin County. Trails, 5:2. (Mar.) 

Flowers noted on the trail to Little Carson on March 5th. Trails, 5 :2. (Mar.) 
More trees of Mt. Tamalpais. California Out-of-Doors. (Aug.) 

Report of the Department of Botany for the year 1927. Proc. Calif. Acad. Sci., (4), 
16:736-738. (May) 

1929. The Eastwood place on Mt. Tamalpais, Summit Avenue, Mill Valley. California Out- 

of-Doors. (Apr.) 
Report of the Department of Botany for the year 1928. Proc. Calif. Acad. Sci., (4), 

17:321-323. (May) 
After a tire in the Mount Tamalpais region. California Out-of-Doors. (Sept.) 
The escallonias iu Golden Gate Park, San Francisco, California, with descriptions of 

new species. Proc. Calif. Acad. Sci., (4), 18 :385-391. (Sept.) 
Studies in the flora of Lower California and adjacent Islands. Proc. Calif. Acad. Sci., 

(4), 18:39.3-484. (Sept.) 

1930. Report of the Department of Botany for the year 1929. Proc. Calif. Acad. Sci., (4), 

18:549-550. (April) 

1931. The true species of fuchsias cultivated in California. The National Horticultural 

Society, 10:100-104, illus. (April) 
Report of the Department of Botany for the year 1930. Proc. Calif. Acad. Sci., (4), 

19:436-438. (May) 
New species of plants from western North America. Proc. Calif. Acad. Sci., (4), 

20:135-160. (Dec.) 

1932. The pittosporums in California n gardens and parks. Leaflets of Western Botany, 

1:1-7. (Jan.) 
ZjigopliyJlum Fabdc/o L. Leaflets of Western Botany, 1:8. (Jan.) 
O.vnJis crrnun Thunb. Leaflets of Western Botany, 1:8. (Jan.) 



XX CALIF<il;.\I.\ .IC.II>J:M)()F SCIEXCES [Proc. 4th Ser. 

A new Califoniian Montia. Li-aflets of Western Botany. 1:11. (April ) 

Gentiana tenelUi in California. Lcaticts df West c in liotany, 1 :1(). (April » 

Another station for Zyyophijlhun Fabiujo L. Lealiets of Western Botany. 1 : 1 (i. ( April) 

Vfrbenn honarien.sis L. Leatlets of Western Botany, 1 :21. (Sept.~) 

Cyprip( dium fasciriilat inn in California. Lealiets of Western Botany. I:L*i'. (Sept.) 

Bottle-brushes in C.ilitdrni.m j^r.-irdciis anil parks. Leaflets of Western Botany, I :9-ll, 

April 4. 1932; 1:17-19, Sept. 21. l!t.!2: I :25-2(), Nov. 18, l!t:',2: 1:3:^-84, .Ian. IP, 

1933 : 1 :45-48, Mar. 2(i, 1933. 
The fetid adder's tonfjue. Leaflets of Western Botany, 1 :24. (Sept.) 
Concerning Pitfosporinii (hiiilinipliiilloides. Leaflets of Western Botany, I :24. (Sept.) 
Tlic tree ferns in ( ioliirii { l.-ilc Park, liay Cities ( la nlcn .M imt lily, 1 :2. 2 iigs. (Nov.) 

1933. The small ferns of Marin ( 'diinty. California Out-of-Doors. (Jan.) 

Taxus brevifoJid in Marin Cduiity, C.ilifoniia. Leaflets of Western Botany, I:38-39l 

(Jan. ) 
A new yei-iia santa. Leaflets of Western Botany, 1 :4ll-41. (Jan.) 
.\n nndescrilied lily from the Sierra Nevada of California. Leaflets of Western Botany, 

I : 41-43. (Jan.) 
J.I iliou as a poisonous jilant. Leaflets of Western Bot.any, 1:43. (Jan.) 
A new Miniiiliis. Leaflets of Western Botany, 1 :44. (Jan. ) 
Flowers alonjT the hij^hwa.vs. Bay Cities Garden Monthly, 2:2, 8. (Feb./Mar.) 
Two new species of w-esfern CaVnnn. Leaflets of Western Botany, 1 :5.5-56. (Mar.) 
A new Californian FritUlaria. Leaflets of Western Botany, 1 :55. (Mar.) 
Juniperus chincnsis variety Toruluaa, an undescribed variety of Chinese .juniper. Bay 

Cities Garden Monthly, 2:2, 7. 1 fig. (April) 
Bottle-brushes recentl.v introdvu-ed into California. Leaflets of Western Botany, 1:.57. 

(Aug.) 
New manzanitas from Santa Cruz Island, California. Leaflets of Western Botany, 

1:61-03. (Aug.) 
.M;inz;Miitas. C;i lifiirnia Out-of-Doors. (Sept.) 
Madrone and manzanita. Trails, 10:2. (Oct.) 

Escallonias in Golden (}ate Park. Leaflets of Western Botany, 1 :65-69. (Nov.) 
New species of California ArctostupJnjlos. Leaflets of Western Botany, 1:73-80. 

(Nov.) 

1934. Chrirantliiis iiraiiiliflitnis, a correction. I^eaflets of Western l>ot:iny, 1:87—89. (.Tan.) 
A kev to till' i-iiniiniin fiiinilies of flow erinji' plants in Ca lif ornin , ;inil A yuiile fur the 

analysis and description of flowering jilants. Prepared for and puldislied by the 

Ciilifornia Botanical Club, California Academy of Sciences. San Francisco, The 

.lanu's H. Barry Company, 1934. 16 pp. 
I'lsi-allonias in (iolilen (rate Park — 11. Leaflets of Western P>otany, 1 :81-82. (Jan.) 
Gentiana Copelandi Eastwood. Leaflets of Western Botany, 1 :96. (Jan.) 
A revision of tlie genera formerly ini-luded in ArciostaplniJos. Leaflets of Western 

Botany, 1:97-100. (Mayj 
A new species of Solanum. Leaflets of Western I'.utany, 1 :104. (May i 
A revision of Arcinslnpli jilns with key and descriptions. Leaflets of Western r.otany, 

1:10.^-127. (Aug.j 
Glailioliis .scgetiim Ker-Gawl. in Si ma County, California. Leaflets of Western 

Botany, 1:131. (Nov.) 
New species of Liliaceae. Leaflets of Western Botany, 1:132-133. (Nov.) 
A new species of Tnliumii from British Cohinil)i;i. L(>;iflets of Western P.otany, 1:139. 

(Nov.) 
New species of 2'' /"("I'l"'"'- Leaflets of Western lintany, 1:139-141. (Nov.) 



Vol. XXV] WRITINGS OF ALICE EASTWOOD XXI 

19'do. Lrptospffmum and allied gviiera in GuidL'n Gate Park. Leari(.4s of Western Botany, 
1:169-170. (Mar.) 
New species of Calif ornian Caslillcja. Leaflets of Western Botany, 1:174-176. (Mar.) 
A new loco-weed from Oregon. Leaflets of Western Botany, 1 :178-179. (May) 

1936. New lioreal species of Castillejn. Leaflets of Western Botany, 1 : 193-195. (Feb.) 
CasiiUcja multisecta Eastwood. Leaflets of Western Botany, 1:195. (Feb.) 
Adenoplea madagascariensis. Leaflets of Western Botany, 1 :197. (Feb.) 

A new Townsendia from Utah. Leaflets of Western Botany, 1 : 206-207. (Feb.) 

Tlie most minute Mimnhis. Leaflets of Western Botany, 1:207. (Feb.) 

The shrubby malvastruins of California with descriptions of new species and a key 

to the known species. Leaflets of Western Botany, 1 : 213-220. (April) 
Osmanthus aurantiacus (Makino) Nakai. Leaflets of Western Botany, 1:224. (April) 
A new Eibes from the Sierra Nevada. Leaflets of Western Botany, 1 : 230-231. (Aug.) 
Malvastnim fragrans Eastwood. Leaflets of Western Botany, 1 :232. (Aug.) 
Variations in Castilleja. Leaflets of Western Botany, 1:235-238. (Nov.) 

1937. New species of Avestern plants. Leaflets of Western Botany, II : 7-9. (Jan.) 
Quest for lilies. Leaflets of Western Botany, 11:27-29. (April) 

Xolina Parry i in Ventura County, California. Leaflets of Western Botany, 11:29. 

(April) 
New species of Dodecatheon. Leaflets of Western Botany, II :36-38. (Aug.) 
New localities in North America for Sphaerophysa Salsitla. (Pall.) PC. Leaflets of 

Western Botany, 11:38. (Aug.) 
Zjigadenus fontanus, a new species from Mt. Tamalpais. Leaflets of Western Botany, 

11:41-42. (Aug.) 
A brief history of the fuchsia. Journal of San Mateo Garden Club. (Sept. /Oct.) 
Notes on Schizococcus, with a key to the species. Leaflets of Western Botany, II : 49-50. 

(Nov.) 
New species of western plants. Leaflets of Western Botany, II :54-56. (Nov.) 

1938. The trees of Mt. Tamalpais. California Out-of -Doors. (Jan.) 

Two new wallflowers. Leaflets of Western Botany, II : 73-74. (Jan.) 

A key to the common families of flowering plants in California, and A guide for the 

analysis and description of flowering plants. Prepared for and published by the 

California Botanical Club, California Academy of Sciences. San Francisco, The 

James H. Barry Company, 1938. Second edition (revised). 17 pp. 
Excerpt from Illustrated dictionary of botanical terms, by John Lindley, 1848. With 

introduction by Alice Eastwood. Eeprinted 1938. [This booklet is photolithographed 

from Lindley's "Introduction to Botany," Fourth Edition, Book I, p. 319 and Book 

III, pp. 347-383, 346] 
The cestrums cultivated in California. Golden Gardens, 6:8. (Feb.) 
The perennial lupines of California, I. Leaflets of Western Botany, II : 81-86. (April) 
The tobacco collected by Archibald Menzies on the northwest coast of America. 

Leaflets of Western Botany, II : 92-94, pi. II, figs. 1-2. (April) 
Allium lacunosum Wats. var. micranthum Eastwood. [Footnote to "A Botanical visit 

to the Vancouver Pinnacles, by John Thomas Howell."] Leaflets of Western Botany, 

11:101. (April) 
Jamrs-ia americana var. caJifornica (Small) Jepson. Leaflets of Western Botany, 

11:102. (April) 
Two new Scrophulariaceae. Leaflets of Western Botany, II : 104. (April) 
New species of Liliaceae. Leaflets of Western Botany, II : 109-112. (July) 
The yellow-flowered perennial lupines of the Pacific States. Leaflets of Western 

Botany, 11:125-128. (Nov.) 
A new variety of Delphinium calif ornicum. Leaflets of Western Botany, 11:137-138. 

(Nov.) 



XXII CALIFORXIA ACADEMY (U-SCIKXCES | PKOf . 4tii Ser. 

1939. rereiiiiial lupiiius of the I'acilic Htatcs. Leatii-ts of Wcstcni iiotaiiv, 11 : 14(]-15U ; 

180-183. (Feb., Apr.) 
A new plilo.x from Oregon. Leaflets of Western Botany, II :175. (Feb.) 
New California plants. Leaflets of Western Botany, II :18()-188. (April i 
Little known exdtics in Califoi iii;i. (Joldi'ii (jardens, 7 -J). (May ) 
Some unilescribed nortlifrn ( 'alifcirnian valerians. Leaflets of Western r.dt.'iiiy. 11: 

196-199, figs. 1-4. (July) 
Liipiniis fJanauK on Mt. Dana :ind ,it ad.j.-ieent loc.-ilities. Leaflets of Western Botany, 

11:2(11-2(11'. (.Jnlyj 
Eugenia »ii/rcianthes. Leaflets of Western Botany, II :"Jl-i-21;l. (Nov.) 
Two new lupines. Leaflets of Western Botany, II : 215-216. (Nov.) 
Eri/simiim filifoliiiin Eastwood. Leaflets of Western Botany. 11:144. (Nov.) 
Early botanical explorers on the Paeifie Coast and the trees they found there. C.-ili- 

fornia Historical Society Quarterly, 18:335-340. (Dec.) 

1940. The stercnlias cultivated in California. Leaflets (.f Western Botany, 11:217-218. 

(Jan.) 
New s])ecies of Liipinu.s. Leaflets of Western Botany, 1 1 :226-228. (Jan.) 
Studies in Ca.stiUrJa. 1. Ca.stilleja in the Marble Mountains, Siskiyou County, Cali- 
fornia. 2. Miscellaneous new species. Leaflets of Western Botany, 11:241-245. 
(April) 
Tlie Lupiinis Bmrcri aggregate. Leaflets of Western Botany, II :249— 253. ( Ai>ril ) 
New western plants. Leaflets of Western Botany, II : 263-209. (July) 
New western plants— II. Leaflets of Western Botany, 1 1 :281-285. (Nov.) 
Report of the Dej)artment of Botany for the years 1931, 1932. Proc. Calif. Acad. Sci., 

(4), 20:5(17-508. (Dec.) 
Carl Purdy. Herbertia, 7:33-34. 

1941. Late (lowering exotics in the San Fr;tncisco I>ay Region. Golden Gardens, 9:8, 11. 

(Jan.) 

New western plants — III. Leaflets of Western Botany, 111:18-24. (Feb.) 

The Lupiiiii.s hitifoliii.s aggregate. Leaflets of Western Botany, ITT :41— 43. (April ) 

On the golden anniversary of the California Botanical Club. Tjeaflets of Western 
Botany, 111:2(5-27. (April) 

Tlie islands of southern California and a list of the recorded jilants — T. L(>aflets of 
Western Botany, TIT :27-36. (April j 

California, the land of flowers. 12th Annual Califiuiiia Spring (iardi'ii Show Year- 
book, Ajuil .Id to .May 4 (incl.) 1941. p. 29. 

The islands of southern California and a list of the i-ecordeil jil.-ints — II. I>eaflets of 
Western Botany, Til :54-78. (July) 

New species of Ca-stilleja. Leaflets of Western llotaiiy. I 1 1 :S7-92. (Nov.) 

1942. New species of CastiUiju. I^eaflets of Western Botany, 111:11 (i-1 1 7. (Jan.) 

A botanical ex|iediti(in to Log S]iring Ridge. Leaflets of Western I'.dtany, 111:121— 

126. (April; 
Two varieties of TriUhia p( duncnlan.s. Leaflets of Western Botany, 111:137-138. 

(April) 
New western |il;ints -IV. Leaflets of Western P.ot.any, HI : l.')7-l (id. (.Inly) 
A plea for preserv.ation of a magnificent ;izalea grove in Humboldt County. National 

League for Women's Service Magazine, 1():9. port., illus. (Aug. ) 
New species and varieties of Lupiiius. Leaflets of Western Botany, 111:169-175. 

(Nov.) 
SohiiiK)}! hiiiridliil lint in California. Leaflets of Western Butjiny, III : 1 89-190. (Nov.) 

194:'.. fiili<i sdllirold Kastwood. Leaflets of W^^stern T')Otany, 111: 199. (Feb. ) 
New western lupines. Leaflets of Western Botany, 111 :202-204. (Feb.; 



Vol. XXV] WRITIXGS OF ALICE EASTWOOD XXIII 

A new alien in California. Leaflets of Western Botany, III :208. (Feb.) 

I A review] Wild violets of North America. National League for Women's Service 

Magazine, 17:13. (Mar.) 
Some veronicas in Golden Gate Park. Leaflets of Western Botany, 111:220-222. 

(April) 
Some alpine castillejas of the high Sierra of California. Amer. Midland Naturalist, 

30:40-4(1, 1 pi. (July) 
Variations in Castillcja plagiofnma. Leaflets of Western Botany, 111:248-249. (July) 
Two new species from Baja California. Leaflets of Western Botany, 111:257. (Nov.^ 
A new Calif ornian rose. Leaflets of Western Botany, III : 262. (Nov.) 

1944. A collection of poimlar articles on the flora of Mount Tamalpais. | San Francisco], 

Published l)y the autlior, | 1944] 32 pp. Reprinted from "California Out-of -Doors," 

the official publication of the Tamalpais Conservation Club. 
A new Manzanita from Baja California. Leaflets of Western Botany, IV:4-5. (Feb.) 
The botanical colk'ctions of Chamisso and Eschscholtz in California. Leaflets of 

Western Botany, IV: 17-21. (April) 
A group of lupines related to Luphiu.s omisfitx. Leaflets of Western Botany, IV:41- 

43. (July) 
Why is a Fuchsia ? American Fuchsia Society. Bulletin, no. 44-7 : 3. (July ) 
A new lupine from Mt. Rainier. Leaflets of Western Botany, IV :43-44. (Jul}') 
New western plants — V. Leaflets of Western Botany, IV:63-()4. (Nov.) 

1945. Notes on Castanopsis, L'oiiniei/a and tlic sjiring star-flower. .lourn. Calif. Hort. Soc, 

6:195-196. (Jan.) 
New western plants — VI. Leaflets of Western Botany, IV: 148-150. (Mar.) 
The wild flower gardens of San Francisco in the 1890's. Leaflets of Western Botany, 

IV: 153-156. (May) 
New varieties of two well-known California plants. Leaflets of Western Botanv, 

IV: 166-167. (May) 
Tentative key and discussion of the lupines with spurred calyx. Leaflets of Western 

Botany, IV: 186-194, 1 pi. (Aug.) 
An account and list of the plants in the Brackenridge Journal. California Historical 

Society Quarterly, 24:337-342. 1 pi. (Dec.) 

1946. The Islay cherry. California Folklore Quarterly, V: 102-104. (Jan.) 

The lure of Arizona rock-ferns. American Fern Journal, 36 :49-51. ( Apr./June) 

A tentative key to the small-flowered lupines of the western United States. Leaflets 

of Western Botany, IV: 217-222, 1 pi. (Jan.) ; 251-254. (May) 
The prostrate oaks of Marin County, California. Leaflets of Western Botany, IV : 241- 

242. (May) 
A new lupine from Bartlett Mt. [In "On the botany of Bartlett Mountain," by John 

Thomas Howell.] Leaflets of Western Botany, IV: 259. (May) 
Notes on Arctostaphylos Uvn-ursi. Leaflets of Western Botany, IV:208-269. (Aug.) 
Species of the Lupiiuis lepidus group. Leaflets of Western Botany, IV: 281-284. 

(Nov.) 

1947. A collection of plants from the Aleutian Islands. Leaflets of Western Botany, V:9-13. 

(Jan.) 
Private herbariums donated to the California Academy of Sciences. Leaflets of 

Western Botany, V:4.5-48. (July) 
Endemism in the flora of California. Contributions from the Gray Herbarium of 

Harvard University, No. CLXV: 55-62. (Oct.) 
Monoecious junipers in Modoc County, California. Leaflets of Western Botany. 

V:72. (Oct.) 



XXIV CAI.IFOHMA ACADEMY OF SCIENCES [Proc. 4th See. 

1948. I Review] Origins of the flora of Califoniia. 15,v Uoughis 11. ('jiiindii-ll ami 1 la L. 

Wiggins. Stanford university iiublieations (University series) : Biological sciences, 

vol. X, no. 1. Stanfniil T'liivcrsity Press. 1947. Pacific Discovery, 1, 2:IS0-31. 

(March/April) 
[Review] The new world cyiircsses. By Cail i;. WHU' and Willis W. WageiuT. El Aliso, 

a series of papers on tiie native plants of California. Vol. 1. Rancho Santa Ana 

Botanic Oardm, A iialiciiii. r'alif. Ainil 1948. T'acific Discovery, 1, 4:3]-.'',2. (.July/ 

August) 
Studies of Pacific coast lilies — I. Ijcaflets of Western lJ(daiiy, ."> : ln;i-](i4. (May 
Biitaiiii-al (-(dk'ctions |Iii '"'riic i-nast of iKJrllicast Greenland with liydr(iyia|iliic 

studics in the Greenland Sea," by Louise A. Boyd.] American (iroiiraphical Society 

Special Publication, No. 30:270-273. (July) 
Studies of Pacific coast lilie.s — II. Leaflets of Western Botany, ~> : 120-123. (Aug.) 
Studies of Pacific coast lilies— III. Leaflets of Western Botany, 5 : 133-138. (Nov.) 
Monocciiiiis junipci's in Tklodnc T'cjunty. ("a lifdi-nia. Leaflets of Western P>iitaiiy, .1:72. 

(Oct. 1 

1919. Luiiinus Loijliii Gray, a good species. Leaflets of Western Botany, '> : loo-loi). (Jan.) 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
\'oL. XXV, No. 1, pp. 1-138, pis. i-iG December 11, 1943 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 1 

A REVISION OF THE GENUS FUCHSIA 
(ONAGRACEAE) 

BY 

PHILIP A. MUNZ 
Professor of Botany, Pomona College 

INTRODUCTION 

FOR A LONG TIME a garden favorite, the genus Fuchsia seems in recent years 
to have stimulated new interest and enthusiasm, and many new importa- 
tions of botanical species have been propagated and are being used for hybridi- 
zation and culture. The great variation in habit, in size, in color, in climatic 
conditions to which native, — all add to the possibility for use both as an indoor 
plant, and for milder climates, as a garden plant which can be adapted to 
hanging baskets, ground cover, shrub, vine or tree. With flowers ranging in 
size from a few millimeters to many centimeters, through various shades of 
green, red to purple, it is not surprising that it should command attention. 
Strangely enough, so important a genus has had no botanical revision since 
many of the species have become known. For horticultural as well as for botan- 
ical purposes, it seems desirable to know what the species are and where they 
come from. To this end, the present paper is presented. 

Fuchsia was first made known to science by a monk named Father Charles 
Plumier who described a species which he called "Fuchsia triphylla flare coc- 
cineo" in his ''Nova Plantarum americanarum Genera," p. 14, tab. 14, 1703. 
Linnaeus in 1753 based his Fuchsia triphylla on Plumier's description and 
figure. For a long time there was confusion as to the application of this name, 
it being used commonly for species from Colombia, such as F. venusta, but 
now it seems pretty well demonstrated that Plumier's plant came from Santo 
Pomingo. Fuchsia was apparently first introduced into England about the end 

[1] 



2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

of the 18th Century by a sailor who grew it in a mndow where it was observed 
by a niirserj'man of Hammersmith, a ^Ir. Lee, who succeeded in buying it and 
propagating it for the trade. This seems to have been one of the short-tubed 
species such as F. magellanica or coccinea. Later introductions of long-tubed 
ones were crossed with the short ones and our common garden hybrids have 
been developed. 

The name Fuchsia was given by Plumier in honor of Leonhard Fuchs (the 
spelling of whose surname ought to help those who have difficulty with the 
order of the letters in the generic name), a German physician and professor 
of medicine at the University of Tiibingen from 1535 to 1566. He was the 
author of a number of botanical works, particularly the great herbal "De 
historia stirpium" published in 1542, a Latin work dealing with about 400 
native German and 100 foreign plants. Agnes Arber in her "Herbals," page 
60, 1912, writes "Of all the botanists of the Renaissance, Fuchs is perhaps the 
one who deserves most to be held in honor." She rates him as superior in 
scholarship, critical study of the nomenclature of classical authors, and in 
botanical illustration. She feels that in his work is reached "the high-water 
mark of that type of botanical drawing which seeks to express the individual 
character and habit of each species, treating the plant broadly as a whole, and 
not laying more stress upon the reproductive than the vegetative organs." It is 
no mean name that is perpetuated in Fuchsia. 

On account of the great historical interest of Plumier's volume and its 
rarity, photographs of the title page, pages 14 and 15, containing the original 
description of the new genus Fuchsia, and plate 14 with its first published 
illustration are reproduced on Plate 16 of the present account. 

In its fleshy fruit the genus Fuchsia differs from all other Onagraceae; in 
this respect and in possessing a corolloid calyx and colored hypanthium it 
constitutes a distinct tribe (Raimann, in Engler and Prantl', Die nat. Pflanzen- 
fam. Ill, 7 :219, 1898) . I can see no very close relationship to any of the other 
genera in the family. Fuchsia is an immensely interesting group in the beauty 
and variety of its species, in the development of taxonomic differentiation, and 
in geographical distribution. In this paper I am recognizing about 100 species 
without attempting to go into horticultural and hybrid forms. I frankly admit 
that in many cases inadequate material or inaccessibility of types because of 
the present world conditions will undoubtedly make necessary a change of 
concept in the future. But the tremendous advantage that has been given to 
the North American student of South American plants by the great accumula- 
tion of photographs and fragments of types at the Field Museum through the 
long years of effort by J. Francis Macbride, now makes it possible to come to 
conclusions concerning species described long ago, which could not have been 
reached earlier without work in many European herbaria. 

In the preparation of this paper I have had from time to time access to 
various herbaria in Europe and South America, as well as in the United States, 
and have more recently borrowed some or all of the specimens from a numbed- 



Vol. XXV] 



MUNZ: THE GENUS FUCHSIA 



of the last named. To the curators of these institutions, to the John Simon 
Guggenheim Memorial Foundation which made possible the journey to South 
America, to Dr. LAnnan B. Smitli of (Jray Herbarium for notes, and to my wife, 
Alice ]\IcCulIy Munz, wlio did extensive library work on Fuchsia, I liereby 
express my appreciation. I submit the folloAving list of herbaria with the 
abbreviations which are used in the citation of specimens : 

Botanisches Museum, Berlin-Daldem (B) 

Bishop Museum, Honolulu (BISH) 

Bailey Ilortorium, Ithaca, N. Y. (BH) 

California Academy of Sciences, San Francisco (CAS) 

Field Museum, Chicago (F) 

Lillo Herbarium, Tucuman, Argentina (LID 

Gray Herbarium, Harvard University (Gil) 

University of Michigan ( MICH ) 

iMissouri Botanical Garden (MO) 

New York Botanical Garden (NY) 

Jardin des Plantes, Paris (P) 

Pomona College, Claremont, California (POM) 

Museu Nacional, Rio de Janeiro (R) 

Museo Nacional, Santiago, Chile (SGO) 

El Darwinion, San Isidro, Argentina (SI) 

Sao Paulo, Brazil (SP) 

Ignited States National Herbarium (US) 

Dominion Museum, Wellington, New Zealand (WELT). 

Taxonomic Characters in the Genus FUCHSIA 

For the most part the characters which I am using are the traditional ones : 
presence or absence of petals; proportions of the hypanthium; sepals erect, 
divergent, spreading or reflexed ; leaves opposite, alternate or whorled ; pu- 
bescence ; flowers erect or not ; stamens included or exserted ; shape of berry; 
pubescence on style; shape of leaves ; length of petioles, pedicels, etc. ; position 
of flowers, whether solitary or axillary or in inflorescences ; number of veins in 
leaves ; color of various parts ; stipules. Since the genus has never been keyed 
out nor anything like a complete treatment made of it, I have had to use pretty 
much my own judgment in choice of characters and have attempted to select 
such as could be seen with the least difficulty. All measurements and propor- 
tions have of course been taken from pressed specimens and this fact must be 
kept in mind if one tries to identify fresh material with my keys. 

Relationships Within the Genus 

I should think the species included in the section Qucliisio might he taken 
as the more primitive ones in the genus. The short hypanthium, erect stamens, 
many-seeded fruit, mostly spreading sepals seem characteristic onagraeeous 
features. Of the several species, F. magcllanica with its sepals quite free seems 




4 CALIFOBXIA ACADEMY OF SCIEXCES [Proc. -Ith Ser. 

primitive. The range of Quelusia, with F. magellanica in southern Argentina 
and Chile and with the other species occurring in the mountains of south- 
eastern Brazil, is an interesting one, and entirely different from all the rest of 
the genus. The Brazilian species have the sepals more or less connate at the 
base. 

The section Eiifuchsia is the most inclusive one in the genus. To me its most 
primitive species is F. splendens of Mexico, which seems near to Quelusia in 
its long exserted stamens, but has too long a hypanthium which is also peculiar 
in being very wide at the base. This species resembles more closely the section 
EufuoJisia than Quelusia in its large leaves. From it there can be arranged two 
series of species : one with long hypanthium, the second with shorter one. Both 
series run from solitary axillary flowers to well defined inflorescence. In the 
first series I would place nearest to F. splendens the Guatemalan F. cordifolia, 
in which the hypanthium has a broad base and the leaves are wide, but 'the 
hypanthium is longer and the stamens less exserted. From this through F. 
austromontana of Peru there can be derived a group with subcylindric hj'pan- 
thium rather wide at the base, but with narrow petals : F. denticulata, F. lepto- 
poda, F. Woytkowskii, and F. macjelalenae. Another group with wider petals 
and somewhat less broad hypanthium-base, but continuing the subcylindric 
condition contains: F. canescens, F. rivularis, F. loxensis, F. pallescens, F. 
Townsendii, F. platypetala, and F. macrostigma. Then comes a series with 
more ampliate hypanthium and wide petals : F. ayavacensis, F. Pringsheimii, 
and F. tripliylla; these have axillary flowers. With narrower petals and ampli- 
ate hypanthium and with flowers axillary or subterminal we have : F. petio- 
laris, F. Smithii, F. Lleivelynii, F. Asplundii, F. venusta, F. Jahnii, F. Gehri- 
geri, F. simplicicaulis, and F. confertifolia. "With the hj-panthium narrow and 
long, and with the flowers in a short, terminal, definite inflorescence and with 
leaves large, are : F. Asinazui, F. tincta, F. Mathewsii, F. Fischeri, F. Storkii, 
F. furfuracca, F. Munzii, F. hirtella, F. polyaniha, F. corymhiflora, F. 
ahrupta, F. Cuatrecasasii, F. Killipii, F. hoUviana, and possibly F. fulgens. 

The other great series which I would derive from the cordifolia type has a 
short hypanthium. Here again one can begin with axillary flowers with rather 
wide hypanthium-base, such as in F. decussata and F. Jiypoleuca, and then go 
to species with a narrower base, such as F. scahriuscula and F. verrucosa and 
F. sanctae-rosae, this last named having either axillary flowers or almost ter- 
minal inflorescence. From it one passes very easily into a large group with 
definitely terminal inflorescence : F. Osgoodii, F. Andrei, F. Lehmannii, F. 
putumayensis, F. Hartwegii, F. ovalis, F. asperifolia, F. pilosa, F. glaherrima, 
F. macrophylla, F. sessilifolia, and F. sylvatica. These species of Eu fuchsia 
are almost entirely Andean, excepting F. splendens, F. cordifolia, and F. 
fulgens of Mexico and Central America, and F. triphylla and F. Pringsheimii 
of the West Indies. 

The section Kierschlege.ria of Chile with its single species F. lycioides is an 
interesting xerophyte which must be related to 7^. magellanica in its short 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 5 

hypanthium and small flowers. I should suppose too that the section Skinnera 
of New Zealand and Tahiti with small flowers and reflexed sepals is of similar 
relationship, that is to the Patagonian magellanica. In its reduced or even 
obsolete petals it becomes a very distinct group that must have been long 
isolated. 

The same tendency to reduction or suppression of petals is seen in a long- 
tubed Andean series undoubtedly related to Eufuchsia rather than to Quelusia 
and for which I propose the section Hemsleyella. The species I have included 
here are doubtless not all of the same origin. In fact, I doubt that F. cestroides 
should be included with the others. They, in turn, are partly of the type with 
cylindrical hypanthium, for example, F. macrantha and F. Garlepyiana, or 
with ampliate hypanthium as in F. apetala. In addition to the suppression of 
petals, specialization is evidenced by such features as union of sepals at their 
base, loss of leaves at anthesis, etc. 

Perhaps the most highly specialized section is Encliandra which occurs 
from Panama to central Mexico. Small-flowered, with small leaves, and the 
shorter series of stamens bent back into the hypanthium, this group also tends 
in A'arying degrees to polygamy. It has also relatively few seeds in the berry. 
It may be perhaps related to the section SoJiufia (with erect flowers in large 
compound inflorescence) of the same geographical area in its small flowers 
and relatively short hypanthium. Just how these two groups are related to 
the rest of the genus is not clear ; perhaps the short hypanthium ties them in 
with the apparently rather primitive Quelusia which must have come from 
common stock with them. 

THE GENUS Fuchsia Linnaeus 

Fuchsia L., Gen. PI., ed. 1, 27, 1737; ed. 5, 126, 1754,- Sp. PL, ed. 1, 1191, 1753. Endlicher, 
Genera PL, 1193, 1840. Benth. and Hook., Gen. PL, 1, part 3: 790, 1862. Raimann in 
Engler and Prantl, Die nat. Pflanzenfam., Ill, 7:219, 1898. 

Hypanthium prolonged beyond the ovary, usually colored, deciduous in 
fruit. Sepals 4, usually colored, deciduous. Petals 4, or minute, or 0, convolute 
or spreading, deciduous. Stamens 8, usually unequal, the episepalous exceed- 
ing the epipetalous ; filaments more or less filiform ; anthers linear to oblong. 
Pollen grains single, 2- or 3-angled. Ovary 4-loculed ; style elongate ; stigma 
capitate or clavate, subentire to 4-lobed. Suffrutescent to arboreous. Leaves 
opposite to alternate or whorled, with small usually deciduous stipules. Flowers 
solitary in the axils or racemose or paniculate, mostly pendulous and long- 
pedicelled, generally showy ; perfect to polygamous. 

Type species, F. triphylla L. 

Key to Sections of FUCHSIA 

A. Petals lacking or reduced to small scalelike structures; leaves tending to be alternate. 

B. Hypanthium infundibuliform, not more than 1.5 cm. long ; sepals reflexed, separate. 

New Zealand and Tahiti 4. Slciimera. 



6 CALIFOEXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

BB. Hypanthium tubular to funnelforni, mostly more than 2 cm. long; sepals divergent 

to suberect, more or less connate at base. South America 5. Hemsleyella. 

AA. Petals present, almost or quite as long as sepals; leaves mostly opposite or whorled. 

B. Leaves with a thickened persistent petiole-like base which becomes spine-like ; 

sepals reflexed ; flowers small. Chile 3. Eierschlegeria. 

BB. Leaf-bases not persisting as spine-like structures. 

C. Flowers erect in terminal cymose panicles arranged subtrichotomously. Mexico 

to Panama 6. Schufia. 

CC. Flowers mostly pendulous, not in cymes. 

D. Flowers tending toward polygamy, small, inconspicuous ; stamens short, 
the epipetalous reflexed; berry few-seeded. Mexico to Panama. 

7. Encliandra. 
DD. Flowers perfect, usually showy; stamens more or less exserted, erect; 
berry many-seeded. Mostly South American species. 
E. Hypanthium usually not longer than the sepals ; stamens long- 

exserted. Eastern Brazil to Patagonia 1. Quelusia. 

EE. Hypanthium several times as long as sepals ; stamens not much ex- 
ceeding sepals. Andes north of central Argentina to Mexico and West 
Indies 2. Eufuchsia. 

Section 1. QUELUSIA (Vand.) DC. 

Quelusia as genus, Vandeli, F1. Lusit., 23, pi. 10, 1788; and in Eoem., Script. Hisp. Lusit., 

101,1796. 
Nahusia Schneev., Ic, 27, 1793. 
Thilcum Molina, Sagg. Chile, ed. 2, 146, 1810. 
Quelusia as sect., DeCandolle, Prodr., 3 :36, 1826. 

Nectary fused to base of hypanthium ; hypanthium eylindric, usually no 
longer than sepals ; stamens long, exserted. 

Type species : Vandeli described the genus without naming a species. De 

Candolle has F. coccinea under Quelusia, as well as other species ; he included 

F. magellanica as a synonym of F. coccinea. Since Vandeli referred to petioled 

leaves, he evidently had in mind F. magellanica Lam., and I would make it the 

type species. 

Key to Species of Section QUELUSIA. 

A. Hypanthium short, 3-6 mm. long ; younger branches pubescent to pilose. 

B. Leaf -blades 4-10 cm. long, 2-4 cm. wide, quite entire 6. F. regia var. aJpcstris. 

BB. Leaf -blades 1.5-15 cm. long, 0.3-2 (2.5) cm. wide, mostly serrulate. 

C. Young growth strigulosc ; leaves mostly less than mm. wide, coriaceous. 

Itatiaya, Brazil l.F. Campos-Portoi. 

CC. Young growth puberulent to pilose ; leaves mostly more than 6 mm. wide, 

membranous. 

D. Suffrutescent, with herbaceous stems 2-5 dm. tall; pedicels 1-2 cm. long; 

stems very densely shaggy-pilose. Espiritu Santo, Brazil. 2. F. Bracelinae. 

DD. Well developed shrub, 1-3 m. tall ; pedicels 2-4 cm. long ; steins puberulent 

to somewhat pilose. Minas Geraes, Brazil .A.F. coccinea. 

AA. Hypanthium longer, mostly 8-20 mm. long; younger branches usually glabrous or 
nearly so. 
B. Leaf -blades subcntire, 4-10 cm. long, 2-4 cm. Avide; sepals connate at base for 

2-15 mm. So. Brazil Q.F. regia. 

BB. Leaf -blades serrulate, 1-5 cm. long ; sepals mostly free. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 7 

C. Petioles mostly 3-10 mm. long ; hypantliium 8-10 (12) mm. long ; sepals 15-25 

mm. long, 3-4 mm. wide. So. Chile and So. Argentina 3. F. magellanica. 

CC. Petioles mostly 10-25 mm. long; hypanthium 10-20 (or more) mm. long; 
sepals 25-80 mm. long, 8-11 mm. wide. Gardens; naturalized as in Colombia, 
Venezuela 5.F. hybrida. 

(1) Fuchsia Campos-Portoi Pilger and Schulze 

(Plate 1, fig. 1) 
Fuchsia Campos-Portoi Pilger and Schulze, Notizblatt, 12 : 470, 1935. 

Shrub, with woody ascending or divaricate branches, the younger ones very 
slender, strigulose, more or less reddish ; leaves opposite or ternate or quater- 
nate, elliptic-lanceolate, acute at both ends, plainly serrulate with incurved, 
gland-tipped teeth, coriaceous, strongly bicolored, glabrous above or strigulose 
on midrib, strigulose beneath especially on midrib and margins, lateral veins 
inconspicuous, about 5 on each side of midrib; blades 1.5-2.5 (3) cm. long, 
3-5. (8) mm. wide; petioles 2-3 (4) mm. long, puberulent; stipules subulate, 
0.5 mm. long, deciduous; flowers solitary in upper axils; pedicels filiform, 
strigulose, 4r-15 mm. long ; ovary 4-5 mm. long ; hypanthium red, broadly 
fusiform, 4-5 mm. long, sparsely strigulose without, glabrous within ; sepals 
red, somewhat strigulose, oblong-lanceolate, acuminate, 18-20 mm. long, 4-6 
mm. wide, connate for 2-3.5 mm. at base ; petals violet, obovate-cuneate, 
slightly emarginate, 10-12 mm. long; episepalous stamens 20-25 mm. long, 
alternate ones slightly shorter ; anthers 1.5 mm. long; style glabrous, ca. 3 cm. 
long ; stigma subentire, ca. 2 mm. long ; fruit ellipsoid, ca. 1 cm. long. 

From Itatiaya, Serra da Mantiqueira, state of Rio de Janeiro, Brazil, where 
it grows at 2100-2300 m. alt. Material seen, Pilger and Brade 31, type coll. 
Dec. 27, 1934 (POM, R) , Brade 15676, March, 1934 (POM, R) , Loefgren 5775, 
March 13, 1903 (POM, SP) . I am greatly indebted to Dr. Brade of the Botani- 
cal Garden at Rio de Janeiro for information concerning and material of 
this species. The species is characterized by its narrow leaves, woody habit, 
strigulose younger growth, and fascicled appearance of the short branches. 

(2) Fuchsia Bracelinae Munz, new species 
(Platel, fig. 2) 

Suffrutescent, with running underground rootstock and erect, rather 
simple stems 2-5 dm. tall, woody and with exfoliating epidermis at base ; stems 
herbaceous, apparently reddish and densely pilose with light-colored hairs up 
to 1 mm. long ; leaves opposite, in 3's or 4's, ovate-lanceolate, green and some- 
what strigose above, paler, reddish and pilose beneath especially on veins, 
rounded at base, acute-acuminate at apex, subentire to denticulate, membran- 
aceous, distinctly veined (main laterals 6-8 on each side of midrib) ; blades 
2-5 cm. long, 5-14 mm. wide ; petioles 1-3 mm. long, villous ; stipules lance- 
deltoid, 0.5-1.5 mm. long, deciduous ; flowers axillary ; pedicels pilose, filiform, 
1-2 cm. long; ovary oblong, pilose, 5 mm. long; hypanthium dark red, pilose 







8 CALIFOBNIA ACADEMY OF SCIEXCES [Proc. 4th Ser. 

without, glabrous within, 3.5-5 mm. long, 2-2.5 mm. wide at slightly expanded 
summit ; sepals rose, 19-23 mm. long, sparsely pilose, connate at base for 3-4 
mm., then erect-divergent, lance-oblong or -elliptic, subacuminate, 5-6 mm. 
wide, 3-veined; petals purple, obovate, 11-15 mm. long, rounded apically; 
episepalous stamens up to 3 cm. long, epipetalous ca. 3 mm. shorter ; filaments 
reddish ; anthers less than 2 mm. long, reddish ; style reddish, up to 3.5 cm. 
long, glabrous; stigma slightly lobed, narrow, 1.5 mm. long; immature fruit 
oblong, 8 mm. long, 4 mm. wide. 

Suffrutescens; caulibus erectis, 2-5 dm. altis, herbaceis, pilosis; foliis oppositis, ternatis 
aut quaternatis, ovato-lanceolatis, in superficie superiore strigosis, inferiore pilosis, base 
rotundatis, apice acuto-acuminatis, subintegris vel denticulatis ; laminis 2-5 cm. loiigis, 
0.5-1.4 cm. latis ; petiolis villosis, 1-3 mm. longis ; pedicellis axillaribus, pilosis, filiformibus, 
1-2 cm. longis; ovario oblongo, piloso, 5 mm. longo; hypanthio 3.5-5 mm. longo, supra 
2-2.5 mm. lato, externe piloso, interne glabro; sepalis roseis, 19-23 mm. longis, sparse 
pilosis, lanceolato-oblongis aut -ellipticis, subacuminatis, 5-6 mm. latis; petalis purpureis, 
obovatis, 11-15 mm. longis; staminibus 2.5-3 cm. longis, rubris; stylo glabro, 3-3.5 cm. 
longo ; bacca oblonga, 8 mm. longa, 4 mm. lata. 

Type : from rocky, open campo, Serra de Caparao, State of Espiritu Santo, 
Brazil, Nov. 25, 1929, at 2650 m., Ynez Mexia 4013, Gray Herb., isotypes 
(CAS, NY, US) ; distributed as F. pulesccjis. Distinct from F. Campos-Portoi 
and F. coccinea in its more herbaceous habit ; from the former by its broader 
thinner leaves and more hair ; from the latter by its narrower leaves. It is a 
pleasure to dedicate this species to Mrs. H. P. Bracelin of Berkeley, California, 
who has devoted much time and energy to distributing the large collections 
made in Latin America by Mrs. Mexia. 

(3) Fuchsia magellanica Lam. 

Fuchsia magellanica Lam., Encycl., 2:5C5, 1788 ? (cf. Kuntze, Eev. Gen. PL, l:cxxxii, 
1891); pi. 282, fig. 2, 1793. 

Shrub up to 2.5 m. tall, bushy or semiscandent, freely branched, the ultimate 
branchlets very slender (ca. 1 mm. thick), usually reddish, glabrous to finely 
and sparsely puberulent, especially about the nodes, older ones exfoliating ; 
leaves opposite or ternate, sometimes alternate, usually rather crowded, the 
blades lance-ovate to rhombic-ovate to elliptic-ovate, rounded or more fre- 
quently somewhat cuneate at the base, acute to acuminate at the apex, usually 
evidently serrulate, more or less reddish on veins, paler beneath than above, 
glabrous to puberulent on margins and veins of under surface, 15-50 mm. long, 
■4-20 mm. wide, with 5-7 principal veins on each side of midrib; petioles 3-10 
(15) mm. long, glabrous to puberulent; stipules lance-deltoid, barely 1 mm. 
long, deciduous ; flowers 1 or 2 in upper axils ; pedicels filiform, 2-5.5 cm. long, 
glabrous to puberulent; ovary oblong-fusiform, 6-10 (12) mm. long, 1.5-2.5 
mm. wide; hypanthium tubular, 8-10 (12) mm. long, glabrous or nearly so 
Avithout, pubescent within; sepals narrowly lance-oblong, 15-25 mm. long, 
3-4 (5) mm. wide, acuminate, mostly glabrous; petals obovate, 1-2 cm. long; 
longer stamens 2-3.5 cm. long, epipetalous 4-6 mm. shorter ; filaments filiform, 



Vof. XXV] MISZ: Tilt: CKMS FICIISIA 

;rlabrous ; anthers 2-2.5 mm. long; style slemiiT, i)ubi*.scent at ba.se, mostly 3-5 
fin. loiifj; stijjrma capitate, more or less lobeil, 1.5-3 mm. long; berry oblong, 
1-2 cm. long, 4-5 mm. wide. 

This species is exceedingly varial)l(', with a noiiiciu lature much complicated 
by long culture in the garden and much hybridization. Many segregates have 
been prctposed and many attempts have been made to classify such (cf. IMiil- 
ippi. Bot. Zeit., 34:577, 187G; Bailey, Cyclop. Am. Ilort., 3:1301, 1015, for 
exami>les). In the study of wild plants the most that I can (ibserve is: (1) a 
tendency for larger flowers, larger thinner leaves, and longer pedicels in the 
more northern material, (2) opposite tendency in more southern material and 
(3) a possible less highly colored variety described by Prof. Espiuo/a. The 
complication involving hybridization with F. cocciiiea and other species 
cannot be discussed here. 

Key to Varieties of Fuchsia magcUanica 

A. Hvpnnthiiim and scpnls deep rod; ovary and pedicels red. 

B. Leaf blades mostly 15-25 mm. long; pedicels 20-.1.J mm. long 3a. var. typica. 

r>B. LcafMades mostly 25-50 mm. long; pedicels 40-55 mm. long. 31). var. marront' <:. 
\A. Ilvp.nntliium and sepals pink; ovary and pedicels green He. v.ir. V"/ . 

(3a) Fuch.sia magellanica Lam. var. typica Munz, new name 

(Plate 1, fig. 3) 
F. mageJlanica Lam., Encycl., 2:565, 1788; pi. 2S2, fig. 2, 1793. 
F. gracili.i var. muUiflora Lindl., Bot. Reg., 13:pl. 1052, 1827. 
F. gracilis var. icnella LiNDL., I.e.; F. macrostema var. toicUa DC. Prodr.. 3:38, 1828: F. 

lencUa Ilort., 0. Don in Loud., Hort. Brit., 155, 1830. 
F. conica Lindl., Bot. Reg., 13:pl. 1062, 1827 ; F. macrostemma var. conica SwE»rr, Brit. Fl. 

Garden, 6: under pi. 216, 1833; F. magellanica var. conico Bajley, Cyclop. .\m. Ilort., 

614. 1900. 
f F. multiflora LoDD., Bot. Cab., 15 :pl. 1415, 1828. 
F. discolor Lindl., Bot. Reg., 21 :pl. 1805, 1835 ; F. magcUanica var. discolor Bailey, Cyclop. 

Am. Ilort., 614, 1900. 
F.coccinea var. rohustior UooK., Fl. Ant., 2:269, 1^47; l>:isi<l •m ( '..iimiirscni .•oil. in Sir:iif 

of Magnlhacns. 
F. chonotica Phiu, Linnaea, 28:687, 1856; Bot. Zeit., 34:577, figs. 4 and 5, 1«76; F. eoe- 

cinea var. rhnnotica Reiche, Anal. Univ. Chile, 98:485, 1897; Fl. Chile, 2:267, 1898. 
F.araucana Piiiu, Bot. Zeit., 34:577, fig. 6, 1876. 
Dnrvalla cucharis CoMM. ex Lam., Encycl., 2:565, 1788. 

Leaf-blades mostly 15-25 mm. long, 4-10 (12) mm. wide, somewhat cori- 
aceous, often folded along the midrib; pedicels mostly 2-3.5 cm. long, red; 
hypanthium and sepals deep red; .sepals 1.5-2 cm. long; petals purplish; 
stamens 2-3 cm. long, reddish. 

Type locality, "Commerson a observe cet arbris.seau sur les montagnes du 
Magellan, dans presque tous les bois, en Janvier 17G.S." The range of this 
variety is in Chile mostly south of Coneepci«'>n, and in Argentina. Hepresenta- 
tive material, "F. mngrUauim Lam. diet. Dorvalli Commerson herb.," Lam- 
ank Herb. (P) ; F. jvulfiflorn. eiilt. Iiortul. (Jirod. CuMieva. in 1837 (.\Y) ; /•'. 



10 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th Seb. 

discolor, II. B. Glasgow in 1850 (NY) ; F. tenella, cult. Lochowitz in 1845 (F) . 
CHILE : Coneepeion : San Vicente, Pennell 12866 (GH). Araiico: Isl. of St. 
Mary's, Eujliis (US). Malleeo: Angol, Kuntze in 1892 (F). Yaldivia : Corral, 
Gunckel2 (F) ;Ceri'ode\a'Mavum,Gunckel5064 (POM) ; Valdivia, B«c/ihew 
in 1896 (Gil) ; Pangnipulli, Hollermaj'er under Werdermeyer 1364 (CAS, F, 
GH, NY, US). Llanquihue: Ancud, Chiloe, Pennell 12492 (F, GH) ; Isla 
Chiloe, near Castro, Mexia 8009 (F ,GH, NY, US) ; Isla Chiloe, Ruiz (POM) ; 
Peulla, Pennell 12675 (F, GH, NY). ARGENTINA : Neuquen : Puerto Meyer, 
Cabrera and Job 210 (NY). Rio Negro: Lago Nahuelhuapi, Ljimgner 226 
(NY). Patagonia: Port Otway, Lee in 1888 (US) ; Hassler Glacier, Hill in 
1872 (GH) ; Eden Harbor, Blake in 1872 (GH). 

Intergrades with var. macrostema are many, for example, with large leaves : 
Valparaiso, Wilkes Exped. (GH, NY, US) ; road from Curico, Chile to Argen- 
tina, beyond Los Queues, Mexia 7863 (F, GH, NY, US) ; Hualqui, Coneep- 
eion, Kuntze in 1892 (NY); Aysen, Llanquihue, Moreira in 1934 (POM) ; 
Isla Victoria, Lago N. Huapi, Argentina, Burkart 6397 (F) ; Lake Argentina, 
Santa Cruz, Furlong 98 (GH, NY, US). Other intergrades in length of pedi- 
cels, etc., are : Casa Panque, Llanquihue, Chile, Shannon 25 (US) ; Camino de 
Bariloche al Correntoso, Argentina, Cahrera 5044 (F, GH, NY). 

(3b) Fuchsia magellanica Lam. var. macrostema (E. and P.) Munz, new comb. 

F. macrostema R. and P., Fl. Peniv. 3 :88, pi. 324, fig. b, 1802. 

F. gracilis var. m-acrostema Lindl., Bot. Reg., 13 : under pi. 1052, 1827. 

F. coccinea var. macro.itema Hook., Fl. Ant., 2 :269, 1847. 

F. coccinea Curtis, Bot. Mag., 1 :pl. 97, 1789, not Ait. 

F. decussata R. Grah., Edinb. Phil. Journ., 11:206 and 401, 1824; Sims in Bot. Mag., 51: 

pi. 2507, 1824, not R. and P., 1802. 
F. gracilis Lindl., Bot. Reg., 10: pi. 847, 1824, new name for Graham F. macrostemma var. 

gracilis Sweet, Brit. Fl. Gard., 6: under pi. 216, 1833 plant (decussata)  F. magellanica 

var. gracilis Bailey, Cyclop. Am. Hort., 614, 1900. 
F. macrostema var. recurvata Hook., Bot. Mag., 63:3521, 1836; F. recurvata Niven ex 

Hook., I.e. 
Thilcum tinctorium Molina, Sagg. Chile, ed. 2, 146, 1810. 

Leaf -blades mostly 25-50 mm. long, 10-20 mm. wide, membranaceous; 
pedicels mostly 4-5.5 cm. long, red ; hypanthium and sepals red ; sepals 19-23 
mm. long; petals purplish; stamens 30-35 mm. long, reddish. 

Type locality, "Habitat in Conccpcionis Chile locis uliginosis, praesertim in 
Quebrada de Caranio." Ranging mostly in Chile from Aconcagua to Coneep- 
eion. Representative material, without locality ,F. macrostemma, Gay 77, Herb. 
Mus. Paris (F). CPIILE : Aconcagua : 15 km. south of Valparaiso, ^Yest 3980 
(GH). Santiago: Cordillera de Pefialolen, Looser 2299 (POM); Santiago, 
Bro. Claude-Joseph 531 (NY, US). O'Higgins: ]\Ionte la Leona, Rancagua, 
Bertero 280 (GH, NY). Curico: Cordillera Peteroa-Planchoa, Canon Rio 
Claro, Mexia 04406 (GH). Nuble: Banos de Chilian, Pennell 12473 (F, GH, 
US). Coneepeion : Talcahuano, C. and I. Skottsberg 1454 (NY) ; presumably 
from Quebrada de Carcamo, Ruiz and Pavon, as macrostema, Madrid Herb. 



I 



Vol. XXVJ MUNZ: THE GENUS FUCHSIA 11 

(photo F, POM). Bio Bio: Pailalmeqne, Padre Pivian 201 (GH). Cautin : 
Hotel Tolhiiaca on road to Banos Calientes, Morrison and Fagenknecht 17499 
(GH). Magallanes : Port Famine, Lee in 1888 (F, US) ; Eberhard, Donat in 
1931 (CAS, GH, NY). ARGENTINA: Terr, del Neuquen : Puerto Blest, 
Bruch in 1898 (NY) . 

Intergrades have been discussed under var. typica. 

(3c) Fuchsia magellanica Lam. var. Molinae Espinosa 
Fuchsia magellanica Lam. var. Molinae Espinosa, Bol. Mus. Nac. Chile, 12:102, 1929. 

Pedicels and ovaries greenish ; hypanthium, sepals and stamens and style 
pink ; corolla lilac-pink. 

Type locality, CHILE : Prov. Llanquihue : Isla Chiloe, near Castro, Que- 
brada de La Chacra, type coll. by Espinosa, Jan. 31, 1929 (SGO). Another 
collection seen from Isla Chiloe : F. Flaminio Ruiz (POM) . Valdivia : Corral, 
West 4846 (GH), Gnnckel 2133 (GH). Cultivated at Valparaiso, Ooodspeed 
4589 (GH) . There is some question as to whether this is a natural entity or a 
cultivated one, but it seems to be the former. 

(4) Fuchsia coccinea Soland. 
(Plate 1, fig. 4) 

Fuchsia coccinea Soland. in Aitoii, Hort. Kew., ed. 1, 2:8, 1789. 

F. coccinea J. D. Hooker, Bot. Mag., 94: pi. 5740, 1868. 

Nahusia coccinea Schneevogt, Icones, 21, 1793. 

STcinnera coccinea Moench, Meth. Suppl., 270, 1802. 

Fuchsia elegans Salisbury, Ic. Stirp. Ear., 13, pi. 7, 1791 ; said to come from Brazil. 

F. pendula Salisbury, Prodr., 279, 1796; cites plate and page in Stirp. for elegans. 

F. pubescens Cambess., in St. Hil., Fl. Bras. Merid., 2:275, pi. 134, 1829. 

F. montana Cambess., I.e., pi. 135. 

F. Glazioviana Taub., Bot. Jahrb. 15, Beibl., 34:16, 1892. 

Shrub, bushy and up to 1 m. tall, or semiscandent and 3 or more m. tall ; 
freely branched, the younger twigs very slender, scarcely 1 mm. thick, sub- 
terete, densely puberulent, sometimes also pilose, more or less reddish ; leaves 
opposite or in 3's, membranaceous, narrow-ovate, paler beneath, puberulent 
above and beneath, often also pilose beneath along main veins, rounded to 
cordate at base, subacuminate at apex, usually plainly serrulate, the blades 
with 4r-6 main veins on each side of midrib, 15-45 (60) mm. long, 10-20 (25) 
mm. wide ; petioles mostly 1-3 mm. long, frequently pilose ; stipules lance- 
deltoid, less than 1 mm. long, often reflexed, caducous; flowers solitary in 
upper axils ; pedicels filiform, puberulent, sometimes scattered-pilose, 2-A cm. 
long; ovary oblong; hypanthium red, subfusiform- tubular, glandular-puber- 
ulent without, sometimes with few long hairs, pubescent within, 4—6 mm. long, 
ca. 3 mm. wide ; sepals red, glandular-puberulent, oblong-linear to -lanceolate, 
acuminate, 15-20 mm. long, connate at base for 3-6 mm. ; petals violet to 
purplish, obovate, 6-9 mm. long ; episepalous stamens 15-30 mm. long ; epi- 
petalous 2-5 mm. shorter ; anthers reddish, ca. 2 mm. long ; style pubescent at 



12 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

base, 2-4 cm. long; stigma subentire, reddish, 1.5 mm. long; berry mostly 
oblong, about 1 cm. long. 

Type locality : Hooker (Bot. Mag. pi. 5740) says : "The native country of the 
true coccinea is unknown ; it is probably Brazilian, as it resembles the Brazilian 
more than the Western or Southern American species ; Salisbury says it was 
introduced by Vandeli from Brazil, whereas Alton attributes its introduction 
to Captain Frith, from Chile. 

"The evidence of the plant here figured being the true and original plant of 
Alton's 'Hortus Kewensis,' ed. 1, rests on the fact that named specimens of the 
same are preserved in the Banksian Herbarium of the British Museum, and 
in Sir J. Smith's Herbarium at the Linnean Society, all procured at Kew in 
the year of the introduction of the plant, and at the date of its being described 
by Alton." . 

It takes some courage to apply the name coccinea to the Brazilian plant 
commonly passing as puhescens or montana, but I see no alternative. These 
are the only Fuchsias which I can find that agree with all that Hooker says 
above and with specimens seen. ]\Iaterial seen, WITHOUT LOCALITY : F. 
coccinea, Herl). Steud. (NY), Meisner Herh. (NY), Hort. Kew. M. Ait. (G). 
BRAZIL : So. Brazil, Glaziou 17614, F. Olazioviana (fragment from Berlin 
and photograph at F, photograph at POM) ; without locality, F. puhescens 
Camb., St. Hil. C 1545, fragment from Mus. Paris (F). Minas Geraes: F. 
montana Camb., St. Hilaire, fragment from Mus. Paris, must be from Serra 
da Caraca. (F) ; Serra da Piedade, Municipio de Caete, Barreto 7160 (F) , 8803 
(F), 7162 (F), 7158 (F); Serra da Piedade, Hoehne 6278 (POM), 6281 
(POM). There is every intergradation from being merely puberulent as in 
the St. Hilaire specimen of montana, through having some longer hairs at the 
base of the midrib, as in St. Hilaire collection of puhescens, Hoehne 6278, 
6281, Barreto 7160, 7162, 8803, to being pilose on the stems : Barreto 7158, 
garden specimens from Herb. Steudel, etc. The species has been naturalized 
in some places as in Jamaica : Cinchona, Nichols 60 (F, GH) , Harris 9130 (F) 
and Greenhill, Harris in 1899 (F). 

(5) Fuchsia hybrida Hort. 

Fuchsia hyhrida Hort. ex. Vilmorin, Blumeng., ed. 3, Sieb. and Voss, 1 :332, 1894; Behrens, 

Blattenformen, pi. 71, fig. d, e, f, 1900. 
F. speciosa Hort. ex Bailey, Cyclop. Amer. Hort., 614-615, 1900. 

Sturdy, with twigs usually reddish; leaves opposite, broadly ovate to 
oblong-ovate, rounded to cordate at base, apically acute, strongly serrulate, 
3-5 cm. long, 1.5-3 em. wide ; petioles mostly 1-2 cm. long; pedicels commonly 
4r-5 cm. long; ovary ovoid, 6-8 mm. long, 4-5 mm. wide; hypanthium 1-2 or 
more cm. long, 4-6 mm. wide; sepals 2.5-3 cm. long, 8-11 mm. wdde; petals 
1-2 cm. long ; stamens included to exserted ; style 4-5 or more cm. long ; stigma 
3-4 mm. wide. 

Manv manv strains of the garden fuclisias, whicli are almost universally 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 13 

hybrid, are described more or less accurately by the above and are said by 
Bailey (Ilortus, 264. 1930) to be "probably of hybrid derivation from forms 
of F. ynagellanica and F. fulgens." 

(6) Fuchsia regia (Vand.) Munz, new comb. 
QiLclusia regia Vand. ex Veil., Fl. Flum., 149, 1825 and 4:pl. 6, 1827. 

Semiscandent shrub up to 6 m. tall, freely branched, the ultimate branchlets 
slender, 1-2 mm. thick, flexuous, angled, dark red, glabrous to puberulent to 
pilose; leaves opposite or in 3's, oblong-ovate, entire to remotely and incon- 
spicuously denticulate, coriaceous, rounded at base, more or less abruptly 
acuminate at apex (this usually not in same plane as rest of blade and some- 
what folded in pressed specimens), glabrous to puberulent or pilose, often 
reddish on veins ; blades mostly 4-10 cm. long, 2-4 cm. wide, with 5-7 main 
veins on each side of midrib ; petioles 5-10 mm. long ; stipules deltoid, about 
0.5 mm. long, deciduous; flowers solitary in upper axils; pedicels almost fili- 
form, 2-4.5 cm. long, glabrous to pubescent ; ovary oblong; hypanthium deep 
red, cylindric, slightly enlarged toward apex, sometimes somewhat fusiform, 
3-15 mm. long, 3-5 mm. wide, glabrous to pilose without and within ; calyx 
red, glabrous to somewhat pilose, 13-35 mm. long, 4-lobed, the segments con- 
nate for 2-15 mm., the free segments lanceolate, spreading, 10-25 (30) mm. 
long, acuminate; petals purplish, obovate, obtuse, 10-15 (18) mm. long; epi- 
sepalous stamens 20-35 (40) mm. long, epipetalous 16-30 (35) mm. long; 
anthers red, 2.5-3.5 mm. long; style somewhat pubescent at base, 3.5-5.5 (7.5) 
em. long; stigma yellowish, or reddish, capitate, subentire, 2.5-3.5 mm. long; 
berry oblong or more spherical, 1.5-2.5 cm. long. 

The name regia is based on a plate, which shows large leaves and flowers 
and a fusiform hypanthium, sepals connate a short distance. This combination 
of characters and the locality can mean only one thing : the species which has 
been called F. integrifolia. It is fair to admit that the plate shows the leaves 
more dentate than one would like. 

Key to Varieties of Fuchsia regia 
A.Young twigs shaggy-pilose; hypanthium 3-6 mm. long; calyx (united plus free por- 
tions) 13-21 mm. long 6d. var. alpestris. 

AA. Young twigs not pilose; hypanthium 7-15 mm. long; calyx 20-35 mm. long. 

B. Young twigs densely and closely puberulent ; connate portion of calyx 9-15 mm. 

long 6c. var. affinis. 

BB. Young tmgs essentially glabrous. 

C. Connate base of calyx 2-5 mm. long; free segments 18-30 mm. long. 

6a. var. typica. 
CC. Connate base of calyx 9-14 mm. long ; free segments 12-20 mm. long. 

6b. var. radieans. 

(6a) Fuchsia regia (Vand.) Munz var. typica Munz, new name 

(Platel, fig. 5) 
Quelusia regia Vajstd. ex Veil., I.e. 

Fuchsia iniegrifolia Cambess., in St. Hilaire, Fl. Bras. Merid., 2 :273, 1829; Bot. Mag., 68: 

pl. 3948, 1842; MiCHELi in Martins, Fl, Bras. 13, pt. 2:174, pi. 37, 1875. 



14 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Glabrous except sometimes slightly and minutely puberulent about the 
young gro^v1:h ; pedicels 3-4.5 cm. long; hj^panthium 7-15 mm. long; calyx 
24r-35 mm. long, connate at base for 2-5 mm., free portion 18-30 mm. long. 

Tj-pe locality, "ad Alpes Pharmacopolitanas fruticetis, per quae ad oppi- 
dum Cunha iter fit," hence near tlie western end of the state of Rio de Janeiro, 
Brazil. Ranging through the mountains of the states of Rio and Minas Geraes. 
Material seen, BRAZIL : without locality, Burchell2196 (GB.) , Riedelin 1816 
(GH), Gardner 375 (NY). Minas Geraes : Ouro Prieto, Cochran 29191 (US) ; 
Districto Carangola, trail to Areponga, Mexia 4326 (F, CAS, GH, NY, US) ; 
top of Serra da Gramma, Mexia 4278 (F, GH, NY, US) ; Caldas, Claiissen in 
1840 (GH), Regnell in 1878 (NY, US) ; Serra do Cipo, Barreto 9254 (F) ; 
Serra do Caraga, Barreto 7157 (F). Rio de Janeiro: Maceirras, Mt. Itatiaya, 
L. B. Smith 1476 (GH) ; near Rio de Janeiro, Wilkes Exped. in 1838-42 (NY) ; 
Organ Mts., Wilkes Exped. (US). 

Since this is the most common form of the species and is in the Organ Mts. 
and since the original plate shows sepals slightly connate and does not show 
hair, I have to assume that this is typical regia. 

(6b) Fuchsia regia (Vand.) Munz var. radicans (Miers) Munz, new comb. 

F. radicans Miers in Bot. Reg., 27: Misc. no. 167, and pi. 66, 1841. 
F. pyri folia Presl, Symbol, bot., 2 :19, pi. 65, 1858. 

Glabrous throughout; pedicels 2-3 cm. long; hypanthium 9-11 mm. long; 
calyx 2^34 mm. long, connate at base for 9-14 mm. then free for 12-20 mm. 

Type locality, Organ Mts., Rio de Janeiro, Brazil. Material seen, BRAZIL : 
Rio de Janeiro : Organ Mts., 3Iiers 4453, Feb., 1838 (US) ; Serra do Itatiaia : 
Retiro, Dusen 81 (GH, US). Parana : Serra do Mar, Desvio Ypiranga, Dusen 
6766 (GH). 

(6c) Fuchsia regia (Vand.) Munz var. affinis (Camb.) Munz, new comb. 
F. afflnis Cambess., in St. Hilaire, Fl. Bras. Merid., 2 :274, 1829. 

Young twigs densely and closely puberulent (leaves maj- be somewhat 
pilose on under surface along veins); pedicels 2-4 cm. long; hypanthium 
8-10 (13) mm. long; calyx 20-30 mm. long, connate at base for 9-15 mm., 
then free for 10-20 mm. 

Type locality, Serra da Boa Vista, Rio de Janeiro, Brazil. Material seen 
from the states of Sao Paulo and Parana. BRAZIL : without locality, St. 
Hilaire 22, ex Herb. Mus. Paris (F). Sao Paulo : Alto da Serra, at about 900 
m., L. B. Smith 1900 (F, G}l),Mimz 15415 (F, NY, POM, US) ; Est. Campo 
Grande, Edicall 1957 (POM). Parana: Curitiba, Hoehne in 1928 (POIM), 
Dusen 14362 (F, GH, NY). A collection from Horto Botanico Cantareira, 
Puttemans 204 (P) has the puberulence of affinis and the flower of typica. 

(6d) Fuchsia regia (Vand.) Munz var. alpestris (Gardner) IMunz, new comb. 

F. alpestris Gardner, Bot. Mag., 69:pl. 3999, 1843. 
F. mollis Krause, Engl. Jahrb., 37:600, 1906. 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 15 

Young twigs densely shaggy-pilose ; pedicels 2.5-4.5 cm. long; hypanthium 
3-6 mm. long, glandular-pubescent and more or less pilose ; calyx 13-21 mm. 
long, connate at base for 2-6 mm., then free for 10-19 mm. 

T.ype locality. Organ Mts., Rio de Janeiro, Brazil. Material seen ; without 
locality, H. B. Glasgov, F. alpestris, Sept., 1850 (NY). BRAZIL: Minas 
Geraes : Serra do Itacolomy, Municipio de Ouro, Barreto 9168 (F) ; Itacolomi, 
Ouro Preto, de Vasconcellos in 1901 (POM). Rio de Janeiro: bet. Petropolis 
and Theresopolis, Princess Therese von Bayer^i in Herb. Monacense (photos 
F, GH, POM) ; Organ Mts., Gardner 5706, type coll. aliwstris (NY), Nova 
Friburgo, Ule 4418, labelled F. integrifolia var. mollis, a name seeming not 
to have been published, specimen at Rio and Berlin (photo F, POM) . Parana : 
Porto de Cima, Dusen 14133 (US). This entity approaches F. coccinea in 
becoming dentate, but the pilosity is more dense, the leaves usually larger. 
Barreto 9168 is quite like rcgia typica, but more toothed. 

Section 2. EUFUCHSIA Baillon. 

Eiifuchsia Baill., Hist. PI., 6 :467, 1877; Eaimann in Engler and Prantl, Die nat. Pflanzen- 

f am. ,111, 7:219, 1893. 
Fuchsia, as genus, L., Gen. PL, ed. 5, 126, 1754; Spach, Nouv. Ann. Sci. Nat., (2), 4:177, 

1835 ; as section, Endlicher, Gen. PI., 1193, 1840. 
Ellobium Lilja, Linnaea, 15 : 262, 1841. 

Flowers perfect ; petals convolute. Stamens erect, somewhat exserted or not, 
those opposite the petals shorter. Fruit many-seeded. Leaves opposite or 
whorled. 

Type species, F. triphylla L. . • 

Key to Species of Section EUFUCHSIA 

A. Flowers axillary, that is borne in axils of ordinary or somewhat reduced foliage leaves. 
B. Hypanthium 3-30 mm. long. 

C. Hypanthium 3-5 mm. long, shorter than the ovary; young twigs subglabrous. 

Colombia 51. F. verrucosa. 

CC. Hypanthium 12-30 mm. long, longer than ovary. 

D. Younger twigs, pedicels, etc., essentially glabrous (sometimes somewhat 
pilose about nodes) ; leaves mostly glabrous except sometimes along veins 
of under surface. 
E. Petioles 5-20 mm. long; sepals scarlet. S. Peru to Bolivia. 

52. F. sanctae-rosae. 
EE. Petioles 20-35 mm. long; sepals whitish, Ecuador. . . 17. F. pallescens. 
DD. Younger twigs, pedicels, etc., not glabrous. 

E. Hairs on twigs and pedicels spreading ; leaf -blades mostly 5-12 cm. 
long. 
F. Stamens conspicuously exserted ; petals green ; leaves with 7-9 
principal lateral veins on each side of midrib. Mexico to Costa 

Eica 7. F. splendens. 

FF. Stamens not exserted; petals red ; lateral veins 10-15. 
G. Leaves opposite, elliptic to obovate. Ecuador. 

50. F. scabriuscula. 
GG. Leaves mostly ternate, lanceolate to lanceovate. West Indes. 

23. F. triphylla. 






I 
16 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

EE. Ilairs on twigs more or less appressed (if somewhat spreading, then 
leaf -blades usually less than 5 cm. long) ; principal lateral veins 
mostly 3-10 on each side of midrib. 
F. Twigs pubescent; petioles mostly less than 1 cm. long: liypan- 
thium mostly less than 2 cm. long. 
G. Sepals 7-12 mm. long. South America. 

H. Pubescence brownish ; leaf -blades usually not more 
than 3 times as long as wide, pale green beneath, ser- 
rate. Peru 48. F. decussata. 

HH. Pubescence grayish; leaf -blades usually 4 times as 
' long as wide, whitish beneath, subentire to remotely 

serrulate. Ecuador 49. F. hypoleuca. 

GG. Sepals 18-22 mm. long. West Indes 22. F. Fringsheimii. 

FF. Twigs minutely puberulent ; petioles 1-3 cm. long ; hypanthium 

2-3 cm. long. Ecuador - 16. F. loxensis. 

BB. Hypanthium mostly 30-70 mm. long. 

C. Petals narrow, 2-3 times as long as wide. 

D. Young stems essentially glabrous except for the puberulenee near the 

tips ; style glabrous throughout ; petals mostly glabrous. 

E. Hypanthium 35-45 mm. long; sepals long-acuminate. Peru and 

Bolivia. 

F. The hypanthium 5-7 mm. wide at base, usually villous within, not 

much narrowed above ; sepals with green tips ; principal lateral 

veins 12-15 on each side of midrib 10. F. deniiculata. 

FF. The hypanthium 2-3 mm. thick at base, then narrowed; sepals 
apparently red throughout. 
G. Principal lateral veins 7-10 on each side of midrib ; hypan- 
thium glabrous within; sepals 17-20 mm. long. Junin to 

to Cuzco, Peru 11. -F. leptopoda. 

GG. Principal lateral veins 13-15 on each side of midrib ; hypan- 
thium villous within ; sepals 12-14 mm. long. Amazonas, 

Peru 12. F. TToijIl-oicskii. 

EE. Hypanthium 50-60 mm. long ; sepals acute. Colombia. 

13. F. magdalenae. 
DD. Young stems not glabrous ; style more or less pilose on lower portion. 

E. Hypanthium 7-11 mm. wide at summit; petals frequently with some 
hair on backs ; stems puberulent or villous ; fruit 8-10 mm. thick. 
F. The young stems with fine ashy appressed puberulenee ; pedicels 
15-25 mm. long; fruit 8-10 mm. long, subglobose. Central 

Cordillera of Colomliia 24. F. petiolaris. 

FF. The young stems ferruginous-villous; pedicels 25-40 (60) mm. 
long; fruit 12-15 mm. long, ellipsoid. Eastern Cordillera of 

Colombia 25. F. Smithii. 

EE. Hypanthium 5 mm. wide at suiinnit ; petals glabrous; fruit less than 
8 mm. thick. Peru. 
F. Stems, etc., crisp-pubescent ; petioles 10-15 mm. long ; petals 

obtusish. Dept. Piura 25. F. AspJundii. 

FF. Stems, etc., minutely appressed-puberulent; petioles 4-8 mm. 
long; petals subacuniinate. Dept. Amazonas.. .27. F. Llewelynii. 
CC. Petals wider, less than twice as long as wide. 

D.Young stems mostly glabrous, except sometimes (toward tips; style 
glabrous or nearly so to its base. 
E. Petals red. So. America. 



Vol. XXV] MUNZ : THE GEXUS FUCHSIA ' 17 

F. Hypanthium 25-40 mm. long; leaf-blades mostly 2-10 cm. long, 
1-3 cm. wide; petioles 2-10 (15) mm. long. Peru. 

9. F. austromontana. 

FF. Hypanthium 60-70 mm. long; leaf -blades 8-18 cm. long, 4-7. (9) 

cm. wide; petioles 10-30 mm. long. Ecuador. 20. F. macrostigma. 

EE. Petals green. Guatemala 8. F. cordi folia. 

DD. Youug stems hirtellous to pilose ; style pilose or villous near base. 

E. Twigs hirtellous with short subappressed hairs; petioles 1-5 (8) mm. 
long ; pedicels 1-2 cm. long. 
F. Leaf -blades 2-4 cm. long, 1-2 cm. wide ; petioles 3-8 mm. long. 

Colombia and Ecuador 14. F. coneacens. 

FF. Leaf -blades 5-6.5 cm. long, 2.5-3 cm. wide; petioles quite lack- 
ing. No. Peru 15. F. rivularis. 

EE. Twigs pubescent to pilose, with spreading hairs; petioles 5-20 mm. 
long; pedicels 15-45 mm. long. Ecuador and Peru. 
F. Hypanthium narrow, 2 mm. thick at base, 5-6 mm. wide at 
summit; sepals 10-12 mm. long, somewhat divergent. Ecuador. 

18. F. Townsendii. 

FF. Hypanthium -wider, 3-5 mm. thick at base, 7-11 mm. wide at 

summit ; sepals 15-22 mm. long, spreading-reflexed. 

G. Hypanthium gradually ampliate, hence widest at its mouth ; 

sepals 3-4 mm. wide; petals longer than wide. Dept. Apuri- 

mac, Peru 19. F. plafi/petola. 

GG. Hypanthium abruptly ampliate near middle and slightly 
narrowed at summit ; sepals 5-6 mm. wide ; petals scarcely 
longer than wide. Ecuador to Ayabaca, Peru. 

21. F. ayavacensis. 
AA. Flowers in clusters, making a definite inflorescence with bracts or leaves much reduced 
and different from foliage leaves. 
B. Hypanthium 1-3 cm. long. 

C. Young twigs essentially glabrous except for youngest tips. 

D. Leaf -blades generally at least 3 times as long as wide; hypanthium 13-16 
mm. long; petioles 2-4 mm. long. Colombia and N. Ecuador. 

63. F. sessilifolia. 
DD. Leaf -blades usually 2-3 times as long as wide; hypanthium usually 16-30 
mm. long. 
E. Leaf -blades mostly 5-8 cm. long, 1.5-3.5 cm. wide ; style glabrous. 
' F. Flowers in elongate raceme-like inflorescences, with more or less 

reduced foliage-leaves. S. Peru and Bolivia. 52. F. sanctae-rosae. 
FF. Flowers in definite very short and compact racemes. North of 
So. Peru, 
G. Hypanthium 24-30 mm. long; sepals 9-11 mm. long; prin- 
cipal lateral veins on each side of midrib 8-12. 
H. Sepals oblong-ovate, 5-6 mm. wide; berry over 1 cm. 
long ; pedicels 15-25 mm. long. Peru. . . 53. F. Osgoodii. 
HH. Sepals linear-lanceolate, 2.5-3 mm. wide; berry 5-6 
mm. long ; pedicels 8-18 mm. long. Ecuador. 

55. F. Lehmannii. 
GG. Hypanthium 14-18 mm. long; sepals 7 mm. long; principal 
lateral veins about 15. Putumayo, Colombia. 

56. F. pniumayensis. 
EE. Leaf-blades mostly 8-20 cm. long, 3-9 cm. wide, with 12-18 principal 
lateral veins on each side of midrib; style pubescent near base. 



18 CALIFOEXIA ACADEMY OF SCIENCES [Proc.4thSer. 

F. Petioles 2-8 mm. long; pedicels 5-7 mm. long; petals almost as 

wide as long; leaves quite glabrous. Ecuador. 61. F. glaberrima. 

FF. Petioles 10^0 mm. long ; pedicels mostly 10-20 mm. long ; petals 

about 3 times as long as wide; leaves puberulent along veins 

underneath. 

G. Leaf -blades mostly obtuse at base; fruit ca. 6 mm. long. 

Colombia 54. F. Andrei. 

GG. Leaf -blades subacuminate at base; fruit 8-10 mm. long. 

Peru 62. F. macrophylla. 

CC. Young twigs definitely not glabrous. 

D. Leaf -blades mostly 5-8 cm. wide ; pedicels 10-30 mm. long. 

E. Hypanthium 13-15 mm. long ; racemes mostly lateral ; pedicels 10-15 

mm. long. Hiiauuco, Peru 31. F. ovalis. 

EE. Hypanthium 20-30 mm. long; racemes terminal; pedicels 15-30 mm. 

mm. long. Dept. Cuzco, Peru M.F. tincta. 

DD. Leaf -blades mostly 1.5-5 cm', wide; inflorescence often terminal. 

E. Stems with fine close appressed puberulence ; hypanthium very nar- 
row, scarcely bulbous at base, 14r-19 mm. long, 2.5-4 mm. wide at 

mouth. Colombia and Venezuela to Ecuador 64. i^. sylvatica. 

EE. Stems pubescent to pilose ; hypanthium somewhat bulbous at base. 

F. Hairs reddish 37. 2?". StorUi. 

FF. Hairs not reddish, 

G. Flowers in terminal pyramidal panicles; hypanthium 14-20 

mm. long. Colomlna 57. F. Harticegii. 

GG. Flowers in racemes. 

H. Twigs pubescent ; hypanthium 15-20 mm. long. Santo 

Domingo and Haiti 23. F. triphylla. 

HH. Twigs pilose; hypanthium 20-25 mm. long. Peru. 

I. Bracts small, subulate or narrow; petioles of a 

given wliorl subequal 59. F. asperifolia. 

II. Bracts more like reduced leaves ; petioles of a 
whorl very unequal, one much exceeding the otliers. 

60. F. pilosa. 
' BB. Hypanthium 3-6.5 cm. long. 

C. Leaves small, crowded, 12-16 mm. long ; stems with brown hair. Peru. 

32. F. confertifoUa. 
CC. Leaves larger, 30-200 mm. long. 

D. Leaf -blades narrow, more than 4 times as long as wide, in whorls of 4, 

linear-lanceolate. Huanuco, Peru 31. F. simplicicaulis. 

DD. Leaf-blades wider, 2-3 times as long as wide, opposite or ternate (some- 
times quaternate in F. hirtella). 
E, Petals (under a lens) puberulent to sparsely villous on back. 

F. Sepals pubescent on inner or upper surface; fruit oblong; leaf- 

blades rigid. Colombia and Venezuela 28. F. venusfa. 

FF. Sepals glabrous on inner surface ; fruit subglobose ; leaf -blades 
membranaceous. Venezuela. 
G. Young stems appressed-puberulent ; pedicels 20-40 mm. 
lolig; hypanthium 35-40 mm. long; sepals spreading, 18-20 

mm. long; petals strigulose on back 29. F. Jaltnii. 

GG. Young stems glabrous or somewhat villous ; pedicels 15-20 
mm. long; hypanthium 40-50 mm. long; sepals divergent, 
14-15 mm. long; petals sparingly villous on back. 

30. F. Gehriperi. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 19 

EE. Petals glabrous on back. 

r. Young stems glabrous ; leaves subglabrous above, puberulent 
beneath ; petioles 2-8 mm. long. 
G. Leaf -blades 3 times as long as wide ; principal lateral veins 
ca. 20 on each side of midrib; pedicels 25-40 mm. long. 

Peru 43. F. abrupta. 

GG. Leaf -blades less than 3 times as long as wide ; principal 
lateral veins 10-12 on each side of midrib; pedicels 7-13 

mm. long. Colombia 44. F. Cuatrecasasii. 

FF. Young stems not glabrous under a lens. 
G. The young stems finely puberulent. 

H. Principal lateral veins about 12 on each side of midrib ; 
berry 5-6 mm. long; pedicels 10-40 mm. long. Colom- 
bia 45. F. Eillipii. 

HH. Principal lateral veins 14-24 on each side of midrib ; 
berry 6-12 mm. long; pedicels 6-15 mm. long. 

I. Leaves quite glabrous ; berry subglobose, almost 

1 cm. long. Junin, Peru 39. F. Munzii. 

II. Leaves not glabrous under a lens ; berry longer 
than thick. 
J. Leaf-blades densely soft-puberulent, with 14- 
16 principal lateral veins on each side of mid- 
rib; style pubescent near base. Ecuador to 

Huanuco, Peru 42. 2^. corymbiflora. 

JJ. Leaf -blades glabrous on casual inspection, but 
minutely strigulose under a lens, with 18-24 
principal lateral veins ; style almost glabrous. 

Libertad, Peru 30. F. Aspiazui. 

GG. The young stems pubescent to villous (very sparsely pubes- 
cent in F. fidgens). 
H. Stems and leaves pilose or villous with brownish or 
reddish hairs ; pedicels 15-40 mm. long. 

I. Sepals 15-20 mm. long ; hypanthium 35-45 mm. 

long. Bolivia 38. 2^. furfuracea. 

II. Sepals 8-12 mm. long. Peru. 

J. Hypanthium 20-35 mm. long; petioles 10-50 

mm. long. Cuzco 34. F. tincta. 

JJ. Hypanthium 40-45 mm. long ; petioles 2-5 
mm. long. 
K. Leaves ternate, broadlj^ elliptic. Ama- 

zonas 35. F. Mathewsii. 

. KK. Leaves opposite, oblong-lanceolate. Caja- 

marca 36. i^. Fischeri. 

HH. Stems and leaves strigulose to pubescent ; pedicels 
mostly 5-20 mm. long. 

I. Sepals red, pubescent on backs. Mostly So. Ameri- 
can. 
J. Sepals somewhat divergent ; hypanthium 
strigulose without. 
K. Leaves membranaceous, the apex in the 
same plane as the lilade ; berry over 1 cm. 
long. Central Colomhia.. .40. F. hi rfella. 
^ KK. Leaves rigid, rugose, the apex l)ent to 



20 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

one side in pressing ; berry less than 1 cm. 
long. So. Colombia and No. Ecuador. 

41. F. polyaniha. 

J J. Sepals spreading-reflexed ; hypantliium -with 

spreading hairs. West Indies and Central 

America to Argentina 46. F. boJiviana. 

II. Sepals green, glabrous except for the sparse pubes- 
cence near the tips, divergent-spreading. Mexico. 

47. F. fulgens. 

(7) Fuchsia splendens Zuccarini 

(Plate 1, fig. 6) 

Fuchsia splendens Zucc, Flora, 1832, ii, Beibl., 102, 1832. 

F. splendens in Curtis, Bot. Mag., 70:pl. 4082, 1844; Bot Eeg., 28:pl. 67, 1842; Hemsley, 

in Bot. Biol. Centr. Amer., 1:460, 1880. 
F. cordifolia /3 Hooker, Icon. PI., 5:450, 1842. 
F. intermedia IIemsley, Diag. PI. Nov., 1 : 14, 1878 ; Bot. of Biol. Centr. Amer., 1 :457, 1880. 

Based on Hartweg 460 which I have not seen, but the characters given do not hold in a 

series of specimens. 

Shrub 6-25 dm. tall, often pendant, sometimes a small tree, relatively few- 
branched, the ultimate branchlets usually pilose or pubescent, greenish to 
reddish; leaves opposite, ovate to ovate-cordate, paler beneath than above, 
membranaceous, more or less pubescent especially when young, acuminate, 
denticulate, the blades 3-10 cm. long, 2-6 cm. wide, with or without reddish 
tinge, principal lateral veins 7-9 on each side of midrib ; petioles often tinged 
red, pilose to subglabrous, slender, 2-5 cm. long; flowers solitary in axils or 
more or less crowded on short lateral branches with somewhat reduced leaves ; 
pedicels nodding, subfiliform, pubescent, with some gland-tipped hairs, 3-5 
(8) cm. long, not much elongate in fruit; hypanthium 17-20 (40) mm. long, 
somewhat compressed, the flattened side much broader than the ovary and 
gradually ampliate upward, 6-8 mm. wide at base, 7-12 mm. wide at summit, 
more narrow in other dimension, rose to bright red, externallj' pubescent with 
some hairs gland-tipped, internally pubescent-villous ; sepals 4, green except 
for the sometimes reddish base, ovate-lanceolate, 8-15 mm. long, 5 mm. wide 
at base, pubescent, the tips scarcely free or free for 1 mm.; petals green, ovate, 
subacuminate, 6-10 mm. long, 4-5 mm. wide ; episepalous stamens exceeding 
the sepals 10-18 mm. long, epipetalous 8-16 mm. long ; filaments pale yellow, 
anthers yellow, 2-3 mm. long; style glabrous, exceeding the stamens; stigma 
conoid, greenish-yellow ; berry linear, 3-3.5 cm. long, 5 or more mm. thick. 

Type locality, "in imperio mexicano." Ranging from Chiapas to Costa Rica, 
at altitudes of 2000 to 10,000 feet. Variable in pubescence, some plants being 
cinereous (Matuda 2322, Steyermark 36056, Pittier 10501, Standley and 
Valerio 43829) , others with longer pubescence (Skuteh 118, Pittier 44, Steyer- 
mark 32833). But there seems to be no geographical or otlier correlation with 
variability in pubescence. There is also, at least in herbarium material, con- 
siderable variation in relative length and width of hypanthium. Curtis, Bot. 

Mag. 4082, mentioned the striking compression of the hypanthium, "so that 

t 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 21 

ill one direction, the diameter of the tube does not appear to be half what it is 
in another" ; this would account for some of the differences in herbarium 
specimens. However, material from Costa Rica has a long hypanthium which 
narrows appreciably at the summit, at least in Standley and Valerio 43829, 
Pittier 10501, not so much so in Stork 3024. Curtis shows such a condition in 
edge-view, but not in the flat view. More material from Costa Rica may 
warrant nomenclatorial segregation. 

Representative material of F. splendens: H. B. Basil, May 7, 1849 (NY) ; 
"F. grandiflora N." Sesse, Mogiilo et al 5208 (F). MEXICO: F. splendens 
Zuccar. ipse, Hab. in Mexico, Karivinski, Herbarium Regium Monacense 
(photo F, GH, POM). Chiapas, without locality, Ghieslreght 698 (F, GH, 
MO) ; Mt. Tacana, Matuda 2322 (F, MICH, NY) ; between San Cristobal and 
Tencjapa, Souviron and Erianson 83 (US). GUATEMALA: San Marcos, 
Quebrada Canjula, between Sibinal and Canjula, Volcan de Tacana, Steyer- 
mark 36056 (F) . Quiche, Chiul, Heyde and Lux, under J. D. Smith 2890 (NY, 
GH, US) ; Chimaltenango : Chichavac, Skutch 118 (US) ; Santa Elena, Skutch 
220 (US) ; Cerro de Tecpam near Santa Clara, Standley 61000 (F, NY, 
POM) ; Volcan de Acatenango, above Las Calderas, Steyermark 61808 (F). 
Sacatepequez, Volcan de Agua, J. Donnell Smith 2174 (F, GH, NY, US), 
Standley 65133 (F, POM) ; Jalapa, Montaiia Miramundo, Steyermark 32833 
(F). COSTA RICA: prov?, Ojo de Agua, Stork 3024 (F) ; Prov. San Jose, 
Cerro de las Vueltas, Pittier 10501 (US), Standley and Valerio 43829 (F, 
US). 

(8) Fuchsia cordifolia Bentham 

(Plate 1, fig. 7) 
Fuchsia cordifolia Bentham, PI. Hartweg., 74, 1841; Lindl., Bot. Reg., 27:70, 1841. 

Straggling shrub 1 m. or so high, to a small tree, often epiphytic, few- 
branched, the younger branches somewhat red, subglabrous except toward the 
glandular-pub erulent tips ; leaves ovate to cordate-ovate, opposite, paler be- 
neath than above, puberulent when young, nearly or quite glabrate in age, 
more or less abruptly acuminate, obscurely denticulate; blades thin, 3-9 cm. 
long, 2-6 cm. wide, with 9-11 principal lateral veins on each side of the mid- 
rib ; petioles 2-5 cm. long ; flowers solitary in axils of upper often somewhat 
reduced leaves; pedicels pendant, subfiliform, puberulent, 3-8 cm, long at 
anthesis, not much longer in fruit ; hypanthium subcylindric, dull red, puber- 
ulent without, some of the hairs gland-tipped, glabrous within, somewhat 
swollen at base, then slightly narrowed and somewhat broader only near the 
summit, 3.5-5 (6) cm. long, 4-5 mm. wide except at the summit, where 7 mm. 
wide; sepals mostly green except at reddish outer base, ovate-lanceolate, 
puberulent with some hairs gland-tipped, acuminate, 11-15 mm. long, 4-5 
mm. wide at base, free tips 1-2 mm. long; petals green to olive-green, broadly 
ovate, glabrous, acute to subacuminate, 6-8 mm. long; episepalous stamens 
about as long as sepals, epipetalous about two-thirds as long ; anthers yellow, 



22 CALIFOEXIA ACADEMY OF SCIENCES [Proc. 4th 8er. 

2-2.5 mm. long; style glabrous, slightly exceeding the sepals; stigma conic. 
1.5 mm. long; fruit apparently linear-ellipsoid, more than 2 cm. long. 

Type locality, "In monte ignivomo Xetucli infra apicem, altitudine ped. 
10,000," prope Quezaltenango, Guatemala. Tlie species is known from eleva- 
tions of 8000-11000 ft. from the mountains of Guatemala. It is distinguished 
from F. fulgens by its green petals and from F. splendens hy its shorter 
stamens, the hj^Danthium being glabrous within and longer and subcylindric. 
Specimens studied, GUATEMALA : Dept. San Marcos, Volcan de Tajumulco, 
between San Sebastian and Todos Santos, Steyermark 36942 (F) ; Dept. de 
Quezaltenango, Volcan de Zunil, Steyermark 34686 (F); Cerro Quemado. 
Kellerman 5953 (US) ; Volcan de Santa Maria, Nelson 3723 (F, US), Skntch 
867 (F, NY), Standley 67592 (F), 67648 (F) ; Xetuch, Hartiveg 528, frag- 
ment of type number (F) , photo of type no. (F, POM) ; Dept. Chimaltenango, 
San Martin, Lcivis 936 (F.) 

(9) Fuchsia austromontana Johnston 
(Plate 1, fig. 8) 

Fuchsia austromontana Johnston, Journ. Arnold Arboi-etum, 20:242, 1939. 
F. serrati folia Hook., Bot. Mag., 71 :pl. 4174, 1845 ; not of K. and P. 

Loose bushy shrubs up to 4 m. tall; twigs reddish, 1.5-3 mm. thick, glabrous 
except for the pilose tips ; leaves opposite or ternate (or quaternate ?) , elliptic 
to elliptic-lanceolate or -ovate, acute to obtuse at base, acuminate at apex, 
serrulate, strigulose (at least along the veins) and light green on upper sur- 
face, paler beneath and villous on veins, rather fleshy, with 8-15 principal 
veins on each side of midrib, the submarginal vein often inconspicuous ; blades 
2-10 (12) cm. long, 1-3 (4) cm. wide, upper ones considerably reduced; 
petioles 2-10 (15) mm. long, villous; stipules lanceolate, 2-3 mm. long, decidu- 
ous ; flowers in upper axils ; pedicels stoutish, 1-3.5 cm. long, sparsely strigu- 
lose; ovary fusiform, sparsely strigulose, 8-10 mm. long, 2-2.5 mm. thick; 
hypanthium light red, subcA'lindric, 2.5-4 cm. long, enlarged at base (3.5-5 
mm. thick) , then narroAved for about 1 cm. (i.e. 2-3 mm. thick) , then gradually 
wider toward summit (ca. 7 mm. wide), sparsely villous without, villous 
wdthin lower portion ; sepals red, lanceolate, 15-17 mm. long, 4 mm. wide, 
subacuminate (bud subacuminate, but scarcely apiculate), pubescent to vil- 
lous without, glabrous within ; petals broadly obovate, rounded at apex, 1^15 
mm. long, 11-12 mm. wide, red, sometimes purple on drying; stamens 16-17 
and 10-11 mm. long; anthers 4-5 mm. long; .style glabrous, 2-3 mm. longer 
than sepals; stigma 2-3 mm. thick; berry broadly ellipsoid, about 15 mm. 
long, 8 mm. wide. 

Type locality, between Pillahuata and Acanacu, prov. Paucartambo, dept. 
Cuzco, Peru. Near to F. dcnticuJata in the broad hypanthium and glabrous 
style and to F. cordifolia in the broad hypanthium-base. It seems to be rather 
local in Paucartambo ; I have seen the following collections : PERU : Paucar- 
tambo : between Pillahuata and Acanacu, at 2800 m., West 7083, isotype 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 23 

(GH) ; Pillahuata, Cerro de Cusilluyoc, 3000-3300 m. Pennell 14110 (F, GH, 
US) ; Cabecera de Montana, eamino a Tambomayo, 2000 m.. Vargas 10-7083 
(F) ; between Acliirani and Medias-Mayu, 2600 m., Vargas 11121 (F) ; Acca- 
naco, 4000 m., Balls 6708 (US) . 

(10) Fuchsia denticulata Rniz and Pavon 

(Plate2, fig. 9) 

Fuchsia denticulata Euiz and Pavon, F1. Peruv., 3:87, pi. 325, fig. b, 1802. 

F. fierrati folia Euiz and Pav6n, I.e., 86, pi. 323, fig. a. Since tins name has page priority, 
I had intended to use it, but must follow Art. 56 of the International Eules, since Mac- 
bride took up this name (Field Mus. Nat. Hist.-Bot. Ser., 13, part 4:553, 1941). 

F.tacsoniiflora Krause, Fedde Eep. Nov. Sp., 1:172, 1905; based on Weberbauer 252, 
which seems to belong here. 

F.grandiflora Euiz ex Dahlgren, Field Mus. Nat. Hist.-Bot. Ser., 21:212, 1940, nomen 
nudiim. Have seen photograph of Geneva specimen (F, POM). 

Semiscandent shrub to tree, up to 12 m. tall, the young twigs green to red- 
dish, mostly 3-5 mm. thick, subterete, mostly glabrous, more or less puberu- 
lent or pubescent toward tips, about younger nodes, etc. ; leaves opposite or 
ternate, elliptic to oblong, rather fleshy, subentire to somewhat serrate, acute 
to obtuse at base, acute to acuminate at apex, darker above than beneath, 
subglabrous to puberulent above, subglabrous beneath or somewhat villous 
along veins, the principal lateral veins 12-15 on each side of midrib, sub- 
marginal vein evident; blades 4-11 cm. long, 2-4 (5) cm. wide; petioles 5-15 
mm. long, 1-2 mm. wide, glabrous to pubescent ; stipules lance-deltoid, ca. 1 
mm. long, deciduous; flowers solitary in axils of upper somewhat reduced 
leaves; pedicels rather stout, 1.5-4 cm. long, usually minutely strigulose; 
ovary subfusif orm, 10-12 mm. long, 2-3 mm. wide, minutely strigulose ; hy- 
panthium tubular, red, 3.5-4 cm. long, 5-7 mm. wide at base, very little nar- 
rowed, then very gradually widened toward mouth where 7-10 mm. wide, 
subglabrous to sparsely villous without, usually densely villous within lower 
portion ; sepals red with green tips, lanceolate, divaricate, 18-24 mm. long, 
4-5 mm. wide, acute to acuminate (bud acuminate) in the terminal 3-4 mm. 
but scarcely apiculate, glabrous to somewhat villous on backs, glabrous on 
inner surface; petals crimson to scarlet, oblong to oblong-lanceolate, acute to 
obtuse, 15-17 mm. long, 4-6 mm. wide, glabrous ; stamens 17-20 and 13-16 
mm. long, glabrous ; anthers white, 4-5 mm. long ; style stout, glabrous, equal- 
ling or exceeding sepals by a few mm. ; stigma white, subglobose, ca. 3 mm. 
thick ; berry ellipsoid, 16-19 mm. long, ca. 8 mm. thick, subglabrous to min- 
utely strigulose. 

Type locality, Muiia, dept. Huanuco, Peru. AVith F. leptopoda, etc. this 
species forms a long-tubed, axillary-flowered group related to F. cordifolia, 
F. austromontana, etc. I am unable to separate F. serratifolia and F. denticu- 
lata, type material of both of which has been available. They are characterized 
by their stout twigs, fleshy, elliptic subglabrous leaves, tubular hypanthium, 
non-apiculate buds, ellipsoid berry. Found in Peru and Bolivia ; repreSenta- 



24 CALIFORXIA ACADEMY OF SCIEXCES [Proc. 4th Ser. 

tive material as follows: F. denticulata, ex Herb. Madrid, Rtiiz and Pavon, 
type material (F), photograph of Geneva material (F, POM) ; F. serratifoUa 
ex Herb. Madrid, Ruiz and Pavon, type material (F), photograph (F, POM) ; 
F. grandiflom, photograph of Geneva specimen (F, POM). PERU : Huaniico, 
15 mi. se. of Huanuco, 3200 m., Machride and Featherstone 2082 (F, GH, US) ; 
Mito, 2800 m., Machride and Featherstone 1403 (GH, F, US) ; west of Carpish, 
Stork and Horton 9893 (F) ; Muiia, trail to Tambo de Vaca, 2500 m., Machride 
4284 (F, GH, US) ; Carpis, between Acamayo and Chinehao, 2800 m., Mexia 
04127 (GH) ; Lima to Oroya, 3200 m., Wcherhauer 252, type tacsoniiflora at 
Berlin (photo F, PO:\r) ; Rio Blanco, Lima at 3600 m., Machride and Feather- 
stone 723 (F, GH) ; Quebrada de San Mateo, Isern 2546 (F). Ayacucho, 
Prov. Huanta, Choimacota Valley, 2900 m., Wcherhauer 7587 (F, GH, US). 
Huaneavelica : Prov. Tayacaja, Montifungo, east of Surcubamba, 3000 m., 
Stork and Horton 10367 (F) ; Salcabamba, 3250 m., Stork and Horton 10261 
(F). Cuzco: Prov. Paucartambo, Rio Tambomayo, 1700-2000 m., West 7093 
(GH). BOLIVIA : La Paz : Nor Yungas : Unduavi, 3000 m., Eyerdam 25137 
(F) ; Sur Yungas : Unduavi, 3100 m., Buchtien 143 (F, GH, 1<^Y),2923 (NY, 
US) ; Yungas, Ba^ig 731 (F, GH, NY, US), Rnshij 1801 (NY) ; Combaya, 
Prov. Larecaja, 3000-3300 m., Mandon 623 (GH). 

(11) Fuchsia leptopoda Krause 

(Plate 2, fig. 10) 

Fuchftia leptopoda Krause, Pedde Eep., Nov. Sp., 1:171, 1905. 
F. siphonantha Krause, I.e., 173. 

Scandent to erect shrub, up to 3 m. tall, the twigs dark, glabrous to strigu- 
lose toward tips, terete, 1-3 mm. thick; leaves opposite or ternate, elliptic- 
lanceolate or -oblanceolate, acute at base, acuminate at apex, serrulate above, 
light green and glabrous or strigulose on veins, paler beneath and glabrous 
except for pilose veins, the principal lateral veins mostly 7-10 on each side 
of midrib, the submarginal vein more or less developed; blades 4-9 cm. long, 
1.5-2.5 (3) cm. wide; petioles slender, 5-15 mm. long, somewhat pubescent; 
stipules lanceolate, 3 mm. long, deciduous ; flowers in axils of often reduced 
foliage leaves; pedicels slender, subglabrous, 3-5 cm. long; ovary fusiform, 
glabrous to puberulent, 5-8 mm. long ; hypanthium dark red, tubular, 3.5-4.5 
cm. long, somewhat bulbous at base and ca. 3 mm. thick, then narrowed to 
width of 1.5-2 mm. for about 1 cm., then widened toward mouth where 6-7 
mm. wide, glabrous to sparsely villous without, densely villous within lower 
portion ; sepals deep red, lance-linear, 17-20 mm. long, 3.5-4 mm. wide, acumi- 
nate (bud very slender and long-acuminate, with tip 4-5 mm. long), glabrous 
to pubescent on back, glabrous within, petals fiery red, oblong, 17-18 mm. 
long, 4-7 mm. wide, acute, glabrous or sparsely villous on back near base ; 
stamens 18-22 and 12-15 mm. long, glabrous; anthers 3-4 mm. long; style 
glabrous, equalling sepals or exceeding them by 1 cm., stigma subglobose, 
obscurely lobed, 2-3 mm. wide ; fruit narrow, over 1 cm. long. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 25 

Type locality, PERU : Juniii : Prov. Tarma, between Huacapistana and 
Palea. I have seen the following examples : PERU : Junin : Prov. Tarma : 
between Huacapistana and Palea, 2200-2500 m., Weherhauer 1772, type at 
Berlin (photo at F, POM) ; Montibus ab Huacapistana ad orientem versus, 
2600-2700 m., Welerlauer 2178 type siphona7itha at Berlin (photo F, POM) . 
Ayacucho : Prov. De La Mar : between Tambo and the Apurimac, Yanamonte, 
2700-2800 m., Weherhauer 5641 (F, GH, US) ; Prov. Huanta, Choimacota 
Valley, 2900 m., Weherhauer 7587a (F, GH). Cuzeo : Paucartambo: Tres 
Cruces, above Cosnipata, Weherhauer 6934 (F, GH, US). In general, this 
species is very near to F. denticulata but with a more narrow tube and leaves, 
long-pointed buds, etc. 

(12) Fuchsia Woytkowskii Macbride 

(Plate2, fig. 11) 

Fuchsia WoyfkowsTcii Macbride, Field Mus. Nat. Hist., Bot. Ser., 13: part 4:566, 1941. 

Shrub up to 4 m. tall, the young twigs glabrous, purplish red, slender; 
leaves opposite or ternate, elliptic-lanceolate to -ovate, acute at base, acute to 
acuminate at apex, subentire, rigid, dark and glabrous above, somewhat paler 
and glabrous beneath except for the minute appressed puberulence on the 
veins, principal lateral veins 13-15 on each side of the midrib, submarginal 
vein evident; blades 4-8 cm. long, 1.5-3 cm. wide; petioles 3-6 mm. long; 
stipules subulate, 2 mm. long, deciduous; flowers solitary in axils of upper 
leaves or apparently subterminal on short branches ; pedicels purplish, sub- 
filiform, nearly glabrous, 1-2.5 cm. long; ovary fusiform, 5-6 mm. long, 2 mm. 
wide; hypanthium deep vermilion, 3.5-4.7 cm. long, bulbous at base, 2-2.5 
mm. wide, then narrowed for one-third its length then gradually widened until 
6-7 mm. wide at mouth, glabrous without, pilose within the basal two-thirds ; 
sepals vermilion, spreading, 12-14 mm. long, 4-5 mm. wide, glabrous, sub- 
ulate for 2-3 mm. at apex (the tips apparently not connivent in the bud) ; 
petals bright red, oblong-elliptic, 12-13 mm. long, glabrous, erose-rounded at 
apex; stamens 10 and 8 mm. long; anthers 2 mm. long; style glabrous, sur- 
passing sepals by 2-3 mm. ; stigma 2 mm. thick. , 

Type, PERU : Dept. Amazonas : near Almirante, at 1900 m., F. WoytkowsH 
(F). This one and only specimen is rather inadequate, but apparently does 
not fit into any of the previously described species. It seems near F. leptopoda, 
but the hypanthium is more slender and not glabrous within, and the sepals 
are shorter and with free tips in the bud. 

(13) Fuchsia magdalenae Munz, new species 

(Plate2, fig. 12) 

Shrub; young twigs glabrous, purplish, 1.5-3 mm. thick; leaves opposite, 
ternate or quaternate, coriaceous, elliptic, rounded at base, abruptly acumi- 
nate at apex, inconspicuously denticulate, quite glabrous except for the ciliate 
margins, somewhat paler beneath than above, principal lateral veins dark. 



26 CALIFOIIXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

8-10, the submarginal vein more or less distinct; blades 3-6 cm. long, 1.5-2.5 
cm. wide, the upper reduced; petioles dark, glabrous, 5-6 (10) mm. long; 
stipules subulate, 1 mm. long, deciduous; flowers solitary in upper axils, sub- 
filiform, glabrous, 4-5 cm. long, dark; ovarj' glabrous, 7-11 mm. long, 2-2.5 
mm. wide ; hypanthium purplish at base, brigliter red beyond, 5-6 cm. long, 
about 4 mm. wide at base, then not constricted but tubular and gradually 
enlarged until 8-9 mm. wide at mouth, glabrous without and within; sepals 
lance-oblong, 16-19 mm. long, 4.5-5 mm. wide, red, glabrous, acute, not sub- 
ulate at apex; petals scarlet, oblong-ovate, 15-18 mm. long, 7-8 mm. wide, 
acutish ; stamens 17 and 13 mm. long ; anthers 3.5 mm. long ; style slender, 
glabrous, exceeding sepals by 5-6 mm.; stigma 4-lobed, 4 mm. wide; berry 
ellipsoid, ea. 17 mm. long, 8 mm. wide. 

Frutex; ramulis glabris, subpurpureis, 1.5-3 mm. diametro ; foliis oppositis, ternatis vel 
quaternis, coriaceis, ellipticis, basi rotimdatis, apice abrupte aeumiiiatis, subglabris; laminis 
3-6 cm. longis, 1.5-2.5 cm. latis; petiolis glabris, 5-6 (10) mm. loiigis; stipulis siibulatis, 
1 mm. longis, caducis; floribus solitariis in axillis superioribus ; pedicellis subfiliformibus, 
glabris, 4—5 cm. longis; ovario glabro, 7-11 mm. longo; hypanthio rubro, 5-6 cm. longo, 
basi 4 mm. lato, tubuloso, supra 8-9 mm. lato, c.xtus intiisquo glabro ; sepalis lanceolato- 
oblongis, 16—19 mm. longis, 4.5-5 .mm. latis, rubris, glabris, acutis; petalis coccincis, oblongo- 
ovatis, 15-18 mm. longis, 7-8 mm. latis, subacutis ; staminibus 17 vel 13 mm. longis ; antheris 
3.5 mm. longis; stylo glabro, sepala excedente; stigmate 4— lobato; bacca ellipsoidea, ca. 17 
mm. longa, 8 mm. lata. 

Type : Colombia : Dept. Magdalena : above San Miguel, at edge of paramo, 

3000 m., July, 1932, Seifriz 392, United States Nat. Herb. no. 1572275. A 

second collection from the same department is from between Pueblo Viejo and 

San Miguel, 900-1700 m., 8eifriz 537 (US). The proposed species is related 

to F. dcnticulata in its tubular hypanthium ; it is characterized by its long 

hypanthium, thickish leaves, glabrous condition, non-acuminate sepals and 

northern range. /-i.N-rr i, • t-. xi 

(14) Fuchsia canescens Bentham 

(Plate 2, fig. 13) 

Fuchsin canescens Benth., PI. Hartweg., 178, 1845. 
F. eorollata Benth., I.e., 179. Type number seen. 

F.vulcnnica Andr^, Rev. Hortic, 233 and 268, 1888; from Yolcan de Azufral, type not 
seen, but characters as given agree. 

Erect or clambering shrubs up to 3 m. tall, freely branched, tlie young twigs 
terete, dark, hirtellous with more or less appressed short hairs; leaves opposite, 
ternate or quaternate, crowded, coriaceous, often with revolute margins, ellip- 
tic-ovate, obtuse or rounded at base, obtuse to acute at apex, serrulate, dark 
green and pubescent to glabrescent above, paler beneath and pilose on veins, 
sometimes more generally, the principal lateral veins mostly 4-7 on each side 
of the midrib, submarginal vein often not conspicuous ; leaf-blades 2-4 cm. 
long, 1-2 cm. wide; petioles 3-5 (8) mm. long, pubescent; stipules lance- 
subulate, 1-1.5 mm. long, deciduous; flowers axillary ; pedicels slender, pubes- 
cent, 1-2 cm. long ; ovary ellipsoid, pubescent, 5-6 mm. long ; hypanthium deep 
scarlet, purplish at base, 3.5-4.5 cm. long, bulbous at base and 2.5-3 mm. thick. 



Vol. XXV] MUNZ: THE GENUS FUCHSIA  27 

then abruptly narrowed to 1-2 mm. for one-third its length, then rather 
gradually widened until 6-7 (9) mm. wide at summit, more or less pilose 
without, pilose within the lower half or almost throughout ; sepals deep scarlet, 
more or less divergent to almost spreading, 10-16 mm. long, 4—5 mm. wide, 
acuminate, with subulate tips about 1 mm. long, pilose without, glabrous 
within; petals scarlet, sometimes drying purplish, rhombic-ovate to -obovate, 
acute to obtuse or almost rounded at apex, from about as long as to exceeding 
sepals, 10-16 mm. long, 6-12 mm. wide ; stamens 10-12 and 7-8 mm. long ; 
anthers 1.5-2 mm. long ; style pilose within much of the hypanthium, about as 
long as sepals ; stigma scarlet, about 2 mm. thick ; fruit apparently ellipsoid. 

Type locality, ascent to Paramo Guanacas, near Popoyan, Cauca, Colombia. 
Extending from Calda, Colombia into Ecuador. Kepresentative material, 
COLOMBIA : Caldas : Paramo del Quindio, 3800-4000 m., Pennell and Hazen 
10058 (GH, NY). Antioquia : Paramo de Ruiz, 2800-3300 m., LeJimann 3067 
(US. Cauca : Paramo de Buena Vista, Huila group, central Cordillera, 3100 
m., Pittier 1188 (US) ; ad Paramo Guanacas, near Popoyan, Hartweg 992, 
type no. canescens at Geneva (photo F, POM, leaf F) ; in sylvis Purace, 3100 
m., ad Paramo de Guanacas, near Popoyan, Hartweg 993, type no. corollaia 
at Berlin (Photo F, POM, leaves F). Nariiio : Volcan de Galera above Pasto, 
2600-2800 m., Lehmann 5615 5 (F) ; Altos Caballos, 3000 m., Triana 3808 
(US). ECUADOR: Carchi: Nudo de Boliche, Voladero, 4000 m., Penland 
and Summers 915 (F, POM) ; Canton Tulcan, road Tulcan to Pun, 3200 m., 
Mexia 7584 (POM, US) ; Paramo del Azufral, east of Angel, Mexia 7512 (NY, 
US). Bolivar : Simiatuz : Hacienda Talahua, 3200 m., Penland and Summers 
623 (F, POM). Tungurahua: Paramo de Minza, Minza Chica, 3800 m., Pew- 
kmd and Summers 337 (F, POM). 

F. canescens comes in a broad-petaled group which seems to me to be related 
to the F. cordifolia group in its broad hypanthium-base and in haA'ing the 
hypanthium quite inflated in its upper third. The species is near F. ayavacensis 
but differs in the smaller thicker leaf -blades, shorted pedicels, the petals nearly 
as long as or longer than the sepals, the latter being shorter and more spread- 
ing than in ayavacensis. There is some question as to w^hether my treatment of 
canescens is too inclusive ; Penland and Summers 623, andPJ'5 and Mexia 7512 
having the petals rather round and shorter than the sepals ; Hartweg 992, 
Pittier 1382, and Penland and Summers 623 having the upper surface of the 
leaves finely strigose. But the whole range of variation seems such as not to 
make possible a taxonomic differentiation with the amount of material avail- 
able. The leaves from the type collections of canescens and coroUata are quite 
alike. 

(15) Fuchsia rivularis Macbride 

Fuchsia rivularis Macbride, Candollea, 8:24, 1940; Field Mus. Nat. Hist., Bot. Ser., 13, 
part 4:562, 1941. 

Branchlets puberulent; leaves ternate, ovate-elliptic, abruptly acuminate, 
entire, glabrous above, loosely pilese beneath, the blades 5-6.5 cm. long, 2.5-3 



28 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

cm. wide; petioles lacking or nearly so; flowers solitary in axils of somewhat 
reduced upper leaves; pedicels 1 cm. long; hypanthium 4.5 cm. long, sparsely 
puberulent without, pilose within lower portion, gradually ampliate ; sepals 
narrowly lanceolate, long-acuminate, 2 cm. long; petals oblong, about as long 
as sepals; stamens shorter. 

Based on a Mathews collection from Chachapoyas, Amazonas, Peru and said 
to be near F. ayavacensis or F. hirtella, but with narrower petals and more 
sessile and wider leaves. I have seen no material nor photographs, but except 
for the larger leaves and longer petioles it seems much like F. canescens. 

(16) Fuchsia loxensisH.B.K. 

(Plate 2, fig. 14) 

Fuchsia loxemis H.B.K., Nov. Gen. et Sp., 6:106, pi. 536, 1823. 

F. umbrosa Bentii., PI. Ilartw., 176, 1845 ; I have seen a photograph of the type. 

F. apiculata Johnson, Contr. Gray Herb., 75:34, 1925. Type seen. 

Shrub, slender, stiff and bushy or more or less scandent or leaning, com- 
monly up to 2 or 3 m. tall, the younger twigs slender, 1-3 mm. thick, purplish 
red, loosely appressed-puberulent, with whitish or yellowish hairs ; leaves op- 
posite to ternate, sometimes quaternate, firm, elliptic to elliptic-oblong, 
rounded to acute at base, mostly acute at apex, subentire to remotely denticu- 
late, shining, deep green and minutely appressed-puberulent to glabrescent 
above, paler and puberulent beneath, also more or less pilose beneath espe- 
cially along or near the midrib, principal lateral veins often reddish beneath, 
prominent, about 10 on each side of the midrib, the submarginal vein present ; 
leaf -blades 2.5-6 (7) cm. long, 1.2-2 (3) cm. wide ; petioles slender, strigulose, 
1-3 cm. long; stipules subulate-deltoid, 1-2 mm. long, deciduous: flowers 
solitary in axils ; pedicels slender, strigulose, and sometimes pilose, 1-2.5 cm. 
long ; ovary ellipsoid, strigulose, 3-5 mm. long ; hypanthium deep red, 20-25 
(30) mm. long, the base 2-3 mm. thick, then very slightly narrowed, then 
gradually widened until 5-6 mm. wide at the mouth, shining and somewhat 
strigulose without, densely white-pubescent within the lower half; sepals 
scarlet, lance-oblong, 8-10 mm. long, 3.5-4 mm. wide, divergent, puberulent 
to sparsely pilose, the subulate tips about 1 mm. long ; petals ovate, dull red, 
6-9 mm. long, 5-6 mm. wide, obtuse ; stamens 7-8 and 5-6 mm. long, anthers 
2 mm. long; style pubescent in lower part, red, exceeding sepals by 3.5 mm.; 
stigma 2 mm. wide ; fruit ellipsoid, 8-15 mm. long, 5-6 mm. thick. 

Type locality, near Loja (Loxa), Ecuador. The species is apparently con- 
fined to the highlands of Ecuador, growing at from 2000-3400 meters. Repre- 
sentative material seen, ECUADOR: without definite locality: Eastern Cor- 
dillera. Rimhach 6 (GH, NY, US) ; in Andibus Ecuadorensibus, Spruce 5203 
(GH, NY). Pinchincha: Hacienda de Pinantura near Quito, Hartiveg 983, 
type no. umbrosa at Geneva (photo F, POM); near Tambillo, Penland and 
Summers 959 (P, POM) ; Qiiitig-Tesalia (iMaehachi), Firmin 666 (F, GH, 
US). Leon: Cotopaxi : road Pilaloa to Macuchi, Hmiglit 2955 (POM, US). 



Vol. XXV J MUNZ : THE GENUS FUCHSIA 29 

Tunguraliua : west of Ambato, Heinrichs 828 (NY) ; south of Baiioa, Penland 
and Summers 93 (F, POM). Bolivar: Simiatug: Hacienda Talahua, Penland 
and Summers 520 (F, POM) . Azuay and Caiiar : between Cuenca and Huigra, 
Hitchcock 21667, type no. apiculata (GH, NY, US). Loja : mts. near Loxa, 
Hartweg 733 (NY) ; F. loxensis, Humboldt in "Willdenow Herbarium at 
Berlin (photo, F, POM). 

F. loxensis seems near to F. canescens in its leaves, broad petals, etc., but 
has shorter hypanthia and somewhat narrower petals; Spruce 5203, above 
cited, is quite intermediate. It is near F. decussata in some ways, but has 
longer hypanthia, less serrate leaves and less evident stamens. 

(17) Fuchsia pallescens Diels 

Fuchsia pallescens Diels, Notizbl. d. bot. Gart. u. Mus. zu Berlin-Dahlem, 14:34, 1938. 

Medium sized shrub, younger branches subglabrous ; leaves opposite or 
ternate, membranaceous, subovate, subacuminate at both ends, callous- den- 
ticulate, almost glabrous ; blades 6-7 cm. long, 3.5-4 cm. wide ; petioles 2-3.5 
cm. long; flowers axillary, but arranged subcorymbosely near tips of branches ; 
pedicels 8-15 mm. long ; ovary attenuate at apex ; hypanthium pale carmine, 

2 cm. long, nodose at base, then constricted, then gradually widened until 4 
mm. wide at mouth, glabrous without; sepals white, lanceolate, very acute, 
10-12 mm. long, 3 mm. wide at base; petals purplish, 6-7 mm. long; stjde with 
globose stigma. 

Type locality, ECUADOR: Prov. Tunguragua : east of Patate, montane 
forest above Leito at 2750 meters. I have seen no material, but the plant 
would seem to be near F. loxensis and distinct from it in being more glabrous, 
having wider more plainly toothed leaves, and in its white sepals. 

(18) Fuchsia TownsendiiJohnston 

(Plate 3, fig. 15) 
Fuchsia Townsendii Johnston, Contr. Gray Herb., 75:33, 1925. 

Shrub, the young twigs purplish, densely spreading-pubescent ; leaves 
crowded, ternate, elliptic-oblanceolate, membranaceous, acute at base, acum- 
inate at apex, remotely denticulate, green and sparsely and finely pubescent 
above, paler and densely canescent-villous beneath, the principal lateral veins 
10-12 on each side of midrib ; blades 6-13 cm. long, 2-3 cm. wide ; petioles 
pubescent, 5-20 mm. long; stipules subulate, 1-2 mm. long, deciduous ; flowers 
few, axillary; pedicels slender, pubescent, 1.5-2.5 cm. long, bulbous at base 
where 2 mm. thick, then narrowed for one-third its length to 1.5 mm., then 
gradually widened to 5-6 mm. at mouth, densely pubescent without and 
within the lower half; sepals 10-12 mm. long, ascending, lanceolate, acute, 
4 mm. wide, pubescent on back, glabrous on upper or inner surface ; petals 
ovate, 9-10 mm. long, 8 mm. wide, obtuse; stamens 13 and 9 mm. long ; anthers 

3 mm. long ;- style pubescent in lower part, slender, not exceeding sepals ; 
berry hairy, narrowly ellipsoid, over 1 cm. long. 

Type locality, Sabiango Hill, Ecuador, tj^pe collected by C. H. Townsend, 



30 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

number A93 (type at US, fragment GH). The type collection is the only 
material I have seen.; it is distinct in its large thin hairy leaves, long- narrow 
hypanthium and short sepals and petals. F. Schcrffiana Andre (Rev. Ilortic, 
1888, 233, 268) may be an earlier name for this species, but since I have not 
seen certain material I am not sure as to width of petals. 

(19) Fuchsia platypetala Johnston 

(Plate3, fig. 16) 
Fuchsia platypetala Johnston, Journ. Arnold Arbor., 20:241, 1939. 

Shrub 2-3 m. tall, the younger branches 2-4 mm. thick, terete, somewhat 
reddish, pubescent; leaves opposite or ternate, elliptic-lanceolate, membrana- 
ceous, acute at both ends, serrulate, pilose-pubescent on both surfaces espe- 
cially when young, principal lateral veins 8-11 on each side of midrib, with 
submarginal vein present ; blades 2.5-7.5 cm. long, 1-2.3 cm. wide ; petioles 
slender, 4-15 mm. long, pilose; stipules subulate-deltoid, ca. 1 mm. long; 
flowers in upper axils; pedicels slender, 2-3.5 cm. long, somewhat pilose; 
ovary ellipsoid, 8-10 mm. long, pilose; hj^Danthium red, 4.5-5 cm. long, 
bulbous at base where 3-4 mm. thick, then 2-3 mm. wide for one-third its 
length, then gradually widened until 8-9 mm. wide at mouth, sparsely pilose 
externally, densely so within for two-thirds its length ; sepals lanceolate, 
crimson, 15-20 mm. long, 3—4 mm. wide, apiculate for 2-3 mm., sparsely pilose 
without, glabrous within ; petals crimson but with central white oblong blotch, 
obovate, rounded or broadly obtuse, 14-16 mm. long, 10-12 mm. wide ; stamens 
16-17 and 12-13 mm. long; anthers 3 mm. long; style pilose through most of 
its length, reddish ; stigma slightly lobed, 3 mm. thick. 

Type locality, Chincheros, Apurimac, Peru. Material seen, PERU: Dept. 
Apurimac : Chincheros, at 2930 m.. West 3705 (GH, type). Cuzco, cultivado 
en los jardines, Herrera 1514 (GH). The plant would seem in its wide-based 
hypanthium to be in the denticulata, austromontana, leptopoda group but 
differs in its hairy style, thinner leaves, wider petals, and whitish blotch on 
each petal. 

(20) Fuchsia macrostigma P>entham 
Fuchsia macrostigma Bentham, PI. Hartw., 129. Dec, 1844. 

Erect shrub, 0.5-1.5 m. tall, apparently rather openly branched, the young 
twigs green to reddish, herbaceous, terete or angled, 1.5-3 mm. thick, glabrous 
to pubescent ; leaves opposite, membranaceous, broadly to narrowly elliptic, 
acute to obtuse at base, abruptly acute to subacuminate at apex, obscurely 
denticulate, ciliate, dark .green and subglabrous to puberulent above, lighter 
and subglabrous to puberulent beneath, purplish on veins, principal lateral 
veins 12-16 on eacli side of midrib, submarginal vein developed ; blades 8-18 
cm. long, 4-7 (9) cm. wide; petioles 1-3 cm. long; stipules deltoid, 2-3 mm. 
long, more or less persistent ; flowers solitary in upper axils ; pedicels coarse. 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 31 

ascending, 1-2 cm. long; ovary linear, 6-10 mm. long, 2-3 mm. thick; hypan- 
tliiuni tubular, reddish purple, especially on lower half, 6-7 cm. long, 2-3.5 
mm. thick at base, then very slightly narrowed for one-third its length and 
gradually widened to 6-8 mm. at mouth, puberulent to pilose without, gla- 
brous within; sepals lance-oblong, purplish red or lighter, spreading, 16-23 
mm. long, 7-8 mm. wide, subulate for about 2 mm. at tip, pilose to puberulent 
without, glabrous to pilose within ; petals orbicular-obovate, 12-17 mm. long, 
somewhat wider, cerise to crimson, apically rounded ; stamens 12-13 and 
9-10 mm. long; anthers light, about 3 mm. long; style stout, sparsely pubes- 
cent to glabrous, extending about to end of sepals ; stigma subglobose, 4-lobed, 
green, about 4 mm. wade ; fruit elongate, more than 1 cm. long. 

Characterized by its large leaves, purplish color on stems, veins, hypan- 
thium, etc., almost glabrous condition, wide petals and long cylindrical 
hypanthium, this species seems near F. platypetala, rividaris, etc. 

Key to Varieties of Fuchsia macrostigma 

A. Leaves subglabrous except on veins ; hypanthium more or less pilose. Provinces CarcM 

to Pichincha, Ecuador 20a. var. longiflora. 

AA. Leaves minutely puberulent; hypanthium densely close-pubescent. Provinces Chim- 
borazo to El Oro, Ecuador 20b. var. typica. 

(20a) Fuchsia macrostigma Benth. var. longiflora (Benth.) Munz, new comb. 

F. longiflora Bentham, PI. Hartw., 177, 1845. 

F. spectabilis Hook., Bot. Mag., 74:pl. 4375, 1848. 

Leaves subglabrous except on veins ; hypanthium more or less pilose. 

Type locality, Andes of Quito, Ecuador, toward Nanegal. Material seen, 
ECUADOR : Carchi : Olivos, 3200-3500 m., Mexia 7462 (US) . Bolivar : Simi- 
atuz : Hacienda Talahua, 2300 m., Penland and Summers 657 (F, POM), 614 
(POM). Pichincha: Quitensian Andes, Couthouy in 1855 (GH) ; Montis 
Pichincha, Jameson in 1856 (GH, NY) . I have not seen the type. 

(20b) Fuchsia macrostigma Benth. var. typica Munz, new name 

(Plate 3, fig. 17) 

F. macrostigma Bentham, PI. Hartw., 129, 1844. 

F. speciahilis Hook., var. puhens Johnston, Contr. Gray Herb., 75 :34, 1925. 

Leaves minutely puberulent ; hypanthium densely close-pubescent. 

Type locality, "In montibus Pacha," southern Ecuador. Benth., I.e., 113 
says for nos. 708-869 : "in montibus circa Loxa" ; since this species comes 
between nos. 733 and 734, one would infer that it came from that region. I 
have not seen the type number, but the description fits very well the following 
plants : ECUADOR : Chimborazo : Huigra : Hacienda de Licay, /. N. and G. 
Rose 22479, type no. of pubens (GH, NY, US). Oro : between La Chorita and 
Portovelo, ffifc/icocA^ 5:Z:Z^8 (GH, NY, US). 



32 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

(21) Fuchsia ayavacensis H.B.K. 
(Plate 3, fig. 18) 
Fuchsia ayavacensis H.B.K., Nov. Gen. et Sp., 6:107, 1823. 
F. ampliata Bentham, PI. Hartw., 178, 1845. 
F. Hitchcockii Johnston, Contr. Gray Herb., 75: 33, 1925. 

Branching, sometimes scandent shrub up to 3 m. high, younger branches 
hirtellous-pilose, more or less purplish, terete, 1.5-3 mm. thick; leaves ternate 
or quaternate, rather crowded, membranaceous, elliptic-lanceolate to -oblance- 
olate, or -ovate, acute to obtuse at base, acute to acuminate at apex, remotely 
denticulate, principal veins purplish, laterals 6-10 on each side of midrib, 
the submarginal present ; leaf-blades pilose or pubescent and dark green 
above, paler and pilose beneath especially along veins, 4—9 cm. long, 1.5-3 cm. 
"vvide ; petioles pilose, slender, mostly 5-20 mm. long ; stipules subulate-lance- 
olate, 1.5-2 mm., long, deciduous; flowers in upper axils; pedicels slender, 
pubescent, 2.5-4 (5) cm. long; ovary oblong-fusiform, hairy, 6-7 mm. long; 
hypanthium deep red, 3.5-5 (6) cm. long, bulbous at base and 3.5-5 mm. wide, 
then constricted for 1-2 cm. and 2-3 mm. wide, then quickly widened until 
7-11 mm. wide just below the slightly constricted apex, sparsely pilose with- 
out, densely so within most of its length ; sepals acuminate, subulate for 1-2 
mm. at apex, somewhat pilose on back, glabrous above; petals scarlet, rounded- 
obovate, 10-13 (15) mm. long, about as wide; stamens 13-17 and 10-14 mm. 
long ; anthers 3 mm. long ; style densely pilose throughout its portion within 
the hypanthium, sparsely so on base of exposed portion, about equalling the 
sepals; stigma 2-2.5 mm. thick; fruit apparently ellipsoid. 

Type locality, Ayavaca (Ayabaca), Piura, Peru. There is some question 
as to whether I have seen the original material ; number 3115 of Humboldt 
in the Willdenow Herbarium at Berlin labeled Fuchsia is interpreted by 
Lewin as ayavacensis. A photograph was distributed by the Field Museum as 
F. hirta, which name has never been published. No. 3115 in the Bonpland Her- 
barium at Paris is labeled "F. ayavacensis HBK, F. ampliaia Benth. !" as from 
Kucu Pichincha. It agrees well with the description, as does tj^pe material of 
F. ampliata. In the type of F. Hitchcockii, the petals are only half the sepals, 
the hypanthium is rather long, but both these conditions are matched by some 
of the Jameson collections. The reflexed-spreading sepals and round petals 
shorter than the sepals, the pilose condition of the largish leaves characterize 
F. ayavacensis. Representative material. ECUADOR : without locality, Jame- 
son (US). Pichincha: Canton Quito: road from Cotocollao, near Nono, 2987 
m., Mexia 7661 (POM, US) ; Volcan de Pichincha, 3500-4000 m., Mexia 7654 
(POM, US) ; La Chorera, above Quito, Balls 7234 (US) ; Vi(t\\mc\m, James on 
73 (NY) ; sive Andium Quitensium, Jameson 42 (GH), 30 (NY) ; Andes of 
Quito, Jameson (US) ; in declivitate montis Pichincha, 3000 m., Harttveg 988, 
type no. ampliata at Berlin (photo F, PO^M) ; Rucu Pichincha, Bonpland 
3115, Herb. Mus. Paris (F), Humholdt 3115 at Berlin (photo F, POM). 
Azuay : between Oiia and Cuenca, Hitchcock 21603, type no. Hitchcockii (GH, 
NY, US). 



Vol. XXV] MUNZ : THE GENVS FUCHSIA 33 

(22) Fuchsia Pringsheimii Urban 

(Plate 4, fig. 19) 
Fuchsia Pringsheimii Urban, Symb. Antill., 1:375, 1898-1900. 

Shrub 1-2 m. tall, openly branched, the ultimate branchlets purplish, 1-2 
mm. thick, fine-pubescent ; leaves opposite, fairly near each other, not much 
reduced upward, subcoriaceous, strongly bicolored, ovate-elliptic, to rhom- 
boid-elliptic, acute at base, acute to acuminate at apex, somewhat remotely 
serrulate, in maturity mostly subglabrous except on veins of both surfaces, 
principal lateral veins 3-4 on each side of midrib ; petioles 4-7 mm. long ; 
blades 1.5-2.5 cm. long, 0.5-1 cm. wide ; flowers in axils of upper leaves ; pedi- 
cels subglabrous, 2-5 cm. long; hypanthium funnelform, 23-26 mm. long, red, 
glabrous without and within, constricted above ovary, then with tubular 
portion about 2 mm. wide, 10-15 mm. long, the upper expanded portion 10-15 
mm. long, 8-12 mm. wide ; sepals red, lanceolate, acuminate, glabrous, 18-22  
mm. long, 7-8 mm. wide at base ; petals red, equalling or slightly exceeding 
sepals, obovate ; episepalous stamens almost as long as petals, anthers 3 mm. 
long; epipetalous slightly shorter, the anthers 2.5 mm. long; style glabrous, 
equalling petals or up to 5 mm. longer ; stigma clavate, slightly 4-lobed, 2 mm. 
long ; berry ellipsoid, 10-14 mm. long, half as thick. 

Type locality, "Valle nuevo," Santo Domingo, at 2100 m., the type Eggers 
2159. The species differs from F. triplxyUa by having the flowers in axils of 
reduced leaves not in an inflorescence, leaves more glabrous, and by sepals 
and petals being longer in relation to the hypanthium. It and F. iripliylla and 
ayavacensis seem to be related in their ampliate hypanthium and wide petals. 
I have seen the following specimens of Pringsheimii : HAITI : Massif de la 
Pelle, Petionville, Morne La Visite, at 2100 m., Ekman H 1450 (US) . SANTO 
DOMINGO : prope Constanza in Valle nuevo, 2300 m., von Tuercklieim 3151 
(F, GH, NY, US) . A collection from Massif de la Pelle, Marigot, near Jardins 
Bois-Pin, Haiti, Ekman H 1260 (US) has leaves of triphyUa and flowers of 
Pringsheimii. 

(23) Fuchsia triphylla L. 

(Plate 4, fig. 20) 
Fuchsia triphylla L., Sp. PL, 1191, 1753. 

Apparently semishrubby, 3-5 dm. tall; stems 1 to few, simple or few- 
branched, the yoimger ones reddish, densely pubescent, 2-A mm. thick; leaves 
ternate or quaternate or opposite, the lower ones remote, middle and upper 
crowded, coriaceous, lanceolate to lance-ovate, a'cute to acuminate at both 
ends, subentire to serrulate, often with inrolled margin, bicolored, often red- 
dish on veins, fine-pubescent on both surfaces; petioles pubescent, 3-10 (20) 
mm. long; main blades 3-8 (10) cm. long, 1-3 (4.5) cm. wide; uppermost 
reduced to leafy lanceolate or lance-ovate bracts; flowers several, pendulous, 
in dense terminal racemes; pedicels 15-20 (30) mm. long, rather slender, 
pubescent ; hypanthium slightly contracted then expanded, funnelform, red, 



34 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

pubescent without, glabrous within, the tubular part 1.5-2 mm. wide, 8-16 
mm. long, the expanded part 4-7 mm. wide, 15-20 mm. long ; sepals red, lance- 
olate, erect, acuminate, 8-13 mm. long, 4-5 mm. wide at base, puberulent; 
petals red at apex, apparently lighter toward base, ovate-oblong, 8-10 mm. 
long ; stamens as long as or two-thirds as long as petals ; anthers 2-2.5 mm. 
long; stj'le glabrous, scarcely equal to sepals; stigma conic, not much lobed, 
1.5 mm. long ; berry subglobose, at least 1 cm. long. 

Tj-pe locality, "Habitat in America" according to Linnaeus w^ho refers to 
Plumier, Gen., 14, 1703, where the genus Fuchsia was founded with a rude 
drawing (cf. PI. 16, figs. 92-95) Linnaeus basing F. triphylla on Plumier. 
Lamarck, Encyc. 2 :565, 1786, said that Plumier had found the species in the 
Island of Santo Domingo, in uncultivated places, in going from the region 
"de la Bande du Sud a celui qu'on nomme le grand Cul-de-sac." Linnaeus 
placed the genus under Tetrandria Monogynia, but in the Gen. PI. ed. 2, 53, 
1764, observed: "Plumier in Generibus stamina quator, in Historia vere 
saepius octo pingit." Presumably then, the species should have been described 
as having eight stamens and coming from Santo Domingo. Of the specimens 
listed below, there seem to be two forms : one with leaves heavy, lanceolate, 
1-2 cm. wide ; the other less thick, 2-4.5 cm. wide. In the first may be cited : 
HAITI : vicinity of Mission, Fonds Varettes, at 1000 m., Leonard 3681 (NY, 
US), 3701 (NY, US). Belonging to the second group are : HAITI : Massif de 
la Selle, Petionville, at 1600 m., Ekman H 1170 (US). SANTO DOMINGO : 
Prov. de la Vega, ad ripam fontis Yaque at 1200 m.. Padre Miguel Fuertes 
1720 (NY) ; prope Jarabacoa, 700 m., Fuertes 1619 (NY) ; Constanza, Ablot 
in 1919 (US), von Tuerckheim 2956 (F, GH, NY, US), 554i (NY). Perhaps 
more material may show that these forms should be separated taxonomically. 

(24) Fuchsia petiolaris H.B.K. 

Fuchsia petiolaris H.B.K., Nov. Gen, et Sp., 6:104, 1823. 

Erect bushy shrubs or somewhat scandent, up to 2 m. tall ; younger twigs 
reddish, but covered with a fine ashy appressed puberulence, terete, 1-3 mm. 
thick ; leaves elliptical to elliptic-ovate, rather dense, membranaceous, touched 
with purple, obtuse to acute at both ends, remotely serrulate, with 5-7 main 
veins on each side of midrib and with submarginal vein, minutely puberulent 
on both surfaces, somewhat paler on lower surface than upper; blades mostly 
1.5-5 (8) cm. long, 1-2.5 cm. wide; petioles slender, 5-15 mm. long; stipules 
subulate, 1-1.5 mm. long, fairly persistent ; flowers in uppermost axils, some- 
times almost racemose; pedicels minutely strigulose, slender, 1.5-2.5 cm. long ; 
ovary strigulose-puberulent, 3-4 mm. long; hypanthium red, 3-3.5 mm. wide 
at base, then 2-3 mm. wide for length of 1 cm., then rather abruptly ampliate 
until 8-11 mm. wide just below the slightly constricted summit, 3.5-5 cm. 
long, strigulose-puberulent without, somewhat pilose within lower portion ; 
sepals deltoid-lanceolate, 16-18 mm. long, 6-7 mm. wide, acuminate, more or 
less subulate for terminal 1-2 mm., red, finely strigulose-puberulent on backs, 



Vol. XXV] " MUNZ: THE GENUS FUCHSIA 35 

glabrous on upper surface; petals deep red, lance-obloug, 15-17 mm. long, 
4-5 (8) mm. wide, acute, frequently strigulose-puberulent on backs; stamens 
12 and 8 mm. long ; anthers 2.5-3 mm. long; style sparsely pilose in lower por- 
tion, equalling sepals ; stigma 2 mm. wide; berry almost globose when pressed, 
8-10 mm. long. 

This and the species that follow differ from the three previous ones by their 
narrower petals, and resemble them in the long, more or less ampliate hypan- 
thium and the axillary or subterminal flowers. 

Key to Varieties of Fuchsia petiolaris 

A. Leaf -blades 2-3 times as long as wide; petals 4-5 mm. wide 24a. var. typica. 

AA. Leaf -blades 4-5 times as long as wide; petals 7-8 mm. wide 24b. var. bolivarensis, 

(24a) Fuchsia petiolaris H. B. K. var. typica Munz, new name 

(Plate4, fig. 21) 

F. petiolaris H. B. K., Nov. Gen. et Sp., 6:104, 1823. 

F. quinduensis H. B. K., I.e., 105 ; I have seen a portion of the type collection. 
F. curviflora Benth., PI. Hartweg., 177, 1845 ; type from near Bogota, and not seen by me. 
Mutis 1365 compared with type by Johnston seems to agree thoroughly and was seen. 

Leaf -blades, 1.5-5 cm. long, one half to one third as wide ; petals 4r-5 mm. 
wide. 

Type locality, Santa Fe de Bogota, Cundimarca, Colombia. Eepresentative 
material, all from COLOMBIA: without locality, Mutis 1365 (US), 1366 
(US), 1360 (US). Cundimarca: Bogota, Dawe 93 (US), Ariste-Joseph 56 
(US) ; Paramo de Cruz Verde, near Bogota, Pennell 2056 (NY) ; Santa Fe 
de Bogota, Humboldt, Humboldt Herb, at Berlin (photo F, POM) and leaf 
from Bonpland Herb, at Paris (F) ; Rio San Cristobal, near Bogota, Pennell 
2379 (GH, NY, US). Caldas: sw. of Ruiz, Termales, 3400 m., Cuatrecasas 
9218-A (US) ; Quindio La Linea, Dryander 1939 (US). Caldas or Tolima? : 
Quindiu, Humboldt, Willdenow Herb, at Berlin, type no. quinduensis (photo 
F, POM), fragment from Bonpland Herb, at Paris (F). Tolima : Volcancito, 
Holton 896 (GH, NY) ; Ibague, at 1300 m., Andre K 817 (F, GH,NY) ;Rosa- 
lito near Paramo de Ruiz, Pennell 3131 (F, GH, NY, US) ; along divide near 
Quindio Highway, 3300-3500 m. Killip and Varela 34600 (POM, US). Comi- 
saria del Putumayo : Corregimiento El Encano, Laguna de la "Cocha," Garcia 
Barriga 7836 (US) ; Paramo El Tabano, 3300 m., Garcia Barriga 7843 (US) ; 
Paramo de Faliano, Garcia B. no. 4560 (US). 

(24b) Fuchsia petiolaris H. B. K. var. bolivarensis Munz, new var. 

(Plate 4, fig. 22) 

Leaf -blades lanceolate, 5-7.5 cm. long, 12-14 mm. wide ; petals obovate, 7-8 
mm. wide. (Folia lanceolata, 5-7.5 cm. longa, 1.2-1.4 cm. lata; petalis obovatis, 
7-8 mm. latis). 

Type : Colombia : Bolivar : below Paramo de Chaquiro, Cordilerra Occi- 
dental, 2800-3100 m., 11-24-1918, F. W. Pennell 4324 (NY). Like var. tijpica 
in puberulence, flower-size, etc. 



36 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th See. 

(25) Fuchsia Smithii Munz, new species 
(Plate 4, fig. 23) 

Low shrub to Avoody vine up to 2-3 m. tall, with drooping branches, the 
ultimate ones reddish to purplish, more or less ferruginous-villous, terete, 
1.5-2.5 mm. thick, older ones with exfoliating epidermis; leaves mostly ter- 
nate, sometimes opposite, elliptic to elliptic-ovate or -obovate, obtuse to acute 
at both ends, or abruptly acuminate at apex, plainly remote-serrulate, pubes- 
cent and green above or glabrescent except on veins, paler beneath and gen- 
erally pilose or only so along the veins, membranaceous, with 9-11 principal 
veins on each side of midrib and with submarginal vein ; leaf-blades 2.5-8 cm. 
long, 1-2.5 (3.5) cm. wide; petioles slender, villous-pubescent, 5-12 mm. long; 
stipules subulate-lanceolate, 1-1.5 mm. long, deciduous; flowers in upper 
axils; pedicels rather slender, pubescent, 2.5—4 (6) cm. long; ovarj^ oblong, 
pubescent, 6-7 mm. long, 1.5-2 mm. thick; hypanthium tubular, pinkish red 
to purplish red, 4-5.5 cm. long, 3-4 mm. wide at base, then narrowed to 2-3 
mm. for 1.5-2 cm., then rather abruptly widened to 7-9 mm. until near the 
sometimes slightly constricted apex, loosely pubescent to almost glabrous 
without, pilose within most of its length ; sepals divergent, lanceolate, light 
to deep red, acuminate with subulate tips ca. 1 mm. long, subglabrous to some- 
what pilose on both surfaces, 13-16 (20) mm. long, 5-6 mm. wide ; petals dark 
red to lighter, narrowly oblong-ovate, 13-18 mm. long, 5-6 (8) mm. wide, 
acute, often erose-margined, usually pilose on back along median line ; stamens 
11-13 and 9-10 mm. long, red; anthers cream, 2-3 mm. long; style pilose on 
lower portion, equaling petals ; stigma 2.5 mm. thick ; berry dark, broadly 
ellipsoid, 12-15 mm. long, 8-10 mm. wide. 

Frutex erectus vel scandens, ad 2-3 m. ; ramulis ferrugineo-villosis, 1.5-2.5 mm. diametro ; 
foliis tematis vel oppositis, ellipticis, elliptico-ovatis vel -obovatis, obtusis vel acutis vel 
abrupte acuminatis, in superficie siiperiore pubescentibus, inferiore pilosis, membranaceis ; 
lamiiiis 2.5-8 cm. longis, 1-2.5 (3.5) cm. latis; petiolis tenuibus, villoso-pubescentibus, 5-12 
mm. longis; floribus axillarbus; pedicellis pubescentibus, 2.5-4 (6) cm. longis; ovario 
oblongo, pubcscente, 6-7 mm. longo; hypantliio tubuloso, 4-5.5 cm. longo, basi 3-4 mm. 
lato, supra 2-3 mm. lato, versus apicem 7-9 mm. lato, exteriore pubescente ad glabro, 
intcriore piloso ; scpalis divergentibus, lanceolatis, acuminatis, 13-16 (20) mm. longis, 5-6 
mm. latis, subglabris vel pilosis; petalis oblongo-ovatis, 13-18 mm. longis, 5-6 (8) mm. 
latis, acutis, pilosis; staminibus 11-13 et 9-10 mm. longis; antheris 2-3 mm. longis; stylo 
piloso, petalis aequalibus; stigmate 2.5 mm. lato; bacca late ellipsoidea, 12-15 mm. longa, 
8-10 mm. lata. 

Type: Dept. Santander: vicinity of Vetas, Colombia at 3100-3250 m., 
open rocky hillsides, Jan., 1927, E. P. Killip mid Albert C. Smith 17300, at 
Gray Herbarium; istoypes (F, NY, US). The proposed species is near F. 
petiolaris of the western Cordillera of Colombia in having pubescence on the 
back of the petals and in many features, but differs in the coarser longer 
pubescence, larger leaves, longer pedicels, larger fruit and eastern range. 
Other material representative, COLOMBIA : Santander del Norte : west slope 



Vol. XXV] MUNZ: TEE GENUS FUCHSIA 37 

of Paramo del Hatico, Toledo to Pamplona, 2800 m., KilJip and Smith 20714 
(GH, NY, US) ; between Mutiscua and Pamplona, 3400 m., Killip and Smith 
19734 (GH, NY, US) ; east slope of Paramo de Santiirban, toward Mutiscua, 
3600-3900 m., Killip and Smith 19599 (GH, NY, US) ; Pica-Pica Valley above 
Tapata north of Toledo, Killip and Smith 21121 (GH, NY). Santander: 
Paramo de Romeral, 3800-4100 m., Killip and Smith 18577 (GH, NY, US) ; 
Paramo de las Puentes, above La Baja, 3500-3700 m., Killip and Smith 18211 
(GH, NY, US) ; Paramo Rico, Killip and Smith 17771 (GH, NY, US), 17755 
(F, GH, NY, US) ; vicinity of Vetas, Killip and Smith 17361 (GH, NY, US), 
17335 (GH, US), 17386 (GH, US) ; edge of Paramo de Las Vegas, Killip and 
Smith 15734 (F, GH, NY, US), 15610 (GH), between California and Vetas, 
Killip and Smith 17236 (GH, NY, US). Boyaca: between Soata and Cocuy, 
Paramo del Alto Caiiutal, Cuatrecasas 1183 (US) ; Valle de la Uvita, Cuatre- 
casas 1146 (US), 1161 (US). 

(26) Fuchsia Asplundii Macbride 
(Plate 4, fig. 24) 
Fuchsia Asplundii Macbride, Field Mus. Nat. Hist., Bot. Ser., 13, pt. 4, no. 1:548, 1941. 

Shrub ; young twigs apparently dark in color, densely crisp-pubescent, 1-2 
mm. thick ; leaves ternate or quaternate, membranaceous, broadly elliptic- 
lanceolate, acute to obtuse at base, acute to subacuminate at apex, subentire, 
strigose above, more or less pilose beneath especially along veins, principal 
lateral veins 8-12 on each side of midrib, the submarginal vein well developed ; 
blades 3-8 cm. long, 1-2.5 cm. wide ; petioles appressed-pubescent, 1-1.5 cm. 
long on main cauline leaves, 4-5 mm. on reduced upper leaves; stipules lance- 
subulate, 1 mm. long, strigose, deciduous ; flowers solitary in axils of some of 
principal leaves, or appearing subterminal ; pedicels slender, crisp-pubescent, 
1-2.5 cm. long; ovary linear-fusiform, crisp-pubescent, 6-8 mm. long; hypan- 
thium 4-5 cm. long, 2 mm. thick at base, then somewhat narrowed for one- 
third its length, then gradually ampliate until 5 mm. wide at mouth, crisp 
pubescent without, pilose within lower half ; sepals divergent, 12-14 mm. long, 
3 mm. wide, subulate for about 1 mm. at apex, crisp-pubescent on backs, 
glabrous on upper surface ; petals oblong-ovate, 10-12 mm. long, 5-6 mm. 
wide, obtusish, glabrous; stamens 10-11 and 7-8 mm. long; anthers 3 mm. 
long ; style pilose on basal half, exserted beyond sepals 4-6 mm. ; stigma about 
1 mm. thick ; fruit narrow-ellipsoid, 4-angled, at least 12 mm. long, and 3 mm. 
wide. 

Type from PERU : Dept. Piura : Prov. Huancabamba : above Palambla, at 
3000 m., April, 1912, Welerlauer 6054 (F, GH, US). This species suggests 
F. ayavacensis in the almost terminal flowers, but the hypanthium is narrower, 
as are the petals. In its narrow petals and pubescent stems it resembles F. 
petiolaris and F. Smithii, but the pubescence is crisped, the fruit is more 
narrow and the range more southern. 



38 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

(27) Fuchsia Llewelynii IMacbride 

(Plate 5, fig. 25) 
Fuchsia Llewelynii Macbride, Field Mus. Nat. Hist., Bot. Ser., 13, part 4, no. 1 :556, 1941. 

Shrub 1 m. tall; young twigs finely and closely appressed-puberulent ; 
leaves opposite, rigid, broadly oblong-oblaneeolate to -lanceolate, acute at 
both ends, plainly serrulate, minutely appressed-puberulent above and be- 
neath especially on veins, apparently paler beneath, with about 11-14 prin- 
cipal lateral veins on each side of midrib and with submarginal vein ; blades 
4-9 cm. long, 1.5-3 cm. wide ; petioles 4—8 mm. long; stipules subulate, 2 mm. 
long, fairly persistent; flowers apparently racemose; pedicels subfiliform, 
strigulose, 3-5 cm. long; ovary fusiform, strigulose, 5-6 mm. long; hypan- 
thium about 5 cm. long, 2 mm. wide at base, then about 1 mm. wide for 1.5 cm., 
then rather gradually ampliate until 5 mm. wide at summit, strigulose with- 
out, pilose within the lower half ; sepals linear-lanceolate, divergent, 14--17 mm. 
long, 4 mm. wide at base, subulate for 1-2 mm. at apex, strigulose without, 
glabrous within ; petals oblong-lanceolate, 15-16 mm. long, 4^5 mm. wide, 
acuminate; stamens 12-13 and 10-11 mm. long; anthers 2-2.5 mm. long; 
style densely pilose on basal portion, exceeding sepals by few mm. ; stigma 1.5 
mm. thick. 

Tj^e from PERU : Dept. Amazonas : La Ventana Road, Chachapoyas to 
Moyobamba, Llewelyn Williams 7594 (F). The type specimen is-much broken 
and rather unsatisfactory ; apparently from a plant with long slender racemes. 
It is near F. petiolaris in the puberulence and long pedicels, but the leaves 
seem larger and with more veins, and the hypanthium is more narrow. It is 
near F. Asplundii in the narrow hypanthium but has much finer puberulence. 

(28) Fuchsia venusta H.B.K. 

Fuchsia venusta H. B. K., Nov. Gen. et Sp., 6:105, 1823. 

Shrub or shrubby vine, the young twigs hirtellous to puberulent or sub- 
glabrous, often dark red, 2-6 mm, thick ; leaves opposite or ternate, elliptic, 
rounded to acute at base, acute to acuminate at apex, subentire, subcoriaceous, 
not much paler beneath that above, subglabrous to puberulent or pubescent 
on both sides, especially on veins, principal lateral veins mostly 10-15 on each 
side of midrib, with evident submarginal vein; blades 4.5-8 (10.5) cm. long, 
2-3.5 (4) cm. wide ; petioles 6-10 mm. long, puberulent to pubescent ; stipules 
triangular-subulate, barely 1 mm. long, deciduous ; flowers in a terminal 
pendant corymbose raceme or in panicles scarcely 1 dm. long; bracts elliptic- 
lanceolate, 1-2 cm. long ; pedicels 1-3 cm. long, almost filiform, glabrous to 
pubescent ; ovary oblong, 6-7 mm. long, ca. 2 mm. thick ; hypanthium tubular, 
red, subglabrous to glandular-pubescent without, villous within lower portion, 
3-4 em. long, 2.5-3 mm. wide at base, then about 2 mm. wide for one-third its 
length and gradually widened until 6-7 mm. broad at summit; sepals lance- 
olate, acuminate, the connivent tips subulate for 3-4 mm., sepals red, glabrous 



Vol. XXV] MVNZ : THE GENUS FUCHSIA 39 

to pubescent on backs, pubescent within, more or less spreading, 15-17 (20) 
mm. long, 3-4 mm. wide ; petals carmine, oblong-lanceolate, abruptly acute, 
more or less pubescent on backs, 15-18 (20) mm. long; stamens 12-13 (15) 
and 9-10 (12) mm. long, glabrous; anthers 2.5-3 mm. long; style slender, 
densely villous through much of its length, extending to about tips of sepals ; 
fruit oblong, up to 14 mm. long and 7 mm. wide. 

This species is characterized by its short elliptic coriaceous leaves, terminal 
inflorescence, short petioles, long pedicels ; sepals hairy on inner surface as 
well as outer, petals hairy on backs. In the more or less ampliate hypanthium 
and subterminal flowers it agrees with those immediately preceding and 
following in this treatment. There are two forms as to pubescence which I am 
recognizing as varieties. 

Key to Varieties of Fuchsia venusta 
A. Young twigs, petioles, hypanthia pubescent with short spreading hairs. 28a. var. typica. 
AA, Young twigs, petioles, hypanthia glabrous to minutely puberulent with more or less 
appressed hairs 28b. var. huilensis. 

(28a) Fuchsia venusta H. B. K. var. typica Munz, new variety 

(Plate 5, fig. 26) 
F. venusta H. B. K., Nov. Gen. et Sp., 6:105, 1823. 

Hairs on young growth and flowers spreading. 

Type locality: near Guayavalito, Colombia. Ranging in the eastern 
Cordillera of Colombia in Santander and Cundimarca. Representative mate- 
rial, COLOMBIA: Guayavalito (exact locality not determined), Humholdt, 
ex Willdenow Herb. 7301 (photo F, POM), Bonpland Herb, ex Mus. Paris 
(fragment F ) . Without locality, Mutis 1370, 1362, 1367, and 1369 (all at US) ; 
eastern Cordillera, Bro. Ariste-Joseph A893 (US). Santander: Chiquinquira, 
Ariste-Joseph A 842 (US). Cundimarca: Tena, Dawe 23 (US); Bogota, 
Schultze 69 (JJ^), Ariste-Joseph A229 (US) ; Salto de Tequendama, 2500 m., 
Cuatrecasas 48 (US) ; Cordillere de Bogota, 1800-2600, Triana 3814 (NY, 
US) ; ad cataractam Tequedamam, Holt on 894 (GH, NY) ; Paramo de San 
Fortunato et prope pagum Fusagasuga, Hartweg 995 (NY). 

(28b) Fuchsia venusta H. B. K. var. huilensis Munz, new variety 

Young growth, hypanthia, etc., glabrous to minutely appressed puberulent. 
(Ramula, petiola, et hypanthia glabra vel minute striguloso-puberulenta) . 

Type: from Huila, Colombia: Cordillera Oriental, east of Neiva, forest at 
1800-2300 m., Rushy and Pennell 874, (NY) ; isotypes (F, GH, US). Mate- 
rial to be referred here comes from two widely separate regions, VENE- 
ZUELA: Merida: Paramo de Aricagua, 2700 m., Jalin 1029 (GH, US). 
Tachira: Paramo de Angaraveca, 2400 m., J aim 129 (US). COLOMBIA: 
Cundimarca: Dintel (Facatativa-La Vega) 2700-2300 m., Perez A. and 
Cuatrecasas 5291 (US) ; Une, in montibus ad orient. Bogotae, Holton 895 
(NY). Tolima : Libano, 1900-2200 m., Pennell 3197 (GH, NY, US). Huila: 



40 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Cordillera Oriental, east of Neiva, Rushy and Pennell 641 (GH, NY, US) ; 
"Balsillas" on Rio Balsillas, 2000-2300 m.. Rushy and Pennell 918 (NY) ; 
722 (GH, NY, US), 809 (NY, US). 

(29) Fuchsia Jahnii ]Munz, new species 
(Plate 5, fig. 27) 

Apparently a scandent shrub with yoiinger twigs very slender, 1-2 mm. 
thick, drooping, somewhat purplish, puberulent with appressed hair ; leaves 
opposite or ternate, elliptic, rounded to acute at base, short-acuminate at 
apex, remotely denticulate, minutely pubescent on both surfaces, darker green 
above than beneath, membranaceous, with about 10-12 principal veins on 
each side of the midrib and with submarginal vein; blades 3-8 cm. long, 1-3 
cm. wide ; petioles 6-15 mm. long, more or less strigulose ; stipules lance- 
subulate, ca. 2 mm. long, deciduous; flowers few in terminal corymbose ra- 
ceme, sometimes also axillary; bracts 1-2 (3) cm. long; pedicels filiform, 
strigulose, 2-4 cm. long; ovary broadly fusiform, strigose, 5-6 mm. long; 
hypanthium red, tubular, 3.5-4 cm. long, strigulose without, villous within, 
2.5-3 mm. wide at base, then 1.5-2 mm. thick and very gradually widened 
until 7-8 mm. wide toward the mouth ; sepals red, lanceolate, spreading, 18-20 
mm. long, 3.5-4 mm. wide, acuminate (the connivent tips subulate for 3^ mm. 
in bud), strigulose without, glabrous within; petals light red, oblong-oblance- 
olate, 18-20 mm. long, abruptly acute, somewhat strigulose on back toward 
base; stamens 16-18 and 12-14 mm. long, glabrous; anthers 2.5-3 mm. long; 
style slender, retrorse-\'illous, projecting about as far as petals; stigma sub- 
globose, slightly lobed, 2 mm. wide ; berry almost globose, strigulose, about 
12 mm. long, 10-11 mm. wide. 

Frutex scandens; ramulis subpurpureis, strigulosis ; foliis oppositis vel ternatis, ellip- 
ticis, basi rotundatis vel acutis, apice breviter acuminatis, remote denticulatis, minute 
pubescentibus, membranaceis, laminis 3-8 cm. longis, 1-3 cm. latis; petiolis 6-15 mm. 
longis, snbstrigulosis; floribus paucis, subracemosis; bracteis 1-2 (3) cm. longis; pedicellis 
filif ormibus, strigulosis, 2-4 cm. longis ; ovario late f usif ormi, strigoso, 5-6 mm. longo ; 
liypantliio rubro, tubuloso, 3.5-4 cm. longo, externe striguloso, interne villoso, base 2.5-3 
mm. late, inde 1.5—2 mm. lato, inde gradatim ampliato, usque ad apicem 7-8 mm. lato; 
sepalis rubris, lanceolatis, divergentibus, 18-20 mm. longis, 3.5-4 mm. latis, acuminatis, 
strigulosis, cum apicibus subulatis 3-4 mm. longis ; petalis roseis, oblongo-oblanceolatis, 
18-20 mm. longis, abrupte acutis, basi et extrinsecus substrigulosis; staminibus 16-18 et 
12-14 mm. longis; antheris 2.5-3 mm .longis; stylo tenui, retrorsovilloso, petalis aequalibus 
aequescente; stigmate subgloboso, 2 mm. lato; bacca subglobosa, strigulosa, 11-12 mm. 
longa. 

Type: Merida, Venezuela, Paramo de la Sal, 2800 m., Sept. 2, 1921, 
John 506 U. S. Nat. Herb. 1186509 ; isotype Gray Herb. Another collection, 
INIerida: Paramo de Pinango, 2600 m., Jakn 402 (US). Apparently closely 
related to F. venusta and characterized by a few flowers, strigulose condition, 
spreading sepals with long subulate tips, hair on back of petals, and big round 
fruit. 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 41 

(30) Fuchsia Gehrigeri Munz, new species 

(Plate 5, fig. 28) 

Subscandent shrub 3-5 m. tall, with slender, ultimate, reddish, drooping 
branchlets 1-2 mm. thick, subglabrous or somewhat villous about the nodes 
and toward the inflorescence, somewhat reddish ; leaves opposite or ternate, 
membranaceous, elliptic, acute to subacuminate at both ends, serrulate, green 
and minutely strigulose above, pale beneath and pubescent along the veins, 
principal lateral veins 7-10 on each side of midrib, submarginal vein scarcely 
evident; blades 3-6 cm. long, 1.5-2.5 (3) cm. wide ; petioles slender, 10-18 mm. 
long, subglabrous to pubescent ; stipules subulate, 1 mm. long, deciduous ; 
flowers few, in terminal clusters of about 2 whorls, pendant, dark red ; bracts 
1—1.5 cm. long; pedicels slender, 1.5—2 cm. long, more or less spreading-villous ; 
ovary broadly fusiform, 5 mm. long, densely spreading-villous ; hypanthium 
tubular, 4—5 cm. long, 2 mm. wide at base, then narrower for one-third its 
length, and rather abruptly widened until 7-8 mm. wide at summit, villous 
without and within ; sepals lanceolate, divergent, 14-15 mm. long, 4 mm. wide, 
acuminate (subulate tips connivent and 1 mm. long in bud), pubescent on 
back, glabrous on inner surface ; petals dark red, elliptic-oblong, obtuse-erose, 
ca. 14 mm. long and 6 mm. wide, sparsely villous on back near the base; 
stamens 9-10 and 7-8 mm. long ; anthers 2 mm. long ; style pubescent on lower 
portion, exceeding sepals by 3-5 mm., quite thick toward the somewhat lobed 
stigma (1.5-2 mm. wide) ; fruit subglobose, somewhat villous. 

Frutex subscandens, 3-5 m. altus; ramulis tenuibus, subrubris, 1-2 mm. diametro, sub- 
g]abris vel subvillosis in nodis et ad inflorescentiam ; f oliis oppositis vel ternatis, membran- 
aceis, ellipticis, basi et apice acutis vel subacuminatis, serrulatis, in superficie superiore 
viridibus et minute strigulosis, inf eriore pallidis et in venis pubescentibus ; laminis 3-6 cm. 
longis, 1.5-2.5 (3) cm. latis; petiolis tenuibus, 10-18 mm. longis, subglabris vel pubescent- 
ibus; floribus paucis, in infloreseentia terminali, subrubris; bracteis 1-1.5 cm. longis; 
pedicellis tenuibus, 1.5-2 cm. longis, subvillosis ; ovario late fusif ormi, 5 mm. longo, villoso ; 
hypanthio tubuloso, 4-5 cm. longo, basi 2 mm. lato, inde angustiore, inde abrupte ampliato, 
ad apicem 7-8 mm. lato, externe et interne villoso; sepalis lanceolatis, divergentibus, 14-15 
mm. longis, 4 mm. latis, acuminatis, externe pubescentibus, apicibus subulatis 1 mm. 
longis; petalis elliptico-oblongis, obtuso-erosis, 14 mm, longis, 6 mm. latis, sparse externe 
villosis ; staminibus 9-10 et 7-8 mm. longis ; antheris 2 mm. longis ; stylo pubescenti, quam 
sepala 3-5 mm. longiore; stigmate 1.5-2 mm. lato; fructu subgloboso, subvilloso. 

Type: from Merida: Mucuruba, Venezuela, 2800-3100 m., July 14, 1930, 
W. Gehriger 322, U. S. Nat. Herb. 1515306; isotypes F, GH, NY. This species 
is near F. JaJinii in its leaves, few flowers, round fruit, and petals with hair on 
the back, but the pubescence is longer and more spreading, the pedicels are 
shorter ; sepal-tips shorter ; flowers smaller, and sepals are less spreading. 

(31) Fuchsia simplicicaulis Ruiz and Pa von 

(Plate5, fig. 29) 
Fuchsia simplicicaulis Euiz and Pav6n, F1. Peruv., 3 :89, pi. 322, 1802. 

Shrub, apparently growing up through other shrubs, up to 4.5 m. tall ; 
stems slender, the ultimate branchlets pendant, scarcely 1 mm. thick, glabrous 



42 CALIFORNIA ACADEMY OF SCIENCES [Pboc. 4th Ser. 

except for the piiberulent inflorescence ; leaves in remote whorls of 4, linear- 
lanceolate, coriaceous, glabrous, subentire, obtuse to acute at base, acuminate 
at apex, not strongly bieolorecl, with 20 or more principal lateral veins on 
each side of midrib, and with submarginal vein, 8-15 cm. long, 1.5-2 (2.5) cm. 
wide; petioles 2-4 mm. long; stipules subulate, ca. 1 mm. long, deciduous; 
inflorescence a simple or few-branched hanging puberulent raceme 2-3 dm. 
long, with remote whorls of 4 submembranaceous lance-ovate, acuminate, 
green bracts 2-4.5 cm. long and 6-14 mm. wide ; flowers one in each axil, bright 
red with slightly purplish tinge ; pedicels 4-7 mm. long, 12 mm. in fruit ; ovary 
puberulent, ellipsoid, 5-6 mm. long, 1.5-2 mm. thick; hypanthium tubular, 
4-5.5 cm. long, puberulent without and within, ca. 3 mm. wide at very base, 
then 1.5 mm. for two-fifths its length, then rather gradually ampliate and 
tubular, 5-6 mm. wide at summit; sepals 4, linear-lanceolate, puberulent, 
acuminate, 16-18 mm. long, 4 mm. wide, somewhat divergent; petals red, 
linear -lanceolate, acute, 11-12 mm. long, 2-2.5 mm. wide; stamens ca. 14 and 
10 mm. long; anthers 3 mm. long; style pubescent in lower hypanthium, 
slightly exceeding petals ; fruit oblong, puberulent, at least 8 mm. long. 

Type locality, Muna, Peru. Specimen seen, PERU : Huanuco : Muiia, at 
2200 m.. May 23-June 4, 1923, J. F. Macbride 4014 (F, GH, US) ; photo of 
Ruiz and Pavon specimen from Muiia at Madrid (F, POM). This species 
stands apart from all others in its almost linear leaves and bracts in wliorls 
of 4, in its long pendant twigs, and in its puberulence. It agrees with F. petio- 
laris and F. ve^iusta in its subterminal inflorescence, slightly inflated hypan- 
thium and narrow petals. 

(32) Fuchsia confertifolia Fielding and Gardner 

(Plate6, fig. 30) 

Fuchsia confertifolia Fielding and Gardner, Sert. PI., pi. 28, 1844. 

F. dolichantha Krause, Fedde, Eep., 1:172, 1905; based on Weberbauer 4390 from the 

Chachapoyas region in which Mathews collected, confertifolia being based on a Mathews 

type. 

Erect shrub up to 2 m. tall, much branched, the young twigs 1.5-3 mm. 
thick, densely f errugineous-hirsute ; leaves opposite to ternate or quaternate, 
crowded, spreading-reflexed, oblong-ovate, rounded to acute at base, acute at 
apex, entire to sub denticulate, glabrous on both surfaces or sparsely hairy 
beneath on veins, principal lateral veins obscure, 3-4 on each side of midrib ; 
blades 12-16 mm. long, 3-4 (6) mm. wide; petioles 1-2 mm. long; stipules 
dark, glabrous, subulate, 1-1.5 mm. long, persistent; flowers very few, sub- 
terminal, pendant ; bracts about 5 mm. long, 1.5 mm. wide ; pedicels 1-1.5 cm. 
long, up to 2 cm. in fruit, villous; ovary oblong, 4—5 mm. long; hypanthium 
dark red, tubular, 4r-5 cm. long, slightly enlarged at very base and 1.5-2 mm. 
wide, then narrowed for one-third its length, then abruptly widened and sub- 
cylindric until 6-8 mm. wide at and below the summit, sparsely villous with- 
out, more densely so within; sepals linear-lanceolate, divergent, 16-18 mm. 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 43 

long, acuminate, scarcely subulate at tips, sparsely villous without, glabrous 
within ; petals red, elliptic-lanceolate, acuminate, glabrous, 12-13 mm. long, 
3 mm. wide; stamens about 13 and 9 mm. long; anthers 2.5 mm. long; style 
pilose near base, equal to sepals; stigma capitate, slightly lobed, 2 mm. wide; 
fruit ellipsoid, ca. 1 cm. long. 

Type locality, "Peru." Specimens seen, PERU: Dept. Amazonas: between 
Jumbilla and San Carlos, 3000 m., WeherUuer 7153 (F, GH) ; ab Chacha- 
poyas at orientem versus Tambo Ventillas, Weberhauer 4390, type doUchantha 
at Berlin (photo F, POM). 

(33) Fuchsia AspiazuiMacbride 
(Plate 6, fig. 31) 
Fuchsia Aspiazui Macbeide, Field Mus. Nat. Hist., Bot. Ser., 19, part 4, no. 1 -.547, 1941. 

Shrub up to 2 m. tall, the younger branches 4-6 mm. thick, minutely strigu- 
lose under a lens, greenish, terete ; leaves opposite, elliptic-ovate to -obovate, 
subentire, apparently rather tleshy, glabrous on casual inspection, but min- 
utely strigulose under a lens, cuneate at base, rather abruptly acuminate at 
apex, yellowish green and much alike on both surfaces, conspicuously veined, 
the main laterals 18-24 on each side of the midrib, with evident submarginal 
vein; blades 8-20 cm. long, 4-10 cm. wide ; petioles 1-2 (3.5) cm. long ; stipules 
lance-deltoid, 2-3 mm. long, sometimes fused in pairs, deciduous ; inflorescence 
a terminal strigulose pendant raceme with rather numerous flowers ; bracts 
reflexed, lanceolate, 1-2 cm. long; pedicels 8-12 mm. long; ovary linear, 5-8 
mm. long, 1 mm. thick, glabrous to somewhat strigulose ; hypanthium blood- 
red, tubular, 3-5 cm. long, slightly bulbous at base, then about 1 mm. wide 
and very gradually ampliate toward the summit where it is 4-5 mm. wide, 
practically glabrous without, retrorse-pubescent within basal part; sepals 
divergent, red, lanceolate, 12-15 mm. long, 3-4 mm. wide, acute, without free 
tips in bud; petals oblong, obtuse, about 1 cm. long, 3.5 mm. wide; stamens 
about 8 and 6 mm. long; anthers barely 2 mm. long; style quite glabrous, 
extending about to the end of the petals; immature berry linear-oblong, 12 
mm. long, 3.5 mm. wide. 

Based on a collection from valley of the Rio Mixiollo, Prov. Pataz, Dept. 
Libertad, PERU, 2300-2500 m., Weherhauer 7042 (F, GH, US). Character- 
ized by its glabrous condition, large leaves with many veins, short pedicels, and 
glabrous style, it comes in the group of species with large leaves, short terminal 
inflorescence, and narrow long hypanthium. 

(34) Fuchsia tincta Johnston 
(Plate 6, fig. 32) 
Fuchsia tincta Johnston, Journ. Am. Arb., 20:242, 1939. 

Shrub, 1-1.5 m. tall, the young twigs more or less broAvnish red, 1-5 mm. 
thick^, puberulent or finely pubescent with somewhat brownish hairs; leaves 
opposite, membranaceous, elliptic-ovate to -obovate, rounded to obtuse at base, 



44 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

acute to subaeuminate at apex, remotely serrulate, more or less purplish espe- 
cially beneath, finely pubescent on both sides, even somewhat short-pilose on 
veins beneath, the principal lateral veins 16-18 on each side of midrib, the 
submarginal vein present; blades 5-12 (15) cm. long, 3-8 (10) cm. wide; 
petioles 1-4 (6) cm. long; stipules subulate, 1 mm. long, deciduous; flowers 
few to several in open terminal finely pubescent corymbose racemes; bracts 
rather leafy, 1.5-3 cm. long ; pedicels slender, pubescent, 1.5-3 cm. long; ovary 
linear-fusiform, pubescent, 4r-5 mm. long ; hypanthium 20-25 (35) mm. long, 
deep crimson, 1.5-2 mm. wide at base, then narrowed for one-third its length, 
then gradually widened to 3^ mm. at summit, rather sparsely villous with- 
out, villous within lower two-thirds ; sepals erimspn, divergent, 8-10 mm. long, 
3.5 mm. wide at base, subulate for 0.5-1 mm. at apex, villous on backs; petals 
crimson-scarlet, oblong-ovate, 7-8 mm. long, 4-5 mm. wide, erose-rounded to 
obtusely mucronate at apex ; stamens about 8 and 6 mm. long ; anthers 1.5 miu. 
long; style pilose in basal portion, equaling sepals; stigma 1-1.5 mm. wide; 
berry ellipsoid-oblong, 8-12 mm. long, 4.5-6 mm. wide. 

Type locality, PERU : dept. Cuzco, prov. Paucartambo : Rio Tambomayo 
at 1800-2200 m., the type collection being West 7092 (GH) , another collection 
from the same locality: West 7110 (Gil). I have seen also from Cabecera de 
montana de Paucartambo, at 2000 m., Vargas 73/7110 (F, POM) and "Pilla- 
huata," Cerro de Cusilluyoc, 2000-2400 m., Perinell 13956 in large part (F, 
NY, US). The species is near F. ovalis in its wide leaves, but the racemes are 
terminal, the pubescence shorter, the stipules shorter and the hypanthium 
longer. When the hair is not too brown, it suggests F. holiviana, but the sepals 
are not reflexed-spreading. In its long-tubed flowers it resembles F. furfur- 
acea, but the sepals are shorter and the plant is less pilose. 

(35) Fuchsia Mathewsii Macbride 

Fuchsia Mathewsii Macbride, Candollea, 8:24, 1940; Field Mus. Nat. Hist., Bot. Ser., 13, 
pt. 4, no. 1:558, 1941. 

Branches and petioles and lower leaf-surface densely rusty-pilose ; leaves 
ternate, broadly elliptic, abruptly narrowed at base, acute at apex, obscurely 
denticulate to subentire, minutely pilose above, densely so beneath along 
midrib; blades 1 dm. long; petioles 4-5 mm. long; inflorescence a terminal 
short-bracted panicle about 5 cm. long; hypanthium 4 cm. long, glabrate 
without, hirsutulose within lower half; sepals 10 mm. long; petals oblong- 
obovate, almost as long as sepals. 

Based on a collection by Mathews, from Chachapoyas, Amazonas, Peru. 
Said to be near F. pilosa, but differing in its short-petioled leaves, denser 
shorter rusty pubescence and its larger flowers. I have seen no specimens. The 
meager description suggests F. tincta of a more southern distribution and 
with apparently longer petioles and shorter hypanthium. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 45 

(36) Fuchsia Fischeri Macbride 

Fuchsia Fischer i Macbride, Field Mus. Nat. Hist., Bot. Ser., 13, part 4,no. 1 :554, 1941. 

Shrub, suberect, branched, the young twigs reddish, densely hirsutulous ; 
leaves opposite, oblong-lanceolate, subcuneate at base, acute at apex, subentire 
to obscurely callose-denticulate, densely soft-pilose on both sides, veins prom- 
inent beneath ; petioles 2-5 mm. long; stipules subulate, somewhat persistent ; 
inflorescence a terminal leafy -bracted corymb 5-6 cm. long ; pedicels 1.5 cm. 
long; ovary velutinous, 4 mm. long; hypanthium about 4.5 cm. long, swollen 
at base, then constricted, then ampliate until 4-5 mm. wide at apex, somewhat 
villous without and within; sepals lanceolate, acuminate, 10-12 mm. long; 
petals narrowly oblong-lanceolate, short-aciiminate, about 9 mm. long; sta- 
mens shorter than petals; style pubescent in lower portion, more or less 
exserted. 

Type not seen, based on Welerhauer 4097 from Hualgayoc, Chugar, Caja- 
marca, Peru. No specimens available and description meager, but this seems to 
be a distinct entity near F. Mathewsii, perhaps with more narrow leaves. 

(37) Fuchsia Storkii Munz, new species 

(Plate6, fig. 33) 

Tree, or shrub up to 3 m. tall ; twigs reddish, pubescent or pilose with red- 
dish hairs ; leaves ternate, narrowly elliptic to elliptic-oblanceolate, firm, 
obtuse to acute at base, acute at apex, remotely serrulate, light green above, 
pilose to subgiabrous or pubescent on veins above, slightly paler and more 
pilose beneath, principal lateral veins 8-9 on each side of midrib, submarginal 
veins present; leaf -blades 2-7 cm. long, 1.3-2 cm. wide; petioles reddish- 
pilose, 4-8 mm. long; stipules lance-subulate, 1.5-2 mm. long, glabrous, fairly 
persistent ; flowers few, in short terminal drooping glandular-pubescent 
corymbose racemes; bracts lanceolate; pedicels stout, red-pubescent, 2-5 mm. 
long ; ovary ellipsoid, glandular-pubescent to red-pilose, 4-5 mm. long ; hypan- 
thium dark red, about 3 cm. long, somewhat bulbous and 1.5-2 mm. wide at 
base, then somewhat narrowed and then gradually ampliate until 5 mm. wide 
at apex, glandular-villous without, villous within lower half; sepals deep red, 
lanceolate, about 12 mm. long, 4 mm. wide, subulate for 1.5-2 mm. at apex, 
glandular-pubescent ; petals deep red, oblong-lanceolate, 7-9 mm. long, 3 mm. 
wide, sharply acute; stamens about 8 and 6 mm. long; anthers 2 mm. long; 
style pubescent in lower portion, extending about to sepal-tips ; fruit appar- 
ently ellipsoid, at least 1 cm. long. 

Arbor vel frutex; ramulis rubescentibus, pubescentibus vel pilosis, cum pubescentia 
rubida ; f oliis ternatis, anguste ellipticis vel elliptico-oblanceolatis, base obtusis vel acutis, 
apice acutis, remote serrulatis, in venis superficiei superioris pilosis vel pubescentibus vel 
subglabris, inferiore pilosioribus; laminis 2-7 cm. longis, 1.3-2 cm. latis; petiolis rubido- 
pilosis, 4-8 mm. longis; stipulis lanceolato subulatis, 1.5-2 mm. longis, glabris, subpersis- 
tentibus ; floribus paucis in racemis brevibus, terminalibus, glanduloso-pubescentibus ; 
In-acteis lanceolatis; pedicellis rubido-pubescentibus, 2-5 mm. longis; ovario ellipsoideo, 



46 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

4-5 mm. longo ; hypanthio rubido, 3 cm. longo, base 1.5-2 mm. lato, inde angustiore, inde 
gradatim ampliato, apice 5 mm. lato, externe glanduloso-villoso, interne baseque villoso ; 
sepalis rubris, lanceolatis, 12 mm. longis, 4 mm. latis, glanduloso-pubescentibus, apice sub- 
ulatis; petalis rubris, oblongo-lanceolatis, 7-9 mm. longis, 3 mm. latis, acutis; staminibus 
8 et 6 mm. longis; antheris 2 mm. longis; stylo pubescente, sepala aequescente; bacca 
cllipsoidea, ca. 1 cm. longa. 

Type: Cajamarca, prov. Chota, Peru: pass south of Conchan, 2500 m., 
Dec. 7, 1938, Stork and Horton 10073, Field Museum Herb. 1052248. The 
reddish condition of the rather long hairs and the narrow leaves are charac- 
teristic of this proposed species, which seems near to F. FiscJieri, but it has a 
shorter hypanthium and pedicels and red hairs. It is a pleasure to name this 
for Dr. Harvey Stork of Carleton College. 

(38) Fuchsia furfuracea Johnston 

(P],ate 6, fig. 34) 
Fuchsia furfuracea Johnston, Contr. Gray Herb., 75:39, 1925. 

Shrubby, of unknown size, the branchlets dark, sparsely villous-hirsute ; 
leaves opposite, oblong-ovate, obtuse to rounded at base, acuminate at apex, 
denticulate, brownish-pilose on both surfaces, especially on the veins beneath, 
with 11-13 principal lateral veins on each side of the midrib ; blades 6-9 cm. 
long, 2.5-4 (5) cm. wide; petioles 8-20 mm. long, pubescent; stipules dark, 
subulate-lanceolate, 2-3 mm. long; flowers few in terminal racemes; bracts 
pilose, 1-2 cm. long, lanceolate, dentate ; pedicels pubescent-pilose, 2-4 cm. 
long; ovary elliptic, villous, 6-8 mm. long; hypanthium red, tubular, 3.5^.5 
cm. long, slightly enlarged at base, then constricted so as to be scarcely 2 mm. 
wide for about half its length, then rather abruptly widened to 6-7 mm. near 
the cylindrical summit, villous without and within; sepals red, somewhat 
spreading, lanceolate, 15-20 mm. long, 4 mm. wide, villous on backs, long- 
acuminate, with free tips 2 mm. long in bud ; petals purplish red, oblong-ovate, 
abruptly acutish, 9-10 mm. long; longer stamens two-thirds the sepals, 
shorter scarcely one-half the sepals ; anthers 2 mm. long ; style villous, slightly 
exceeding sepals; stigma 1.5 mm. thick, subcapitate, slightly 4-lobed; im- 
mature berry ellipsoid, barely 1 cm. long. 

Type locality, Yungas, La Paz, Bolivia. Material seen, BOLIVIA : La Paz : 
Yungas, Bang 674, type collection (F, GH, NY, US) ; Unduavi, at'3000 m.. 
Rushy 2511 (NY), Characterized by its long stipules and pedicels, brown 
pubescence, long sepal-tips, it seems near to F. Storkii, F. hirtella, etc., in its 
large leaves, narrow hypanthium, short terminal inflorescence. The "F. 
peruviana" of Johnston's text accompanying the original description is un- 
doubtedly a misprint for F. holiviana. 

(39) Fuchsia Munzii Macbride 

(riate7, fig. 35) 
Fuchsia Munzii Macbride, Field Mus. Nat. Hist., Bot. Ser., 13, part 4, no. 1:559, 1941. 

Shrub, the younger branches 1.5-3 mm. thick, very finely puberulent, ap- 
parently somewhat dark ; leaves mostly opposite, sometimes ternate, broadly 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 47 

elliptic-lanceolate to -ovate or -obovate, subentire to inconspicuously denticu- 
late, the principal lateral veins about 20 on each side of midrib, blades sub- 
cuneate at base, acuminate at apes, somewhat bicolored, quite glabrous, 6-15 
cm. long, 2.5-4.5 cm, wide; petioles 4-10 mm. long; stipules subulate-lanceo- 
late, about 1 mm. long, caducous; inflorescence a small terminal raceme or 
panicle, up to 1 dm. long, with rather few flowers, very minutely strigulose 
bracts linear to lanceolate, 5-20 mm. long; pedicels 5-15 mm. long; ovary 
linear, 4—5 mm. long, strigulose; hypanthium red, tubular, 4-5 cm, long, 
slightly enlarged at base, then 1 mm. wide, then very gradually ampliate and 
tubular, 4 mm. wide, minutely and rather faintly puberulent without, pubes- 
cent within ; sepals lanceolate, somewhat puberulent, 12 mm. long, about 3 mm. 
wide, acute, somewhat spreading; petals narrowly oblong, 11-12 mm. long, 
acute ; stamens 9 and 6 mm. long; anthers 2 mm. long ; style pubescent in lower 
portion, extending about to sepal-tips ; stigma subglobose, somewhat 4-lobed, 
2 mm. thick ; berry subglobose, almost 1 cm. long. 

Type collection, PERU: Dept. Junin: Rio Masamerich, 2100-2200 m., 
Weherhauer 6648 (F, GH, US). This species suggests F. corymhiflora from 
which it differs in its more glabrous leaves and rounder fruit. 

(40) Fuchsia hirtella H.B.K. 

(Plate 7, fig. 36) 

Fuchsia hirtella H. B. K., Nov. Gen. et Sp., 6 : 107, 1823. 

F. caracasensis Fielding and Gardner, Sert. PI., pi. 29, 1844; based on Linden 368, sup- 
posed to be from Caracas, Venezuela, in 1842. The plate and the specimen at US indicate 
synonymy with hirtella. 

Fuchsia miniata Planch and Linden, PL des Serres, Ser. I, pi. 8, 1852-1853; based on 
Linden collection from "Nouvelle Grenade." In my key would seem to run to F. Killipii 
on basis of puberulent stems, but cannot be that, since the young growth is not canescent, 
so must be hirtella. I have seen no specimen of Linden, labelled miniata. 

Shrub to low tree, the younger branches apparently mostly reddish, 2-4 
mm. thick, terete, mostly rather densely pubescent with short somewhat ap- 
pressed and interwoven hairs, older branches exfoliating freely; leaves in 
whorls of 3 or 4, lance-oblong to elliptic-ovate, rounded to cuneate at base, 
acute to acuminate at apex, remotely and indistinctly denticulate, strigulose 
and dark green above, more densely pubescent and paler beneath, membrana- 
ceous, with 10-15 principal lateral veins on each side of midrib and rather 
inconspicuous submarginal vein, the blades 6-12 (14) cm. long, 2.5^ (5) cm. 
wide; petioles densely pubescent, 5-20 (35) mm. long; stipules lance-linear, 
1-2 mm. long, deciduous; flowers in terminal pendant corymbose racemes 
which may be grouped in a panicle 1-2 dm. long and equally wide ; bracts 
lance-ovate, mostly 1-2 cm. long, strigulose ; pedicels more loosely pubescent, 
slender, mostly 5-10 mm. long ; ovary oblong-fusiform, strigulose-pubescent, 
5-7 mm. long, 1.5-2 mm. thick; hypanthium tubular, rose-red, 3-3.5 (4,5) 
cm. long, 2.5-3 mm. thick at base, then 1.5-2 mm. for about one-third its 
length and very gradually widened until 6-8 mm. at summit, strigulose with- 



•48 CALIFORXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

out, villous witliiu lower half ; sepals 4, red, somewhat divergent, lanceolate, 
acuminate, subulate for terminal 2 mm., 12-13 mm. long, 2.5-3 mm. wide, 
strigulose on backs; petals red, lance-oblong, glabrous, acutish, 11-14 mm. 
long ; stamens 10-14 and 6-10 mm. long, glabrous; anthers 2-3 mm. long ; style 
red, filiform, almost or quite glabrous, exserted 4—8 mm. beyond sepals; 
stigma subglobose, 4-lobed, ca. 2 mm. thick; berry ellipsoid, puberulent, 14 
mm. long, 6 mm. wide. 

Type locality, in mts. near Fusagasuga, Cundimarca, Colombia. This species 
seems near to F. polyantha and F. holiviana in pubescence, leaves, inflores- 
cence, hypantliium, etc., but distinct in its membranaceous leaves, ascending 
sepals, and long fruit. It ranges rather widely in Colombia as can be seen from 
citation of representative material : COLOMBIA : without locality, Mutis 
1364 (US) ; Bonpland Herharium 1783, probably type material, it is rather 
fragmentary, a condition referred to in the original description (fragment F) . 
Antioquia : Tamesis, near Medellin, Toro 965 (NY) ; San Pedro, Tomds and 
Daniel 1306 (F). Caldas : Pinares, above Salento, 2600-2900 m., Pennell 9195 
(Gil), 9324 (GH, NY, US); Volcancito, Holton 893 (NY); Mediacion- 
Quindio, Andre 2152 (NY) ; "Laguneta" to "Magana," old Quindio trail, 
3000-3300 m., Pennell 9418 (GH, NY, US) ; Salento to "Laguneta," Pennell 
9141 (GH, NY, US) ; "Laguneta" to "Magaiia," Pennell 9425 (GH). Tolima ; 
"La Lora" to "Cucarronera," new Quindio trail, Hazen 9679 (GH). Cundi- 
marca : near Bogota, Ariste- Joseph (US) ; Sibate, 2900-3000 m., Pennell 2517 
(GH, NY, US) ; El Peiion, Pennell 2435 (NY) ; Cortesito, pro. de Mariquita, 
Triana 3809 his (US) ; San Miguel, extremo W. de la sabana de Bogota, 
Ciiatrecasas 6657 (US). Cauca : "Paletara" to "Calaguala," 3000-3200 m., 
Pennell 7083 (GH, NY, US), 7094 (GH, NY, US); "Canaan," Mt. Purace, 
3100-3300 m., Pennell 6669 (GH, NY, US). Narino : Vulcan Galera, near 
Pasto, Lehmann 5497 (GH, F, US) : Putumayo : Corregimiento El Encano, 
Laguna de la Cocha, Paramo El Tabano, Garcia B. 7822 (US). There is con- 
siderable variation in amount of hairiness and the more southern more hairj^ 
specimens may represent F. miniata, but at present I am unable to separate it. 

(41) Fuchsia polyantha Killip, new species 
(Plate?, fig. 37) 

Bush, up to about 1 m. high; young stems quite simple, purplish red, min- 
utely and closely appressed-puberulent under a lens, 2-4 mm. thick ; leaves not 
crowded, ternate or quaternate, rigid, more or less rugose, elliptic to broadly 
oblong-lanceolate, rounded to obtuse at base, acuminate at apex (the apex 
not in the same plane as the rest of the blade and bent at an angle in drying) , 
subentire, light green and glabrescent above except for the appressed puber- 
ulence along the veins, about the same color beneath, minutely appressed- 
puberulent to almost glabrous and with prominent veins beneath, principal 
lateral veins 14-18 on each side of midrib, the submarginal not well formed ; 
blades 6-10 (12) cm. long, 2-3.5 cm. wide; petioles purplish, strigulose, 2-6 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 49 

mm. long; stipules dark, glabrous, subulate, 1.5-2 mm. long, promptly de- 
ciduous ; inflorescence a compact, pendulous, strigulose, many-flowered pan- 
icle, 10-15 cm. long; bracts lance-ovate, 1-2 cm. long; pedicels strigulose, 
purplish, 8-12 mm. long; ovaries strigulose, purple, fusiform, 6-7 mm. long; 
liypanthium purplish-red, tubular, 3.5 cm. long, but little enlarged at base, 
1.5 mm. wide, then 1 mm. wide for one-third its length and very gradually 
ampliate until 5 mm. wide at summit, minutely strigulose without, pubescent 
within basal portion; sepals scarlet, divergent, linear-lanceolate, strigailose 
on backs, 13-15 mm. long, 2.5-3 mm. wide at base, subulate at apex for 1.5-2 
mm. ; petals crimson, oblong-oblanceolate, 10-12 mm. long, 3.5-4 mm. wide, 
subacuminate, glabrous; stamens 11-12 and 7-8 mm. long; anthers 2 mm. 
long; style quite glabrous, equalling or slightly exceeding sepals; stigma 
slightly longer than wide, 1-1.5 mm. thick; berry ellipsoid, 8 mm. long, 4.5-5 
mm. wide. 

Frutex, 1 m. altus; ramulis subpuri3ureis, minute strigulosis, 2-4 mm. diametro; foliis 
ternatis vel quaternis, rigidis, subrugosis, ellipticis vel late oblongo-lanceolatis, basi rotun- 
datis vel obtusis, apice acuminatis, subintegris, subglabris vel in venis strigulosis ; laminis 
6-10 (12) cm. longis, 2-3.5 cm. latis; petiolis subpurj)ureis, strigulosis, 2-6 mm. longis; 
stipulis glabris, subulatis, 1.5-2 mm. longis, caducis; inflorescentia paniculata floribus 
multis; bracteis lanceolato-ovatis, 1-2 cm. longis; pedicellis strigulosis, subpurpureis, 8-12 
mm. longis; ovario striguloso, fusiformi, 6-7 mm. longo; hypanthio purpureo-rubro, tubu- 
loso, 3.5 em. longo, basi 1.5 mm. lato, inde 1 mm. lato, gradatim ampliato, apice 5 mm. lato, 
externe striguloso, interne pubescente; sepalis coceineis, divergentibus, lineari-lanceolatis, 
strigulosis, 13-15 mm. longis, 2.5-3 mm. latis, apice subulatis; petalis rubris, oblongo- 
oblanceolatis, 10-12 mm. longis, 3.5-4 mm. latis, subaeuminatis, glabris; staminibus 11-12 
vel 7-8 mm. longis ; antheris 2 mm. longis ; stylo glabro, sepalis aequalibus ; stigmate 1-1.5 
mm. lato; bacca ellipsoidea, 8 mm. louga, 4.5-5 mm. lata. 

Type: Dept. Narino : between Mayasquer and Tambo, Colombia, 2800 m., 

Aug. 2, 1935, Mexia 7571, U. S. Nat. Herb. 1662416. Another collection from 
ECUADOR: Prov. Carchi: between Moran and Olivos, 3200-3500 m., July 
10-13, 1935, Mexia 7474 (US). The proposed species is near to F. JiirteJla in 
puberulence, divergent sepals, inflorescence, etc., but has the leaves more 
rigid and heavily veined with a sharper apex, the style less exserted, the fruit 
shorter. 

(42) Fuchsia corymbiflora Ruiz and Pavon 
(Plate7, fig. 38) 

Fuchsia corymbiflora Ruiz and Pav^n, F1. Peruv., 3:87, pi. 325, 1802. 

F. corymhosa Pritzel, Ind. Ic, 1:469, 1866, apparently misspelling. 

F.macropetala Presl, Eel. Haenk., 2:28, 1835; from Huanuco, Peru. No specimen seen, 

but description fits here. 
F. dependens Hook., Ic. PL, l:pl. 65, 1837; based on Jameson 81, I am not sure of having 
seen it. 
F. velutina Johnston, Contr. Gray Herb., 75:36, 1925. 

Semiscandent shrub up to 4 or 5 m. tall, the branches rather few, simple, the 
ultimate branchlets commonly 3-5 mm. thick, densely canescent-puberulent 
or -velutinous ; leaves opposite, the blades elliptic to broadly oblong-lanceolate, 



50 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

subentire to inconspicuously denticulate, densely and minutely soft-puber- 
ulent on both surfaces, rounded to obtuse at base, acuminate at apex, 6-11 cm. 
long, 3-5 cm. wide, the principal lateral veins 1-1—16 on each side of the mid- 
rib, with evident submarginal vein ; petioles 6-14 (18) mm. long, velutinous ; 
stipules lance-deltoid, about 1 mm. long, caducous; flowers terminal, few to 
rather many, racemose to corymbose, the bracts oblong-linear to -lanceolate. 
8-15 (20) mm. long ; pedicels 6-10 (12) mm. long, velutinous ; ovary ellipsoid, 
puberulent, 5-6 mm. long; hypanthium tubular, scarlet, 3.5-6 cm. long, 
slightly dilated beyond ovary, then 1.5-2 mm. wide and very gradually am- 
pliate until 5-7 mm. wide at summit, puberulent without, pubescent within 
much of its length; sepals lance-linear, erect-divergent, scarlet, 12-15 mm. 
long, 3-3.5 mm. wide, acute to acuminate, puberulent on backs ; petals deeper 
red, narrow-oblong, 13-16 mm. long, about 3 mm. wide, acute to obtusish ; 
stamens 10-11 and 8-9 mm. long, with reddish filaments ; anthers 2 mm. long; 
style pubescent in lower part, reddish, exceeding sepals by a few mm. ; stigma 
subglobose, slightly lobed, 2.5-3 mm. wide ; fruit round-ellipsoid, densely 
puberulent, 6-10 mm. long, 4-6 mm. wide. 

Type locality, Chinchao, Huanuco, Peru. As here treated this species is 
quite different from the usual concept. I agree with Johnston (Contr. Gray 
Herb. 75 :37. 1925) that most material which has been treated as cory ml) (flora, 
including illustrations, is loliviana, but I feel he erred in saying that the' 
former had a glabrous inner surface to the hypanthium. In fact, I do not see 
what he did include in corymhiflora. However, the Ruiz and Pavon specimen 
now available at Field Museum reveals definitely what it is and shows it to be 
a plant of very minute soft pubescence, rather short round-ellipsoid fruits, and 
agreeing with material occurring from Ecuador to north central Peru and 
including F. dependens and velutina. 

Representative material is, ECUADOR: In Andibus Ecuadoriensibus, 
Spruce 5471 (NY). Prov. Carchi: Guaca San Gabriel, 3300 m., Balls 7344 
(US) ; Canton Tulcan, near Pun, 2987 m., Mexia 7595 (POM, US). Prov. 
Pichincha : west side, Pichincha, 2700 m., Jameson, possible type coll. depen- 
dens (GH, NY, US) ; Canton Quito, road Cotocallao near Nono, 2987 m., 
Mexia 7658 (POM, US) ; between Calacati and Nono, Hartweg 986 (NY). 
Prov. Cotopaxi ? : Hacienda Sumbagua, 3600 m., Haught 2942a (US) . PERU : 
Uuiz and Pavon, ex Herb. Madrid, probably from Huanuco (F). Huanuco: 
Yanano, 2300m., Machride 3715, type no. velutina (F, GH) ; road Mirador to 
Chanchao, 2300 m., Mexia 7765 (GH, US); Carpis, road Acamayo to Chin- 
chao, 2800 m., Mexia 04129 (GH). 

(43) Fuchsia abrupta Johnston 
(Plate 7, fig. 39) 
Fuchsia abrupta Johnston, Contr. Gray Herb., 75 :37, 1925. 

More or less scandent shrub, with glabrous divaricate twigs 4-5 mm. thick ; 
leaves opposite, subcoriaceous, lance-elliptic to broadly oblanceolate, 7-12 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 51 

em. long, 2.5^ cm. wide, subentire, gradually narrowed at base, acuminate 
at apex, dark green and subglabrous above, paler and puberulent beneath 
especially along the veins, principal lateral veins about 20 on each side of 
midrib, submarginal vein evident; petioles pubescent, 4-8 mm. long; stipules 
lance-deltoid, 2-3 mm. long, deciduous; inflorescence a terminal pendant 
raceme, 1-2 dm. long ; bracts reflexed, lanceolate, 8-18 mm. long, somewhat 
loose-pubescent ; pedicels 2.5-4 cm. long, quite glabrous ; ovary linear, gla- 
broiLS, 6-8 mm. long, 1 mm. wide ; hypanthium scarlet, tubular, 3.5-4.5 cm. 
long, somewhat nodose at very base, then 1 mm. thick, then gradually ampliate 
until 4-5 mm. wide at summit, glabrous without, 'pubescent within lower 
portion ; sepals scarlet, lanceolate, 13-15 mm long, acuminate, erect-divergent, 
the tips subulate for 1.5-2 mm.; petals oblong, obtuse, 14-15 mm. long; sta- 
mens 10 and 8 mm. long; anthers 2 mm. long; stigma subglobose, slightly 
lobed, 2 mm. wide ; fruit cylindric-oblong, 12-14 mm. long, 4 mm. wide. 

Type collection from Cushi, dept. Huanuco, Peru, at 1600 m., Machride 
4541 (F, GH, US). This is the only collection I have seen; it has glabrous 
stems, narrow leaves, long pedicels and glabrous hypanthium. 

(44) Fuchsia Cuatrecasasii Munz, new species 

(Plate 7, fig. 40) 
Shrub; young twig's glabrous except sometimes toward the inflorescence, 
greenish, 2-4 mm. thick ; leaves opposite, membranaceous, not crowded, ellip- 
tic-ovate, rounded to obtuse at base, acute to subacuminate at apex, subentire, 
light green and quite glabrous above, mostly glabrous beneath except for 
some exceedingly minute appressed puberulence along the veins, principal 
lateral veins 10-12 on each side of midrib, submarginal weakly developed ; 
blades 4-16 cm. long, 2-9.5 cm. wide; petioles more or less strigulose, 2-18 
mm. long; stipules deltoid, more or less connate and persistent, 1-1.5 mm. 
long; flowers few, in short terminal racemes; bracts lanceolate, 5-15 mm. 
long; pedicels slender, 7-13 mm. long, almost glabrous; ovary glabrous, ellip- 
soid, purplish red, 4—5 mm. long, 1.5-2.5 mm. wide ; hypanthium bright red, 
29-40 mm. long, 2-3 mm. wide at base, then slightly narrowed for one-third 
its length, then gradually ampliate until 6-8 mm. wide at summit, glabrous 
to sparingly pubescent without, densely pilose within ; sepals scarlet, diver- 
gent, 10-13 mm. long, 4-5 mm. wide, glabrous, subulate at apex for about 1 
mm. ; petals scarlet, narrowly oblong-ovate, 8-10 mm. long, about half as wide, 
somewhat pointed, glabrous; stamens 7-8 and 5-6 mm. long; anthers 1 mm. 
long ; style sparsely pubescent, not equalling petals ; stigma barely 1 mm. 
thick; berry apparently oblong-ellipsoid. 

Frutex; ramulis glabris, 2—4 mm. diametro; foliis oppositis, membranaceis, elliptico- 
ovatis, basi rotundatis vel obtusis, apice acutis vel subacuminatis, subintegris, subglabris, 
laminis 4-16 cm. longis, 2-9.5 cm. latis ; petiolis strigulosis, 2-18 mm, longis ; stipulis del- 
• toideis, connatis, 1-1.5 mm. longis; floribus paucis, in racemis, bracteis lanceolatis, 5—15 
mm. longis; pedicellis tenuibus, 7-13 mm. longis; ovario subglabro, ellipsoideo, 4-5 mm. 
longo, 1.5-2.5 mm. lato; hypanthio rubro, 29-40 mm. longo, basi 2-3 mm. lato, inde 



52 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

angustiore, inde ampliato, apice 6-8 mm. lato, exteriore subglabro, interiore piloso ; sepalis 
coccineis, divergentibus, 10-13 mm. longis, 4-5 mm. latis, glabris, 1 mm. subulatis ; petalis 
coccineis, anguste oblongo-ovatis, 8-10 mm. longis; staminibus 7-8 vel 5-6 mm. longis; 
antheris 1 mm. longis; stylo sparse pubescente; stigmate 1 mm, lato; bacca oblongo- 
ellipsoidea. 

Type : from Comisaria del Caqueta, Colombia : Cordillera Oriental, verti- 
ente oriental Quebrada del Rio Hacha, bosque abierto en Cajon de Pulido, 
1700 m. alt., March 26, 1940, /. Cuatrecasas 8738, U. S. Nat. Herb. 1796401. 
An additional collection from the Com. de Putumayo, COLOMBIA, is from 
between El Sileneio and La Cabaiia (carretera de Sibundoy a Urcusique), at 
2200-2400 m., Dec. 31, 1940, Cuatrecasas 11495 (US). The species seems quite 
near F. abrupt a of Peru, but has shorter leaves with fewer veins, shorter 
sepals, shorter inflorescence and pedicels. 

(45) Fuchsia Killipii Johnston 
(Plates, fig. 41) 
Fuchsia Killipii Johnston, Contr. Gray Herb., 81:94, 1928. 

Somewhat scandent shrub, up to 4 m. tall, the younger branches brownish, 
sub terete, somewhat puberulent ; leaves opposite or ternate, elliptic, acute at 
base, acute to acuminate at apex, subentire, almost equally green on both 
surfaces, somewhat coriaceous, with about 12 principal lateral veins on each 
side of the midrib and evident submarginal vein, subglabrous to minutely 
strigulose on both surfaces, the blades 5-9 cm. long, 2-3.5 cm. wide; petioles 
5-10 mm. long; stipules deltoid, scarcely 1 mm. long, deciduous; flowers in 
upper axils and in terminal drooping racemes or panicles, rather numerous ; 
bracts elliptic-ovate, 1-2 (3) cm. long; pedicels subfiliform, 1-2 cm. long at 
anthesis, 2-4 cm. long in fruit, subglabrous; ovary round-ellipsoid, incon- 
spicuously puberulent, 3—4 mm. long ; hj^ianthium red, tubular, 3-4 cm. long, 
1.5-2 mm. wide at the base, then narrowed for one-third its length, then 
rather abruptly widened until 5—7 mm. at summit, glabrous without, retrorse- 
villous within; sepals lanceolate, divergent, acuminate, 14-15 mm. long, 4 
mm. wide, somewhat puberulent toward apex of outer surface, subulate 1-1.5 
mm. at tips ; petals bright red, oblong-oblanceolate, 13-15 mm. long, 2.5-4 mm. 
wide, obtuse to short-acute, glabrous; stamens 12-13 and 9-10 mm. long, 
glabrous ; anthers white, 2 mm. long ; style villous through much of its length, 
up to 5 mm. longer than sepals; stigma 4-lobed, subglobose, 1.5 mm. long; 
berry globose, 5-6 mm. thick. 

Type locality, Rio Surata above Surata, Santander, Colombia. The species 
is distinguished from its closest relatives by having fewer veins, smaller fruit, 
and longer pedicels. I have seen the following collections from COLOMBIA : 
Santander : Rio Surata valley, above Surata, 2000-2300 m., Killip and Smith 
16695, type coll. (Gil, NY, US), 16604 (GH, US) ; near Charta, 2600 m., 
Killip and Smith 18854 (GH, NY. US), 17459 (GH, NY, US) ; near La Baja, 
2200-2600 m., Killip and Smith 16776 (F, GH, NY, US) . 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 53 

(46) Fuchsia boliviana Carr. 

Fuclisia boliviana Cabr., Eev. Hort., 1876:150, with fig., 1876. 

Bushy shrub or small tree up to 4.5 (6) m. high, with rather open brauch- 
ing-, the branchlets spreading or dependent, mostly 2-4 (5) mm. thick, terete 
to angled, densely pubescent; leaves opposite, sometimes alternate or ternate, 
elliptic to oblong-lanceolate or -ovate, obtuse to rounded at base, acute to 
acuminate at apex, usually somewhat denticulate, soft-pubescent on both 
surfaces to pilose or puberulent or glabrescent, somewhat paler beneath, with 
about 12-24 principal lateral veins on each side of midrib ; leaf -blades 5-15 
(20) cm. long, 3-6 (10) cm. wide; petioles 1.9-3 (4.5) cm. long, pubescent; 
stipules subulate-lanceolate, scarcely 1 mm. long, deciduous; flowers few to 
many, in terminal drooping pubescent corymbs or panicles which elongate 
in fruit; bracts 7-20 (30) mm. long, 3-9 mm. wide, usually lanceolate and 
reflexed; pedicels 5-15 mm. long; ovary oblong-linear, 5-7 mm. long, 1.5-2 
mm. thick; hypanthium tubular, usually dark red, 3-6 cm. long, slightly en- 
larged at base, then 1.5-2 mm. wide and very gradually ampliate until 3.5-7 
mm. at apex, more or less pubescent without and within ; sepals usually dark 
red, lanceolate, acuminate, 10-20 mm. long, 2.5-5 mm. wide, spreading- 
reflexed in anthesis, more or less pubescent without ; petals dark to light red, 
erect to divergent, oblong, 9-13 mm. long, 3-4.5 mm. wide, acute ; episepalous 
stamens equaling or slightly exceeding petals ; alternate three-fourths as long ; 
filaments glabrous, reddish ; anthers light in color, 2-3 mm. long ; style slender, 
pubescent within hypanthium, scarcely reaching tips of sepals ; stigma sub- 
globose, slightly lobed, 1-2 mm. thick; berry oblong-ellipsoid, 1-2 cm. long, 
4-6 mm. thick, pubescent. 

As here understood, this species is common and widespread, including most 
of the material once referred to F. corymbiflora, from which it can be dis- 
tinguished by its usually longer and more spreading hairs, spreading sepals, 
apparently less vinelike habit, more apiculate buds and elongate fruit. 

Key to Varieties of Fuchsia 'boliviana 
A. Hypanthium 3-4 cm. long ; sepals 10-18 mm. long. South of Ecuador. 

B. Plant pilose or pubescent with spreading hairs. Peru to Argentina. 46a. var. typica. 

BB. Plant minutely puberulent. Bolivia 46b. forma. Tpuherulenta. 

A A. Hypanthium 5-6 cm. long; sepals 17-21 mm. long. North of Ecuador. 

46c. var. luxurians. 

(46a) Fuchsia boliviana Carr. var. typica Munz, new name 

(Plates, fig. 42) 

F. boliviana Carr., Kev. Hortic, 1876:150, with pi., 1876; Johnston, Contr. Gray Herb., 

75:37, 1925. 
F. corymbiflora of many authors, not of Euiz and Pavon, 1802. 

Plant usually soft-pilose or -pubescent on all new growth ; hypanthium 
2.5-4 em. long ; sepals mostly 12-18 mm. long. 

Type locality, "sur les montagnes a la Bolivie, a 6000 metres environ d'alti- 



54 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

tucle." Such an altitude is quite impossible ; I have seen specimens from as 
high as 4000 m.; the variety ranges from Peru to northern Argentina. In the 
southern part of the range the plants tend to be less densely pilose and 
smaller-flowered. Representative material may be cited as follows, PERU : 
Huanuco: 5 km. SE. of Carpish, 2850 m., Stork 9921 (F). Huaneavelica : at 
2400 m., Weherhauer 6565 (F, GH). Ayacucho: Prov. de la Mar, Tambo, 
Weberbauer 5555 (F, GH, US) ; Ccarrapa, between Huanta and Rio Apuri- 
mac, 2800 m., Killip and Smith 22280 (NY, US). Apurimac : Abancay : 
Ampuy, 3100 m., Stork, Horton and Vargas 10595 (F) ; Huancarama, 3200 
m., West 3783 (GH). Cuzco : Urubamba Valley, West 6462 (GH, US), Her- 
rera 2094 (US), 1123 (F, US) ; Paucartambo, Balls 6715 (US); OUantay- 
tambo. Cook and Gilbert 290 (US). BOLIVIA: La Paz: Yungas, Bang 327 
(F, GH, NY, US), Rusby 1071 (F, GH, NY, US) ; Sorata, 2600-2800 m., 
Mandon 622 (F, GH, NY, US). Santa Cruz: Samaipata, 1800 m., Steinbach 
3755 (POM); Comarapa, Steinbach 8459 (GH, NY). Cochabamba : Prov. 
Chapare: Incacorral-Aduana, 2700 m., Steinbach 9538 (F); Prov. Sacaba: 
Cerro de Ineachaea, 2500 m., Steinba.ch 5763 (F, GH, US). ARGENTINA: 
Jujuy : San Lorenzo, Joergensen 36041 (POM) . Tueuman : Depto Chicligasta, 
Estancia Las Pavas, Ve7ituri 4657 (F, US) ; Villa Nougues, 1200 m., Munz 
15469 (NY, POM, US); Las Lenguas al Clavillo, 1750 m., Munz 15475 
(POM) ; Depto. Famailla: Quebrada de Lules, Venturi 1295 (F, CAS, GH, 
US). 

(46b) Fuchsia boliviana Carr. forma puberulenta Munz, new form 

Plant with dense minute subappressed puberulence; hypanthium 3-4 cm. 
long; sepals mostly 10-16 mm. long. (Planta cum puberulencia minuta et 
densa : hypanthio 3-4 cm. longo.) 

Type: from La Paz, Bolivia: Nor Yungas: Milluguaya, 1800 m., Buchtien 
732, Field Mus. no. 588700; isotypes F, GH, NY, US. Other material all from 
BOLIVIA : La Paz : Unduavi Valley, 2000-2600 m., Bro. Julio 478 (US) ; Nor 
Yungas: Coroica to La Paz, Mexia 04286 (GH) ; Sur Yungas: SirUpaya near 
Yanaeachi, 2100 m., Buchtien 732 (F, GH) ; Colaya, Mexia 7809 (F, US). 
Cochabamba : Prov. Chapare, 110 km. north east of Cochabamba, near Chi- 
more, 800 m., Eyerdam 24740 (F). This form resembles F. corynibiflora in 
puberulence, though the latter has it less dense, but F. boliviana has spreading 
sepals and cor^m&i^ora does not. 

(46c) Fuchsia boliviana Carr. var. luxurians Johnston 

Fuchsia boliviana Carr. var. luxurians Johnston, Contr. Gray Herb., 75:38, 1925. 

F. cuspidata Fawcett and Rendle, Journ. Bot., 64:105, 159, 1926, as to tj'pe cited: Harris 

5825. 
F. corynibiflora of many authors: LiNDL., Bot. Reg., 26:pl. 70, 1840; Hook,, Bot. Mag., 

69: pi. 4000, 1843; not of R. and P., 1802. 

Plant soft-pilose to -pubescent; hypanthium 5-6 cm. long; sepals 17-21 
mm. long. 



Vol. XXV] MVNZ: IRE GENUS FUCHSIA 55 

Type locality, Colonia Tovar, Venezuela. Representative material, JA- 
MAICA : Cinchona, Harris 7643 (NY), Harris and Lawrence C 15438 (NY) ; 
St. Helen's Gap, St. Andrew, 3Iaxon and Killip 571, 1475 m. (F, GH, NY, 
US). GUATEMALA : Cartago : Cartago, 1350 m.. Smith 4804 (US) ; Zacate- 
IDequez: San Rafael, 2000 m., Smith 2176 (US). San Marcos: Tajumulco, 
northwest slope of Volcan Ta.jmnnlco, 2300 m., Steyermark 36510 (F). 
Chimaltenango : San Marcos, Johnson 1231 (F). EL SALVADOR: bought 
in market of San Salvador, brought from Volcan de San Salvador, C alder on 
2345 (F, US) . COLOMBIA : Antioquia : "Las Minitas," so of Caldas, 2650 m., 
Pennell 10970 (GH, NY); Medellin, Toro 721 (NY). Caldas : Salento to 
"Lagunita," old Quindio trail, 2000 m., Pennell 10594 (GH). Cundimarca : 
Bogota, 2700 m., Schultze 135 (US), Perez 4802 (US). VENEZUELA: 
Colonia Tovar, 1800-2000 m., Pittier 9252, type no. (GH, NY, US), Allart 
354 (NY, US), Ja/iw 439 (US). 

(47) Fuchsia fulgens DeCandolle 
(Plate 8, fig. 43) 

Fuchsia fulgens DC, Prodr., 3:39, 1828; in Sesse and Mo§ino, Icon. Fl. Mex. ined., 362; 

LiNDLEY, Bot. Eeg., 24:1, 1838; Curtis, Bot. Mag., 67:3801, 1840; Hemsley, Biol. Centr. 

Amer., Bot., 1:457, 1880; Standley, Contr. U. S. Nat. Herb., 23:1078, 1924. 
Ellohium fulgens Lilja, Linnaea, 15:262, 1841. 

Shrub, commonly 3-12 dm. tall, with tuberous-thickened roots, soft woody 
stems, few-branched; young branches sparsely pubescent, these and the leaves 
tinged red ; leaves broadly ovate to oblong-ovate, opposite, paler beneath than 
above, fiue-pubescent on both surfaces, rounded to more or less cordate at 
base, abruptly short-acuminate, shallowly and evenly serrulate, the blades 
thin, 5-17 cm. long, 3-12 cm. wide, with about 10 principal lateral veins on 
each side of the midrib; petioles 1.5-10 cm. long; flowers several, in short 
racemes with an axis 5-20 cm. long; lower bracts leafy and ovate, 1-3 cm. long ; 
upper reduced, lanceolate, 0.5 cm. long, puberulent ; pedicels pendant, very 
slender, 1-2 (3.5) em. long; hypanthium 5-6.5 cm. long, about 2 mm. wide at 
base, then scarcely narrowed and then gradually ampliate until 5-7 mm. wide 
near mouth, dull scarlet, subglabrous to puberulent without, pubescent with- 
in ; sepals 4, yellowish to greenish with somewhat pale or yellow or red base, 
ovate-lanceolate, 12-14 mm. long, subglabrous except for the sparse pubes- 
cence near the acuminate tips; petals bright red, oblong-ovate, obtuse, 8 mm. 
long, 6 mm. wide ; stamens about 10 and 7 mm. long ; anthers 2-3 mm. long, 
filaments whitish to reddish ; style filiform, pink, pubescent near base, slightly 
exceeding sepals; stigma greenish, subconic, 2 mm. long; fruit ellipsoid, sub- 
glabrous, dark, 2 or more em. long. 

Type locality not known ; the species is Mexican, is near to such long-tubed 
species as F. holiviana, corymhiflora, etc., but quite distinct in its green sepals 
and leaves of cordifolia type. It may be an epiphyte, may grow on ledges, 
walls, or more open ground. Material seen, "Fuchsia cordifolia N. No. 195. t 




56 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

2. f. 159 i. c. 195," Sesse, Mocino et al 5211 (F) ; photo of Geneva copy of 
Sesse and Mocino plate (F) ; H. B. Basil, Aug. 1841 (NY). MEXICO : Mex- 
ico City, Russell and Souviron 132 (US). Mexico: Temascaltepec District, 
Rincon, Hinton 1144 (GH, US) . Michoacan : near Tarascon, Pringle 11988^2, 
at 2000 m., (F, GH, US) ; near Patzcuaro, Pringle 4123 (F, GH, MICH, NY, 
POM, US) ; Morelia, Campanario, 2200 m., Arsene in 1911 (US) ; Monte San 
Miguel, Arsene 6673 (US) ; 2 miles north of Zitacaura, Hitchcock and Stan- 
ford 7207 (US) ; Tancitaro, Leavenworth 303 (F). Jalisco: Volcano Colima, 
2100 m., Goldsmith 22 {Gil). 

(48) Fuchsia decussata Ruiz and Pa von 

(Plate 8, fig. 44) 

Fuchsia decussata Euiz and Pav6n, PL Peruv., 3 :88, pi. 323, fig. b., 1802. 

F.scandens Krause, Fedde Eep., Nov. sp. 1:171, 1905; based on Weberbauer 3324 from 

Monzon, Huanuco, Peru ; have seen photograph. 
F. fusca Krause, Engl., Jahrb., 37:599, 1906; based on Weberbauer 4975 from near Cuzco; 

have seen photograph ; flowers large like Macbride 4414. 
F. fontinalis Macbr., Candollea, 8:25, 1940. Eeduced by Macbride, Field Mus. Nat. Hist., 

Bot., 13, part 4:552, 1941. 

Shrub, supported by other vegetation, up to 3 ni. high, with long flexuous 
branches, the youngest reddish, 1-3 mm. thick, densely brown-pubescent with 
incurved hairs ; leaves ternate or opposite, numerous, those of main stems 
often remote and supplemented by smaller ones on reduced lateral branchlets, 
coriaceous, elliptic to elliptic-lanceolate or -ovate, cuneate at base, acute at 
apex, serrate to serrulate, dark green and puberulent to strigulose to glabres- 
cent above, paler and short-pilose beneath, especially along veins, principal 
lateral veins 6-10 on each side of midrib, submarginal vein somewhat devel- 
oped, blades 1.5-5 cm. long, 0.5-1.5 (2) cm. wide; petioles pubescent-pilose, 
3-8 (15) mm. long; stipules dark, subulate, 1-1.5 mm. long, persisting; 
flowers solitary in axils; pedicels subfiliform, pilose, 1.5-3 cm. long; ovary 
ellipsoid, 3-4 mm. long, more or less pilose; hypanthium 13-16 (22) mm. long, 
red, 2-3 mm. wide at base, then very little narrowed, 3.5-4.5 mm. wide at 
summit, sparsely pilose without, glabrous within ; sepals divaricate, lance- 
oblong, red with green tips, acute, 7-9 (11) mm. long, 2.5 mm. wide, sparsely 
pilose on backs; petals red, oblong-obovate, obtusish to apiculate, 6-7 (10) 
mm. long, 3-4 mm. wide ; stamens 8-9 and 6-7 mm. long, more or less exserted; 
anthers 1.5-2 mm. long; style glabrous, equaling or exceeding sepals by 2-3 
(6) mm. ; stigma 1.5 mm. wide; fruit red, oblong, obtusely angled, 8-10 mm. 
long, 5-6 mm. thick. 

Type locality, Muila, dept. Huanuco, Peru. The species ranges at elevations 
of 2800-3400 m. from Hiianuco to Cuzco. It has considerable variation in 
width and aeuteness of petals. It falls in a group with axillary flowers and 
short hypanthium and seems to be a possible derivative from the cordifolia 
type. Macbride (Field Mus. Nat. Hist. Bot. Ser. 13, part 4, no. 1 :554, 1941) 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 57 

maintains F. fiisca as a species on the basis of the wider petals and brownish 
pubescence, but this latter character seems to be a general tendency in many 
specimens. Representative material of F. decussata are: PERU: without 
definite locality given, Ruiz and Pavon, F. decussata sp. nov. Herb, at Geneva 
(photo F, POM), ex Herb. Madrid (F). Huanuco: 15 miles SE of Huanuco, 
2900 m., Machride and Featherstone 2124 (F, GH, US), 2081 (F, GH, US), 
2107 (F, GH, US) ; Panao, Machride 3615 (F, GH, US) ; Tambo de Vaca, 
Machride 4414 (F, GH) with petals 10 mm. long; Carpish, Stork and Horton 
9911 (F) with very narrow petals; Carpis, between Acomayo and Chinchos, 
Mexia 04126 (GH), 04128 (GH) ; Monzon, Weherhauer 3324, type scandens 
at Berlin (photo F, POM). Ayucucho : prov. Huanta, between Tambo and 
Apurimac River, Weherhauer 5578 (F). Cuzco : between Cuzco and Santa 
Anna, below Yanamanche, Weherhauer 4975, type fusca at Berlin (photo F, 
POM). 

(49) Fuchsia hypoleuca Johnston 

(Plate 8, fig. 45) 
Fuchsia liypoleuca Johnston, Coiitr. Gray Herb., 75:34, 1925. 

Shrub, 1-2 m. high ; young stems densely canescent-pubescent with more 
or less incurved hairs, slender, 1.5-3 (4) mm. thick; leaves opposite or ternate, 
rigid, crowded, lanceolate, obtuse to acute at base, acute at apex, remotely 
serrulate in upper half, more or less revolute, veiny almost rugose, greenish 
and somewhat strigulose above, pale and densely pilose-pubescent beneath, 
tending to have interwoven fungus mycelium underneath, principal lateral 
veins 6-10 on each side of midrib, submarginal vein present ; blades 3-6 (7.5) 
cm. long, 0.8-1.7 cm. wide; petioles pubescent, 5-10 mm. long; stipules sub- 
ulate, 2-3 mm. long, deciduous; pedicels few, solitary in axils, filiform, pen- 
dulous, pubescent, 1.5-2 cm. long; ovary ellipsoid, pubescent, 4 mm. long; 
hypanthium red, 18-23 mm. long, about 2.5 mm. wide at base, then somewhat 
narrowed and then gradually ampliate until 5 mm. wide, pubescent and 
villous within lower half ; sepals divaricate, scarlet, lanceolate, 9-12 mm. long, 
3 mm. wide, pubescent on backs, subulate for 1 mm. at apex; petals scarlet, 
obovate, 7 mm. long, 4^5 mm. wide, rounded at apex ; stamens 7-8 and 5-6 
mm. long; anthers 1.5 mm. long; style pubescent within the hypanthium, 
equalling the sepals or exceeding them by 2-3 mm.; stigma green, 1.5 mm. 
wide ; fruit apparently oblong-ellipsoid, 4-angled, 8-10 mm. long, 3-4 mm. 
wide. 

Type locality, between Loja and San Lucas, prov. Loja, Ecuador. The 
species seems to me in its floral position and h^^Danthium shape to be near 
F. decussata, but amply distinct in the narrow^ crowded leaves which are 
whitish beneath. I have seen the following specimens, ECUADOR : without 
definite locality, Jameson (F, US), southern provinces, Jameson (US) ; be- 
tween Loja and San Lucas, 2100-2600 m. Hitchcock 21440, type coll. (GH, 
NY, US). 



58 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Rer. 

(50) Fuchsia scabriuscula Bentliam 

(Plate 8, fig. 46) 
Fuchsia scabiuscula Benth., PI. Hartweg., 177, 1845. 

Spreading shrub, 7-20 dm. tall, the younger twigs sometimes reddish, 2-4 
mm. thick, densely pilose with white to somewhat brownish hairs ; leaves 
opposite, elliptic to elliptic-obovate, rigid and very veiny, even rugose, 
rounded to obtuse at the base, acute to subacuminate at the apex, subentire, 
somewhat strigose above, paler and pilose beneath, especially on veins, with 
10-15 principal veins on each side of midrib and with prominent submarginal 
vein; blades 3-10 (13) cm. long, 1.5-5 (7) cm. wide; petioles 5-25 mm. long, 
densely pilose ; stipules subulate, pubescent, 1.5-3 mm. long, quite persistent ; 
pedicels solitary, axillary, slender, curved, pilose, 1-2 cm. long; ovary nar- 
rowly fusiform, densely pilose, 5-7 mm. long; hypanthium bright red, 15-18 
mm. long, 1.5-2 mm. wide at base, then somewhat narrowed, then gradually 
arapliate until 4.5-5 mm. wide at summit, densely pilose without and within 
the basal half ; sepals bright red, divergent, lance-oblong, 8-9 mm. long, ea. 
3 mm. wide, subulate at apex for about 1.5 mm., pilose on back; petals red, 
obovate-oblong, 8-9 mm. long, about 0.5 mm. wide, rounded at apex ; stamens 
about 8 and 6 mm. long; anthers 1.5 mm. long ; style densely pubescent within 
hypanthium, extending about to tips of petals ; stigma greenish, about 2 mm. 
Made ; fruit oblong, 10-12 mm. long, 3-4 mm. wide. 

Tj^pe locality, "In declivities oecidentali Andium Quitensium." Ecuador, 
This species with F. verrucosa seems to me to be near F. decussata, but with 
a somewhat more narrow base to the hypanthium. It is characterized by the 
very veiny, rugulose leaves, pilose stems and flowers, and short hypanthium. 
It seems to be confined to Ecuador, from which I have seen the following 
material: in Andibus Ecuadorensibus, Spruce 5038 (GH, NY). Pichincha : 
in declivitate oecidentali Andium Quitensium, Hartweg 987, type no., Geneva 
Herb, (photo F, POM; leaf F) ; Sive Andium Quitensium, 2500 m., Jameson 
53 (NY), 55 (GH) ; between Baeza and Papallacta, 2000 m., Mexia 7339 (GH, 
US) ; Nanegal District, Couthoiiy in 1855 (GH) ; west declivity of Pichincha, 
Jameson 7 (NY). Tungurahua: Valley of Pastaza River, between Baiios and 
Cashurco, 1300-1800 m., Hitchcock 21781 (GH, NY, US) ; Vulcan Tungur- 
agua, LeJimann 4995 (F). A collection from Huigra, by Rose and Rose, no. 
22477 (US) is near this but the leaves are thin. 

(51) Fuchsia verrucosa HartAveg 

(Plate8, fig. 47) 

Fuchsia verrucosa Hartweg, in Benth., PI. Hartweg., 178, 1845. 
F. perbrevis Johnston, Contr. Gray Herb., 75:30, 1925. 

Shrub 1-2 m. high, young twigs subglabrous, slightly red, 2-4 mm. thick ; 
leaves opposite or ternate, thick and firm, elliptic-ovate to ovate-oblong, acute 
to acuminate at base, acuminate at apex, subentire to somewhat serrulate, 
quite glabrous above, paler and subglabrous beneath, or puberulent on the 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 59 

veins; veins very prominent beneath, the prineipla lateral ones about 15 on 
each side of the midrib, submarginal prominent ; blades 4-12 cm. long, 2.5- 
5.5 cm. vs^ide ; petioles 2-10 mm. long ; stipules more or less connate, reflexed, 
about 2 mm. long; flowers few, solitary, axillary; pedicels firm, glabrous, 
1-2.5 cm. long, recurved at anthesis; ovary subglabrous, 10-12 mm. long, 
about 2 mm. wide, 4-angled ; hypanthium obconic, 3-5 mm. long, subglabrous 
without and within ; sepals 8-10 mm. long, 2-3 mm. wide, lance-oblong, diver- 
gent, acutish, bright red; petals red, obovate, 8-9 mm. long, 4-5 mm. wide; 
stamens 3-4 mm. long; anthers 1.5 mm. long; style glabrous, 7-9 mm. long; 
stigma 1.5 mm. thick; fruit 4-angled, 15-18 mm. long, 3.5-4 mm. wide. 

Type locality, between "Paramo de San Fortunato et pagum Fusagasuga, 
prov. Bogota," Colombia. The species seems to be nearest to F. scabriuscula , 
with which it agrees in its thick veiny leaves, but is remarkable, as has been 
pointed out both by Bentham and Johnston in their descriptions, for its very 
short hypnathium. I have seen the following collections, all from COLOM- 
BIA: without definite locality, Muiis 2077 (US), 2047 (US), 1361 (US). 
Dept. Santander del Norte : road Pamplona to Toledo, 2800-3000 m., KilliiD 
and Smith 19976 (GH, NY, US), 19831 (F, GH, NY, US). Cundimarca : be- 
tween Paramo de San Fortunato and Fusagasuga, prov. Bogota, Hartweg 
991, type no. verrucosa, Berlin specimen (photo F, POM), Geneva specimen 
(leaf F) ; Flora Neo-granadina-Bogotana, Ultra Abate (ultra Ubate ?), 
Holton 892, type perhrevis (GH). Huila : Comisaria del Caqueta : Cordillera 
Oriental, en Gabinete, 2300-2450 m., Cuatrecasas 8465 (US). 

(52) Fuchsia sanctae-rosae 0. Kuntze 

(Plate 9, fig. 48) 

Fuchsia sanctae-rosae O. Kuntze, Eev. Gen., Ill, pt. 2 :98, 1898. 

F. holiviana Britton, Bull. Torrey Bot. Club, 17 : 214, 1890, not Carr., 1876. 

F. Brittonii Johnston, Contr. Gray Herb., 75 :39, 1925, new name for 'boliviana Britton, 

F. Weberbeueri Krause, Fedde Rep. Nov. Sp., 1:170, 1905; based on Weberbauer 661, of 

Avhicli I have seen a photograph, 
F. filipes RusBY, Mem. N. Y. Bot. Gard., 7:317, 1927; based on White 232, which I have seen. 

From suffrutescent or almost herbaceous and 3-5 dm. tall, to shrubby and 
2-3 m. high, rather simple to many -branched, erect to semidecumbent ; young 
shoots mostly glabrous, sometimes somewhat pilose, purplish red, 1.5-4 mm. 
thick; leaves mostly in whorls of 3 or 4, firm, elliptic-lanceolate to almost 
-obovate, obtuse to acute at base, acute to acuminate at apex, sub entire to 
minutely serrulate, deep green and glabrous above, lighter and glabrous be- 
neath except usually somewhat pilose along midrib, principal lateral veins 
8-12 on each side of midrib, submarginal frequently not very evident; blades 
3-10 cm. long, 1.5-4 cm. wide ; petioles 5-20 mm. long, usually purplish red 
and glabrous, sometimes appressed-pubescent ; stipules subulate, 1-2 mm. 
long, sometimes reflexed, sometimes connate, deciduous ; flowers rather nu- 
merous, but solitary in upper axils of somewhat reduced leaves, hence more or 



60 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

less racemose; pedicels very slender, 1-2.5 cm. long, mostly glabrous; ovary 
narrow ellipsoid, glabrous to puberuleut, 4-6 mm. long ; hypanthium deep to 
bright red, 13-18 (22) mm. long, 2-2.5 mm. wide at base, then slightly nar- 
rowed and then gradually ampliate until about 4 mm. wide at summit, gla- 
brous without, pilose within the lower half ; sepals scarlet, lance-oblong, 8-10 
mm. long, 2.5-3 mm. wide, glabrous or sometimes pilose, acutish ; petals 
orange-red to scarlet, oblong-obovate, 6-8 mm. long, 3-6 mm. wide, obtuse to 
acutish, sometimes almost 3-toothed; stamens 7-9 and 5-7 mm. long; anthers 
about 1.5 mm. long; style mostly glabrous, extending about to the sepal-tips; 
stigma red, about 1 mm. wide ; berry varying from globose-ellipsoid to oblong- 
ellipsoid, 6-10 mm. long, 4-5 mm. wide. 

Type locality, Santa Rosa, Bolivia (there are several towns of this name in 
Bolivia). I have not seen the type. The species is variable in every way, but I 
cannot find any constant characters for division. It comes between the race- 
mose and axillary-flowered groups with short hypanthium, so seems to stand 
between the species I have just treated and those like Osgoodii, Andrei, etc., 
which follow. It ranges at altitudes between 2000 and 3000 m., in southern 
Peru and Bolivia. Representative material, PERU : Dept. Cuzco : "Pillahu- 
ata," Cerro de Cusilluyoc, Pennell 13997 (F, GH, NY, US), 13956 in part 
(GH); prov. Paucartambo, valley of Rio Tambomayo, West 7091 (GH) ;• 
Macchu-Picchu, Munz 15548 (POM), West 6414 (GH), 3Iexia 8078 (F, GH, 
US), Balls 6802 (US). Puno : Sandia, Weherhauer 661, type Wederhaueri at 
Berlin (photo F, POM). BOLIVIA : Santa Rosa, as serratifolia, 0. Kuntze in 
1892 (F, GH). La Paz: Haciendo Simaco, sobre el camina a Tipuani, Biich- 
tien 832 (F, POM), 5508 (GH, NY, US) ; Pongo, Pulcheri, White 232, type 
no. filipes (GH, US) ; Prov. Larecaja : near Ananea : Machacamarca, Mandon 
624 (F, GH, NY) ; Prov. No. Yungas, between Unduavi and Chirca, Eyerdam 
25388 (F) ; Yungas, Bmig 675 (F, GH, NY, US), Rushij 1812 (F, NY, US), 
1813 type no. holiviana, Brittoni (NY, US). Cochabamba : prov. Chapare, 
Steinhach 9227 (GH) ; Incachaca, Cdrde^ias 665 (US). 

(53) Fuchsia Osgoodii Macbride 

(Plate9, fig. 49) 

Fuchsia Osgoodii Macbride, Field Mug. Nat. Hist., Botany, 13, part 4, no. 1 :559, 1941. 
F. ovalis E. and P. var. aherrans Macbride, I.e., 560. 

Shrub, up to 4 m. tall ; twigs dark, subglabrous, 1-2.5 mm. thick ; leaves 
opposite, firm, elliptic-lanceolate, acute to acuminate at both ends, quite entire, 
deep green, glabrous above, paler and glabrous beneath except minutely 
puberulent or short-pilose on veins, with 10-12 principal lateral veins on 
each side of midrib, submarginal present; leaf-blades 5-8 (12) cm. long, 
1.5-3 (6) cm. wide; petioles glabrous, 8-10 mm. long; flowers few, in short 
congested racemes; bracts 1-2 cm. long; pedicels filiform, 1.5-2.5 cm. long, 
glabrous ; ovary linear-ellipsoid, ca. 5 mm. long ; hypanthium 23-24 mm. long, 
dark red, somewhat enlarged at base, where 1.5.2 mm. wide, then 1 mm. wide 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 61 

for one-third its length, then gradually ampliate until 4-5 mm. wide at sum- 
mit, glabrous without, villous within lower portion ; sepals red, oblong-ovate, 
10-11 mm. long, 5-6 mm wide, glabrous, subulate for 1-1.5 mm. at apex; 
petals red, oblong, about 10 mm. long, 4 mm. wide, abruptly acute; stamens 8 
and 6 mm. long ; anthers 1.6 mm. long; style glabrous, equalling petals ; stigma 
4-lobed, 1.5 mm. wide; berry oblong, at least 12 mm. long and 4 mm. wide. 
Type locality, PERU : Dept. Libertad : Ucheo, the type Osgood and Ander- 
son 47 (F), From Cajamarca: prov. Cutervo : Arenales, Stork and Horton 
10155 (F) comes the type of ovalis var. aherrans, which has some lower and 
larger leaves than the type of Osgoodii. They are more pilose on the veins 
beneath the leaves, but the upper smaller leaves have the same puberulence 
as in Osgoodii. F. Osgoodii is near F. sanctae-rosae in hypanthium, foliage, 
etc., but has more definitely terminal inflorescence. 

(54) Fuchsia AndreiJohnston 
(Plate9, fig. 50) 

Fuchsia Andrei Johnston, Contr. Gray Herb., 75 :31, 1925. 

Shrubby, the younger twigs more or less purplish red, fairly coarse, sub- 
glabrous except toward the minutely pubescent tips, 3-8 mm. thick ; leaves 
opposite or ternate, oblong-elliptic to elliptic-ob ovate, mostly obtuse at base, 
abruptly acuminate at apex, membranaceous, subentire, sparsely ciliolate, 
green and subglabrous above, paler beneath and puberulent on veins, with 
about 12 principal lateral veins on each side of midrib and with evident sub- 
marginal vein; blades 7-15 cm. long, 3-7 (9) cm. wide; petioles subglabrous, 
1-2.5 cm. long; stipules deltoid, ca. 1 mm. long, deciduous; inflorescence a 
short lateral or terminal raceme; bracts lanceolate, 3-10 mm. long, somewhat 
pubescent; pedicels slender, somewhat pubescent, 10-15 mm. long; ovary 
oblong, subglabrous, about 5 mm. long ; hypanthium 2-3 cm. long, somewhat 
enlarged at base, then 1 mm. wide for about one-third its length, then gradu- 
ally ampliate until 4-4.5 mm. Avide at apex, subglabrous without, pilose with- 
in lower half ; sepals oblong-lanceolate, 7-9 mm. long, divergent, subulate for 
1.5 mm. at tips, almost glabrous; petals lance-oblong, 9-10 mm. long, 2.5-3 
mm. wide, obtuse ; stamens about 8 and 6 mm. long ; style pilose in basal half, 
exceeding sepals by 3^ mm. ; stigma globose, 4-lobed, ca. 1 mm. thick ; berry 
oblong, 6 mm. long and 3 mm. wide. 

Type locality, Rio de Huannamaca, probably southern Ecuador. I have 
seen two collections : Rio de Huannamaca, Andre K820, type no. (F) ; Tambo 
de Savanilla, Andre K 818 (F, GH, NY). The leaves are large and glabrous, 
the flowers in terminal racemes. 

(55) Fuchsia Lehmannii Munz, new species 
(PlateQ, fig. 51) 

Soft-wooded shrub up to 2 m. high, the younger twigs dark red, mostly 
glabrous, 1-3 mm. thick; leaves opposite or ternate, elliptic-lanceolate or 



62 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

-oblanceolate, acuminate or acute at both ends, rigid, subentire, glabrous, and 
dark green above, beneath paler and glabrous except for some short appressed 
pubescence on the reddish veins, the principal lateral veins 8-12 on each side 
of midrib ; leaf -blades mostly 5-10 (12) cm. long, 1.5-3 (4) cm. wide; petioles 
glabrous to somewhat strigulose, 5-15 mm. long; stipules lance-subulate, 1 
mm. or so long, deciduous; flowers numerous, crowded in short lateral and 
terminal racemes arranged paniculately; bracts lance-linear, reflexed, some- 
what pubescent, 1-2 cm. long; pedicels slender, pubescent, 8-18 mm. long; 
ovary narrow-fusiform, more or less strigulose, 3-5 mm. long ; hypanthium 
red, 24-30 mm. long, 1 mm. wide at base, then narrowed for one-third its 
length, then gradually ampliate until 3.5-5 mm. wide at apex, mostly glabrous 
without and pubescent within the narrow basal portion; sepals scarlet, re- 
flexed-spreading, linear-lanceolate, glabrous, 9-10 mm. long, 2.5-3 mm. wide, 
subulate for 1-2 mm. at apex; petals scarlet, oblong-lanceolate, about 1 cm. 
long, 2-2.5 mm. wide, obtusish ; stamens 7 and 5.5 mm. long ; anthers 1 mm. 
long; style glabrous, slightly exceeding petals; stigma 1-1.5 mm. wide; fruit 
ellipsoid, 5-6 mm. long, 3-4 mm. wide. 

Frutex, 1-2 m. altus ; ramulis rubris, subglabris, 1-3 mm. crassis ; f oliis oppositis vel 
ternatis, elliptico-lanceolatis vel -oblanceolatis, acuminatis vel acutis, rigidis, subintegris, in 
superficie superiore glabris, inferiore glabris vel in venis puberulentis, laminis 5-10 (12)  
cm. longis, 1.5-3 (4) cm. latis; petiolis glabris vel substrigulosis, 5-15 mm. longis; floribus 
numerosis, in racemis paniculatis; bracteis lanceo-linearibus, reflexis, pubescentibus, 1-2 
cm. longis; pedicelis tenuibus, pubescentibus, 8-18 mm. longis; ovario angusto-fusiforme, 
strigulose, 3-5 mm. longo ; hypanthio rubro, 24-30 mm. longo, base 1 mm. lato, inde angus- 
tiore, inde ampliato et apice 3.5-5 mm. lato, externe subglabro, interne pubescente; sepalis 
coceineis, reflexo-divergentibus, lineare-lanceolatis, glabris, 9-10 mm. longis, cum apicibus 
subulatis 1.5-2 mm, longis; petalis coceineis, oblongo-lanceolatis, 1 cm. longis, 2-2.5 mm. 
latis, oBtusis ; staminibus 7 et 5.5 mm. longis ; antheris 1 mm. longis ; stylo glabro, petala 
excedentibus ; stigmate 1-1.5 mm. lato; bacca ellipsoidea, 5-6 mm. longa, 3-4 mm. lata. 

Type: from Loja, Ecuador: east Andes of Sigsig, 1600-1800 m., Lehmann 
5498 (equal Pearce's 235), type Field Museum 550994; isotypes GH, US. A 
second collection also from Loja : Rio Savanilla, east Andes of Loja, 800-1300 
m., Lehmann 7858 (P, US) . The proposed species is near F. Osgoodii in leaves 
and flowers, but sepals are more narrow and berry shorter ; near F. Hartwegii 
in flower-shape, but glabrous and with more pointed leaves ; near F. Andrei, 
but with glabrous style and narrow leaves. 

(56) Fuchsia putumayensis Munz, new species 
(Plate 8, fig. 52) 

Shrub or tree, younger twigs guite glabrous, with some purplish red about 
the nodes, 2-4 mm. thick ; leaves opposite, lance-ovate, rather membranaceous, 
acute to obtuse at base, acute to acuminate at apex, subentire, bright green 
and glabrous above, lighter and glabrous beneath except for fine appressed 
puberulence along midrib and veins, the principal lateral veins about 15, 
rather prominent beneath, the submarginal inconspicuous; blades 6-10 cm. 



Vol. XXV] MUNZ: TEE GENUS FUCHSIA 63 

long, 2.5-4 cm. wide ; petioles 5-12 mm. long, purplish red, somewhat strigu- 
lose; stipules not seen; flowers several, in very short compact terminal and 
lateral almost glabrous racemes ; bracts reflexed, linear-lanceolate, 3-10 mm. 
long; pedicels filiform, quite glabrous, 1-2.5 cm. long; ovary linear-fusiform, 
glabrous, 3.5-4 mm. long, red ; hypanthium bright red, 14-20 mm. long, ca. 1 
mm. wide at base, then somewhat narrowed for one-third its length, then 
gradually ampliate until 2.5-3.5 mm. wide at summit, glabrous without, pilose 
within lower portion ; sepals scarlet, divergent, 7-9 mm. long, 3 mm. wide at 
base, subulate at apex for 1-1.5 mm., glabrous; petals scarlet, oblong, 6 mm. 
long, about 3 mm. wide, rounded-obtuse at apex ; stamens 5 and 3.5 mm, long ; 
style glabrous, barely equalling petals; stigma ca. 1 mm. thick; fniit oblong- 
ellipsoid, about 2.5 cm. long. 

Frutex vel arbor ; ramulis glabris, subrubris, 2-4 mm. diametro ; f oliis oppositis, lanceo- 
lato-ovatis, membranaceis, basi acutis vel obtusis, apice acutis vel acuminatis, subintegris, 
in superficie superiore glabris, inferiore in venis strigulosis; laminis 6-10 cm. longis, 2.5-4 
cm. latis ; petiolis 5-12 mm. longis, substrigulosis ; racemis brevibus, compactis, terminalibus 
et lateralibus, subglabris ; bracteis reflexis, lineari-lanceolatis, 3-10 mm. longis ; pedicellis 
glabris, 1-2.5 cm. longis, ovario lineare-fusiforme, glabro, 3.5-4 mm, longo, rubro; hypan- 
thio rubro, 14^20 mm, longo, basi 1 mm, lato, inde angustiore, inde ampliato et apice 
2.5-3.5 mm. lato, externe glabro, interne piloso ; sepalis coccineis, divergentibus, 7-9 mm. 
loiigis, basi 1 mm. latis, glabris ; petalis coccineis, oblongis, 6 mm. longis, 3 mm. latis, apice 
rotundatis vel subobtusis; staminibus 5 et 3.5 mm. latis; style glabro, petala non excedent- 
ibus ; stigmate 1 mm. diametro ; bacca oblongo-ellipsoidea, circa 25 mm, longa. 

Tyije: Comissaria del Putumayo: Cocoa, Colombia, May 23, 1935, H. 
Garcia B. 4639, U. S. Nat. Herb. 1593482. This is rather a fragmentary speci- 
men and had almost finished flowering, so that the last late flowers may be 
somewhat small for the species. A second specimen, Plailada de Minchoy, 
Cuatrecasas 11436 (US). It seems near F. Lehmannii in having lateral and 
terminal compact racemes, being glabrous, having subulate sepal-tips and 
glabrous style. But it has more veins, too long a berry, too long pedicels and 
too short a hypanthium, 

(57) Fuchsia Hartwegii Bentham 

(Plate9, fig, 53) 
Fuchsia Hartwegii Bentham, PI. Hartw,, 179, 1945. 

Shrub or small tree, the younger branches slender, more or less purplish- 
red, pubescent, sometimes also with some longer hairs ; leaves in 3's or 4's, 
elliptic-oblong, rounded to obtuse at base, obtuse to abruptly acuminate at 
apex, subentire or inconspicuously and remotely denticulate, membranaceous, 
darker green and appressed-pilose above, paler and pilose beneath especially 
on veins, principal lateral veins 14-16 on each side of midrib, submarginal 
vein present; leaf-blades mostly 4-10 cm. long, 2-4 (5.5) cm. wide; petioles 
appressed-pilose, slender, 5-35 mm. long; stipules subulate-deltoid, 1.5 mm. 
long, deciduous ; flowers numerous in terminal pyramidal leafy -bracted, 
pendulous panicle 5-15 cm. long; bracts broadly elliptic, 8-25 mm. long; 



64 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

pedicels slender, pilose, 5-10 mm. long; ovary ©void, appressed-pubeseent, 
about 3 mm. long; hypanthium red, tubular, sparsely pubescent without, 
pilose within lower half, 1.4-2 cm. long, 1.5 mm. wide at base, then somewhat 
narrowed, then gradually widened until 4—5 mm. wide at apex ; sepals orange- 
red or scarlet, linear-lanceolate, spreading-divaricate, 8-9 mm. long, barely 
2 mm. wide, acuminate, with subulate tips less than 1 mm. long, pubescent on 
backs; petals red, linear-lanceolate, 8-9 mm. long, 2 mm. wide, acute, gla- 
brous; stamens 10 and 8 mm. long; anthers 1.5-2 mm long; style slender, 
glabrous or somewhat pubescent in lower part, equalling or exceeding sepals 
by 3^ mm ; stigma globose, about 1 mm. broad ; fruit round-ellipsoid, up to 
7 mm. long. 

Type locality, Colombia : "In sylvis prope Pitayo et Huambia." A species 
of southern Colombia, where it grows at between 2000 and 3000 m., it seems 
near F. Lelimanmi, F. ovalis, etc., in its general features: terminal inflores- 
cence, short hypanthium, etc., but has its own characteristic pubescence, leaf- 
size, pedicel-length, and the like. Representative material is as follows, 
COLOMBIA: Caldas: Dawe 741 (NY, US). Huila: "Balsillas" in Rio Bal- 
sillas, Rushy and Pennell 789 (NY, US). El Cauca: near Pitayo and Huam- 
bia, ffarfwegr 994, type no. (fragment F), (photo Berlin specimen F, POM) ; 
central Andes of Popayan, Lehmann 5613 (F) ; Coconuco, Killip 6829 (GH, 
NY, US) ; near Purace, Lelimaym 3503 (F, US) ; San Isidro, Purace, Pennell 
and Killip 6460 (GH) ; Canaan, Mt. Purace, Pennell and Killip 6502 (GH) ; 
del paramo a Purace, matorrales en Chiquin, Perez Arheldez and Cuatrecasas 

5973 (US). 

(58) Fuchsia ovalis Ruiz and Pavon 

(Plate 10, fig. 54) 

Fuchsia ovalis Euiz and Pav6n, F1. Peruv., 3 :87, pi. 324, fig. a, 1802. 

F. polyanthella Johnston, Contr. Gray Herb., 75:36, 1925; I have seen type material of 
both this and ovalis. 

Erect or somewhat scandent, semishrubby, up to 1 m. tall, rather sparsely 
branched, the younger branches 3-6 mm. thick, more or less dark, pilose to 
almost glabrous ; leaves opposite or ternate, membranaceous, broadly elliptic 
to oblong-ovate or obovate, obtuse to acute at base, abruptly acuminate at 
apex, entire to inconspicuously denticulate, darker green and hirsute-strigose 
above, paler and hirsute beneath especially on veins, with about 15-17 prin- 
cipal lateral veins on each side of midrib and with submarginal vein ; blades 
mostly 8-15 cm. long, 5-8 cm. wide; petioles pilose, 2-5 cm. long; stipules 
subulate-deltoid, 2-5 mm. long, quite persistent; flowers in mostly axillary 
racemes up to 1 dm. long ; bracts lanceolate, 1-1.5 cm. long, pilose ; pedicels 
1-1.5 cm. long, glabrous to pilose ; ovary ellipsoid, 5-7 mm. long, glabrous to 
sparingly pilose; hypanthium 13-15 mm. long, scarlet, 1.5-2 mm. wide at 
base, then narrower until gradually widened to summit where 3.5-4.5 mm. 
wide, glabrous to sparingly pilose without, villous within lower half ; sepals 
lanceolate, acute, somewhat divergent, 9-11 mm. long, acute, not subulate, 



1 



Vol. XXV] MUNZ : THE. GENUS FUCHSIA 65 

glabrous to somewhat pilose ; petals obovate-oblong, obtusish, 7-8 mm. long ; 
stamens 5 and 4 mm. long, filaments violet; anthers 1 mm. long ; style glabrous, 
extending to sepal-tips ; stigma subglobose, about 1 mm. broad ; berry oblong, 
red-purple, somewhat 4-angled, about 1 cm. long, 4 mm. wide. 

Type locality, Muna, Huanuco, Peru. The species is characterized by its 
large leaves which are pilose on both surfaces, by its lateral racemes, lance- 
olate bracts, and hypanthium 15 mm. long. I have seen the following collec- 
tions, PERU: the Ruiz and Pavon collection from Madrid Herb. (F), from 
Geneva Herb, (photo F, POM). Huanuco: Huallaga und Muiia, 2800-2900 
m., Weherbauer 6721 (F) ; Muiia, trail to Tambo de Vaca, 2600 m., Machride 
4290, type number polyanthella (F, GH). 

(59) Fuchsia asperifolia Krause 

Fuchsia asperifolia Krause, Fedde Rep. Nov. Sp., 1 :169, 1905. 

Low shrub, up to 5 dm. tall, simple or few-branched, the young twigs dark, 
densely pilose; leaves opposite or ternate, membranaceous, broadly elliptic 
to elliptic-obovate, acute at both ends, subentire, hirsute on both surfaces, 
especially on veins beneath, with about 15 principal lateral veins on each side 
of midrib, submarginal vein present; blades 4-10 cm. long, 2-5 cm. wide; 
petioles hirsute, 2-5 cm. long ; stipules subulate, deciduous ; flowers in dense 
usually terminal hirsute raceme ; bracts apparently lanceolate ; pedicels ap- 
parently scarcely 5 mm. long; ovary about 6 mm. long, pilose; hypanthium 
dark red, 2-2.5 cm. long, slightly bulbous at base, then narrowed and gradu- 
ally widened until 5-6 mm. wide at apex, hirsute without, pilose within; 
sepals ovate, acuminate, 7-8 mm. long ; petals scarlet, narrowly obovate, acut- 
ish at apex, 8-9 mm. long; stamens included; style equaling sepals; stigma 
subglobose, 4-lobed ; fruit not seen. 

Type locality, Peru : Amazonas : between Tambos Bagazan and Almirante, 
east of Chachapoyas, at 2700 m. ; type Weherhauer 4445 at Berlin (photo F, 
POM) . Having seen only a photograph, I do not have many details; the species 
seems near to F. ovalis, but with more terminal inflorescence, is more hirsute 
throughout and with shorter pedicels, 

(60) Fuchsia pilosa Fielding and Gardner 

Fuchsia pilosa Field, and Gardner, Sert. PI., 1, pi. 27, 1844; Macbride, Field Mus. Nat. 
Hist., Bot. 13, pt. 4, no. 1 :561, 1941. 

Shrub, thinly white-pilose on young branches, leaves, petioles, hypanthia, 
etc. ; leaves ternate, oblong-lanceolate, acute at base, acuminate at apex, min- 
utely denticulate, the blades 6-9 cm. long, 1.8-2 cm. wide, one of each whorl 
somewhat larger than the other two ; two of petioles at each node 5-7 mm. 
long, the third at least 2 cm. long ; flowers in terminal racemes which become 
1 or more dm. long; bracts like reduced leaves and 1-2.5 cm. long; pedicels a 
few mm. long ; flowers scarlet, crowded ; hypanthium apparently about 22 
mm. long; sepals oblong, 8-9 mm. long, prominently subulate at apex; petals 



66 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

obtuse, wider than sepals, almost equaling them ; style included ; fruit oblong, 
4-angled, less than 1 cm. long. 

Type locality, Taulia, Amazonas, Peru: type Mathews 1482, which I have 
not seen. Macbride separates this species on the basis of its unequal petioles 
and leaves, one at each node being larger than the other two. It would seem 
from the description to be near to F. asperifoUa and the two are perhaps dis- 
tinct from each other because of the unequal leaves. 

(61) Fuchsia glaberrima Johnston 
 (Plate 10, fig. 55) 
Fuchsia glaberrima Johnston, Contr. Gray Herb., 75:32, 1925. 

Shrub ; younger twigs slightly reddish, 3-5 mm. thick, not branched, gla- 
brous except for very minute scarce appressed puberulence near tips ; leaves 
opposite, firm, elliptic-oblanceolate, tinged reddish purple, obtuse at base, 
acuminate to acute at apex, entire, quite glabrous, with 13-15 principal lateral 
veins on each side of the midrib and with inconspicuous marginal vein ; blades 
10-20 cm. long, 4-8 cm. wide; petioles up to 5 (8) mm. long; stipules connate, 
often reflexed, quite persistent, 2-2.5 mm. long ; inflorescence a sliort terminal 
raceme ; bracts lanceolate, 1-3.5 cm. long ; pedicels stout, 5-7 mm. long, min- 
utely appressed puberulent; ovary fusiform, 6-9 mm. long; hypanthium ' 
bright red, 2.5-3 cm. long, about 2 mm. wide at base, then very little con- 
tracted, then widened until 6-7 mm. wide at apex, almost glabrous without, 
densely pubescent throughout within ; sepals scarlet, obong, 9-11 mm. ong, 
4-5 mm. wide, obtusish ; petals scarlet, broadly obovate, 9 mm. long, rounded 
at apex; stamens 5 and 3.5 mm. long; anthers 1.5 mm. long; style densely 
pubescent much of its length, almost equaling sepals ; stigma 4-lobed, 2.5 mm. 
thick ; fruit not seen. 

Type, ECUADOR: Prov. Tungurahua: Valley of Pastaza River, between 
Banos and Cashurco, 1300-1800 m.; type number Hitchcock 21750 (GH, NY, 
US). Diels, Bibl. Bot. 116 :119, 1937, refers here a Raimondi collection from 
Amazonas, Peru. This species is like F. spectahilis in its broad petals, but has 
a shorter hypanthium; like F. asperifoUa in general appearance, but more 
glabrous and with more sessile leaves. 

(62) Fuchsia macrophylla Johnston 
(Plate 10, fig. 56) 
Fuchsia macrophylla Johnston, Contr. Gray. Herb., 75 : 35, 1925. 

Shrub up to 3 m. tall, openly branched, the younger twigs 3-6 mm. thick, 
with some dark red or purplish tinge, subglabrous to finely strigulose-puberu- 
lent toward the tips ; leaves opposite, thick-membranaceous, elliptic, gradually 
attenuate and subacuminate at base, somcAvhat abruptly acuminate at apex, 
subentire, dark green above, paler beneath, subglabrous on both surfaces 
except for some minute puberulence along the veins, principal lateral veins 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 67 

14-18 on each side of midrib, submarginal well developed; blades 7-15 (18) 
cm. long, 3-5 (6.5) cm. wide; petioles 1-4 cm. long; stipules dark, lance- 
subiilate, 1.5-2 mm. long, deciduous ; inflorescence mostly of short axillary 
sometimes terminal racemes, often only a few cm. long and with few flowers ; 
bracts lanceolate or wider, 5-15 mm. long ; pedicels slender, minutely strigu- 
lose, 5-20 mm. long; ovary ellipsoid, 4—5 mm. long ; hypanthium scarlet, 18-23 
mm. long, 1.5-2 mm. wide at base, then narrowed, then gradually ampliate 
until 4-5 mm. wide at apex, finely appressed-puberulent without, pilose 
within ; sepals divergent, oblong-lanceolate, red with green tips, 7-8 mm. long, 
3 mm. wide, appressed-puberulent on backs, apices subulate for 0.5 mm. ; 
petals bright red, oblong-oblanceolate, 8-9 mm. long, somewhat mucronate at 
apex; stamens 6-8 and 3-5 mm. long; anthers 1.5 mm. long; style pubescent 
near base, scarcely extending to sepal tips; stigma 4-lobed, 1.5 mm. thick; 
fruit dark red, subglobose to somewhat elongate, 8-10 mm. long. 

Type locality. Hacienda Schunke, La Merced, Dept. Junin, Peru. This 
species is near F. Andrei in leaf -size, puberulence, etc., but has more narrow 
and attenuate-based leaves, longer fruit, and more southern range. I have 
seen, from PERU : Huanuco : Distrito Churubamba, trail Puente to Exito, 
Pampa Hermosa, 1750 m., Mexia 8142 (F, GH, NY, US) . Junin : La Merced, 
Maclride 5616, type no. (F, GH, US) ; Pichis Trail, 1600 m., Killip and Smith 
25436 (F, GH, NY) ; Eneiias, Killip and Smith 25784 (GH, NY, US). Aya- 
cucho : Ccarrapa, between Huanta and Rio Apurimac, 1500 m., Killip and 
Smith 22353 (F, GH, NY), 22336 (NY, US). Cuzco: Prov. Paucartambo: 
Rio Tambomayo, bet. Tambomayo and Consuelo, 1100-1400 m.. West 7113 
(GH). 

(63) Fuchsia sessilifolia Bentham 

(Plate 11, fig. 57) 
Fuchsia sessilifolia Bentham, PI. Hartweg., 176, 1845. 

Slender, erect shrubs to small trees, the ultimate branchlets purplish red, 
2-5 mm. thick, glabrous except for the minutely strigulose-puberulent 
younger parts ; leaves rather remote, in whorls of 3 or 4, elliptic-lanceolate to 
narrowly elliptic-ovate or -obovate, rounded to obtuse at base, acuminate at 
apex, remotely serrulate, rather coriaceous, glossy green and quite glabrous 
above, as well as minutely puberulent on the impressed veins, glabrous and 
paler beneath, or puberulent on the veins, principal lateral veins 15-20 on 
each side of midrib, submarginal vein evident ; blades 8-15 cm. long, 2.5-4 
(5) cm. wide; petioles 2-4 mm. long, strigulose-puberulent ; stipules subulate- 
deltoid, about 1 mm. long ; inflorescence terminal, drooping, racemose-panicu- 
late, 1-2 dm. long, canescent-strigose, conspicuously bracted ; bracts mostty 
1.5-2.5 cm. long, ovate-elliptic; pedicels strigulose, slender, 3-5 mm. long^ 
ovary oblong, strigulose, 4—6 mm. long; hypanthium scarlet, 13-16 mm. long, 
1.5 mm. wide at base, then very little narrowed, then gradually ampliate until 
4-5 mm. wide at summit, finely strigulose-puberulent without, pilose within 



68 CALIF OMNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

most of its length; sepals lanceolate, acuminate, divergent, greenish red, 
strigulose on backs, 8-10 mm. long, subulate for almost 1 mm. at tips; petals 
scarlet, oblong, 9-10 mm. long, 3 mm. wide, acutish, glabrous ; stamens ca. 7 
and 5 mm. long ; anthers 1 mm. long ; style glabrous, extending almost to ends 
of sepals; stigma about 1 mm. thick; berry oblong, 10-12 mm. long, 5-6 mm. 
thick, somewhat 4-angled. 

Type locality, "in sylvis Guayan in declivitate occidentali montis Pichin- 
cha," Ecuador. The species seems to me to be near F. Hartwegii in inflores- 
cence, hypanthium, etc., but differs in the long, subsessile, narrow, rather 
glabrous, firm leaves, short pedicels, conspicuous inflorescence. It grows in 
southern Colombia and northern Ecuador at altitudes of from 2000-3600 
meters. Representative material is, COLOMBIA. Antioquia : Dauro, Choco, 
near Medellin, Toro 1168 (NY). Caldas: Buenavista-Quindio, Andre in 1876 
(F, GH, NY). Tolima: Prov. de Mariquita; El Gallego, Triana 3813 (US), 
Holton in 1853 (NY). Huila: Cordillera Central, east of Neiva, Rushy and 
Pennell 687 (NY, US) ; Cordillera Oriental, vertiente occidental, entre Gabi- 
nete y Andalucia, Cuatrecasas 8586 (US). Cauca : Valle de Quintero, near 
Pitaio, Pittier 1427 (NY, US) ; Canaan, near Mt. Purace, Pennell and Killip 
6674 (GH, NY, US); Paletara to Calaguala, Peniiell 7111 (GH, NY, US). 
Nariiio : woods near Pasto, Jameson 432 (US). Putumayo : above El Encavo, 
Laguna La Cocha, Balls 7521 (US). ECUADOR: Carchi: Canton Tulcan 
near Pun, Mexia 7597 (POM, US). Pichincha : sive Andium Quitensium, 
Jameson under Couthouy (GH). 

(64) Fuchsia sylvatica Bentham 

(Plate 11, fig. 58) 

Fuchsia sylvatica Bentham, PI. Hartweg., 176, 1845. 

F. nigricans Linden, F1. de Series, Ser. 1, 5 :pl. 481, 1849; see discussion below. 
F. atrorubra Johnston, Contr. Gray Herb., 75 :31, 1925 ; I cannot maintain, although sepals 
and petals are somewhat short. 

Low shrub, more or less scandent, openly branched, the ultimate twigs pur- 
plish, minutely canescent strigulose, 2-A mm. thick ; leaves opposite to ternate, 
elliptic to elliptic-ovate or -obovate, obtuse to acute at both ends or sub- 
acuminate at apex ( subentire or inconspicuously denticulate, membranaceous, 
deep green and strigulose above, paler and more densely strigulose beneatli, 
veins reddish, the principal lateral ones 11-15 on each side of midrib, sub- 
marginal present; leaf -blades 3-10 cm. long, 1.5-5 (6) em. wide; petioles 
strigulose, 6-20 mm. long; stipules subulate-deltoid, 1-1.5 mm. long, some- 
w^hat persistent ; flowers on terminal or lateral pendant strigulose racemes ; 
bracts ovate to oblong to suborbicular, 1-2.5 cm. long ; pedicels slender, strigu- 
lose-canescent, 5-9 mm. long; ovary narrowly ellipsoid, 4-6 mm. long; hypan- 
thium 14-19 mm. long, ca. 1.5 mm. thick at base, gradually ampliate until 
2.5-3 mm. wide at summit, more or less canescent-strigulose without, densely 
pilose within; sepals pink or pale red, divergent, linear-lanceolate, 6-9 mm. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 69 

long, acute, not subulate at apex, strigulose on backs ; petals crimson to purple 
red, oblong, 7-9 mm. long, acute; stamens 4-6 and 3-5 mm. long; anthers 
1-1.5 mm. long; style glabrous, scarcely reaching sepal-tips ; stigma 4-lobed, 
2 mm. wide; berry oblong, 12-14 mm. long, 4.5-6 mm. thick. 

Type locality, "In sylvis Guayan in declivitate occidentali montis Pichin- 
eha," Ecuador. It is near F. sessilifolia in puberulence and short pedicels, but 
has broader leaves and more pubescent twigs. I have not seen the type of 
F. nigricans from Merida, Venezuela, but the three Jahn specimens seem to 
agree with the description. Though these come from much farther north than 
the general range of F. sylvatica, I cannot separate them unless I see the 
labels. Perhaps these Venezuela specimens have slightly looser pubescence 
and fruit about 2 mm. longer. Representative material of sylvatica, VENE- 
ZUELA: Merida: Paramo de Arieagua, 2500 m., Jahn 1028 (US) ; between 
Bocono and Campo Elias, 2000 m., Jahn 14 (US) ; selva arriba de Palmira, 
2600 m., Jahn 509 (US) . COLOMBIA : Antioquia : Santa Elena, Archer 1195 
(US) ; Medellin, Toro 972 (NY). Caldas : Rio Santa Rita, Salento, Killip and 
Hazen 8966 (GH, NY, US); Salento, Pennell and Hazen 10144 (GH, NY, 
US) ; Quindio, Dawe 769 (NY, US) ; Santa Elena, above Santuario, Pennell 
103 13, type no. atrorulra (GH, NY). Cundimarca : Fomeque, Dawe 555 (US). 
Tolima: La Mediacion, Quindio, Triana 3511 (US) ; Rio Toche to Machin, 
old Quindio Trail, Killip and Hazen 9564 (GH). ECUADOR: Pichincha: 
w. slope Mt. Pichincha, Jameson 190 (GH), Hartweg 984, Geneva specimen 
US) ; between Cosanga and Baeza, 2000 m., Mexia 7337 (US) ; Rio Mapoto, 
(photo F, POM). Tungurahua : Canton Quito : Alaspongo, Mexia 7703 (POM, 
Penland and Summers 282 (F, POM). 

Section 3. KIERSCHLEGERIA (Spach) Munz, new comb. 

Kierschlegeria, as genus, Spach, Nouv, Ann. Sci. nat., (2), 4:176, 1835; as subsect., 
Endlicher, Gen. PL, 1193, 1840. 

Pedicels axillary, pendulous ; seeds angled. Leaves sparse or subverticillate ; 
thickened petiole-base persistent and becoming spine-like. Sepals reflexed, 
little shorter than hypanthium. 

One species, Fuchsia lycioides Andr. 

(65) Fuchsia lycioides Andrews 

(Platell, fig. 59) 

Fuchsia lycioides Andr., Bot. Eep., II, pi. 120, 1800; Bot. Mag., 25:1024, 1807. 

F. rosea Etnz and Pavon, F1. Peruv., 3 :88, 1802. 

F. spinosa Presl, Eel. Haenk., 2 :26, pi. 51, 1835. 

F. rosea var. spinosa Eeiche, Anal. Univ. Chile, 98:486, 1897; PL de Chile, 2:268, 1898. 

Shrub up to 3 m. tall, the older branches stout, grayish, knobby with dwarf 
lateral branches, intricately branched and with some of the twigs dull-spinose 
at their ends, younger twigs more slender (2-3 mm. thick), fairly straight, 
reddish, subglabrous with short sparse pubescence especially toward tips. 



70 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

beset after the leaves are shed by the persistent thickened woody bases of the 
petioles which form heavy dull spines 1.5-2 mm. long; leaves alternate, nu- 
merous, the blades lance-ovate to ovate, 6-24 mm. long, 3-15 mm. wide, sub- 
entire, acute to obtuse at both ends, glabrous except sometimes at tips and 
margins, somewhat paler beneath; petioles slender above the thickened base, 
glabrous to pubescent, 5-15 mm. long ; stipules apparently fused with petiole 
base ; flowers axillary, solitary ; pedicels filiform, 5-8 mm. long, glabrous to 
puberulent; ovary rounded, 1.5-2 mm. long; hypanthium glabrous without 
and within, red, subcylindric, 4-7 mm. long, 2-2.5 mm. wide ; sepals 4, red, 
lanceolate, acuminate, spreading-reflexed, glabrous or somewhat pubescent, 
-4-7 mm. long ; petals purplish, obcordate, 3^ mm. long ; episepalous stamens 
about as long as petals, epipetalous somewhat shorter; style glabrous, reddish, 
about as long as sepals ; stigma somewhat 4-lobed, about 1 mm. wide ; berry 
blue-black, subspheric, 4-6 mm. thick. 

Type locality given as "North-west coast of America," but undoubtedly 
Valparaiso, Chile, near w^hieh most specimens of the species have been col- 
lected. It apparently grows on dry slopes near the sea, mostly in the provinces 
of Coquimbo and Aconcagua. Representative material, CHILE : Fray Jorge, 
Skottsherg 798 (NY), 3Iu7ioz P. no. B-217 (GH), Werdermann 1526 (B), 
Berninger 621 (B) ; Punta de Teatinos, ^Yest 3921 (GH) ; Coquimbo, Gaudi- 
chaud 77 (B, G). Santiago: Santiago, Germain in 1856-57 (G) ; Concon, 
Poeppig 123 (B). Aconcagua : Valparaiso, Macrae in 1825 (G), Bertero 1006 
(G, GH), Gaudickaud 248 (F, G),Rushtj 1800 (F, NY, US) ; Laguna Verde, 
near Valparaiso, Looser 2290 (POM), Garaventa 1641 (GH); Choapa, near 
Los Vilos, Mr. and Mrs. J. N. Rose 19280 (NY, US) ; Papudo, Looser 2291 
(POM). 

Section 4. SKINNERA (Forst.) DeCandolIe 

STcinnera (FoRST.) DC, Prodr., 3:39, 1828; Endlicher, Gen. PI., 1183, 1840; Benth. and 
Hook., Gen PI., 1, pt. 2:791, 1862; Eaimann in Engl, and Prantl, Die nat. Pflanzenfam., 
Ill, 7:219,1893. 
as genus, Forst., Char. Gen., 57, pi. 29, 1776. 

Hypanthium infundibuliform, swollen above ovary. Sepals reflexed. Petals 
small and scale-like or wanting. Filaments long, exserted. Fruit many-seeded ; 
seeds very small, much crowded, in 2 or more rows. Stigma subcapitate, some- 
what 4-lobed. Pedicels axillary, pendulous. Leaves alternate. 

Type species, Fuchsia excorticata (Forst.) L.f. 

Key to Species of Section SKINNEEA 
A. Petals 2-4 mm. long; plant very woody, varying from a liane to an erect shrub or tree; 
leaf -blades whitish beneath, 1-9 cm. long 
B. Trees; leaf -blades 2-9 cm. long, vnt\\ about 10 principal veins on each side of 
midrib. 
C. Leaves oblong-ovate to lance-ovate, acute to acuminate at apex, somewhat 
pilose beneath on midrib. New Zealand 66. F. excorticata. 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 71 

CC. Leaves elliptic to elliptic-obovate, rounded to obtuse at apex, glabrous beneath. 

Tahiti 67. i^. cyrtandroides. 

BB. Not trees; leaf -blades 1-4 cm. long, with 3-9 main veins. 

C. Erect shrub ; leaves oblong-ovate, blades mostly 2-4 cm. long. . . 68. F. Colensoi, 

CC. Liane ; leaves suborbicular, blades mostly 1-2 cm. long 69. F. perscandens. 

AA. Petals lacking ; plant small, procumbent, wth very slender stems ; leaf -blades less white 
beneath, 0.5-1.8 cm. long. 
B. Sepals acuminate; style about as long as stamens; hypanthium tubular-campan- 

ulate, 7-8 mm. long 70. F. procumbens. 

BB. Sepals obtuse ; style not exserted from hypanthium ; hypanthium campanulate, 
6-7 mm. long 71. F. KirTcii. 

(66) Fuchsia excorticata (Forster) L.f. 

(Plate 11, fig. 60) 

SMnhera excorticata FoRST., Char. Gen., 58, pi. 29, 1776. 

Fuchsia excorticata (Forst.) L. f., Suppl., 217, 1781; Lindley, Bot. Eeg., 10:857, 1824. 

Wide-spreading tree 10 m. tall, with papery exfoliating light brown bark ; 
young twigs subgiabrous to puberulent, subterete, more or less reddish in 
color; leaves alternate, oblong-ovate to lance-ovate, rounded to subeordate at 
base, acute to acuminate at apex, usually somewhat gland-denticulate, green 
and almost glabrous above, whitish and generally glabrous beneath except 
for some pilosity along veins and margins ; blades 3-9 cm. long, 1.5-3 cm. 
wide, with about 10 principal veins on each side of midrib and with sub- 
marginal vein ; petioles puberulent, 1-1.5 (4) cm. long; stipules ovate, acumi- 
nate, puberulent, less than 1 mm. long, deciduous; flowers solitary, axillary, 
pendant, at first green, then purplish red ; pedicels capillary, puberulent, 
5-15 mm. long; ovary subgiabrous, 5-6 mm. long, about 1 mm. thick; hypan- 
thium 4-angled, 8-15 mm. long, subgiabrous without and within, 2-5 mm. 
wide at very base for 1—2 mm., then 1.5-2 mm. wide and gradually ampliate 
until 4-10 mm, wide at summit; sepals spreading to reflexed, ovate-lanceolate, 
glabrous, acuminate, 8-14 mm. long ; petals dark purple, erect, 2-4 mm. long, 
elliptic, lanceolate, acutish ; stamens erect, the episepalous 6-9 mm. long, the 
others somewhat shorter; anthers blue, 1-2 mm. long; style glabrous, about 
equalling sepals ; stigma subentire, 2 mm. long, yellowish ; berry ellipsoid, 
about 1 cm. long, dark. 

Type locality not given; species ranging through New Zealand. Representa- 
tive material, NEW ZEALAND : North Island : Auckland : Waikowhai, Miss 
Mackie in 1932 (NY) ; Auckland, Petrie 4110, in 1898 (NY), Meehold 9925 
(NY). Wellington : Feilding, Allan in 1924 (NY), in 1928 (CAS) ; Wainui- 
o-Mata, Meehold 4619 (BISH) ; Khandallah, Self in 1941 (POM, WELT). 
South Island : without definite locality, Kirk (F). Otago : Dunedin town-belt, 
Colby, DN, from Herh. D. Petrie (GH) ; cliff west of St. Clair, Prue Collier, 
per M. Neal, 168 (BISH) ; Nelson, Kirk 253 (GH), Marie Neal 3 (BISH) ; 
Canterbury : Kaiturea Valley, Bank's Peninsula, Cockayne 4111 (NY) ; So. 
Canterbury, edge of forest on Hunter Hills, Anderson 207 (F, NY) . 



72 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

(67) Fuchsia cyrtandroides Moore 

(Plate 11, fig. 61) 
Fuchsia cyrtandroides Moore, B. P., Bishop Museum, Occasional Papers, 16:13, 1940. 

Tree up to 5 m. tall, the young twigs quite glabrous; leaves mostly alternate, 
broadly elliptic to elliptic-obovate, green above, white beneath, glabrous, 
rounded to obtuse at base and apex, subentire to obscurely denticulate, with 
about 10 principal veins on each side of midrib and with indefinite sub- 
marginal vein; petioles 8-14 mm. long; blades 2-9 cm. long, 2-5.5 cm. wide; 
stipules apparently ovate, deciduous; flowers solitary, axillary, rose-magenta ; 
pedicels 1.5-2.5 cm. long ; ovary fusiform-clavate, 6-7 mm. long, 2 mm, thick ; 
hypanthium about 1 cm. long, tubular-campanulate, glabrous, 2-2.5 mm. wide 
at base and gradually ampliate to 5-6 mm. at summit; sepals spreading, 
lanceolate, subacuminate, 10-14 mm. long, 4-5 mm. wide; petals erect, oblong- 
ovate, 4 mm. long ? ; stamens about as long as sepals, the epipetalous slightly 
shorter; style slightly exceeding stamens, glabrous; stigma 1.5 mm. long, 
somewhat 4-lobed; berry black, ellipsoid, about 1 cm. long. 

SOCIETY ISLANDS: Tahiti: Orofena, south side, rain forest, 1500 m.. 
May 14, 1927, MacDaniels 1315, type coll. (BH, BISH), at 1600 m., 8t. John 
and Fosherg 17005 (BISH). The species is very near F. excorticata in its 
arborescent habit, large leaves which are white beneath, and in the size and 
shape of the flowers. It differs from that New Zealand species, however, in 
having the leaves proportionately wider, more obtuse and less pubescent. 

(68) Fuchsia Colensoi Hook. f. 

(Plate 12, fig. 62) 

Fuchsia Colensoi Hook, f., Handb. New Zealand FL, 728, 1867; Trans, and Proc. New Zea- 
land Inst., 1892 : 25, pi. 19, fig. 4, 5, 1893. 

Small branching erect shrub, with ultimate twigs very slender, puberulent ; 
leaves oblong-ovate, rounded to subcordate at base, obtuse to subacuminate at 
apex, subentire, green and glabrous above except for fine strigulosity on 
impressed veins, white and glabrous beneatli except for some pilosity on main 
veins, short-ciliate, principal lateral veins 6-9 on each side of midrib ; petioles 
very slender, 5-10 (20) mm. long ; blades 1-4 cm. long, 0.5-2 cm. wide; pedicels 
filiform, 5-10 mm. long ; ovary ellipsoid, about 5 mm.long, 2-2.5 mm. wide ; 
hypanthium 8-10 mm. long, 2-2.5 mm. wide at base, then 1-1.5 mm. wide and 
rapidly ampliate until 5-7 mm., red, quite glabrous; sepals greenish to reddish, 
spreading-reflexed, lanceolate, 7-15 mm. long ; petals purple, 2-3 mm. long ; 
stamens almost equaling sepals or about two-thirds as long, epipetalous ; style 
exserted by 2-5 mm.; stigma 1.5-2 mm. wide; berry oblong, at least 8 mm. 
long. 

Type locality. North Island, New Zealand. Material seen, NEW ZEA- 
LAND : North Island : Papaitonga, in Kahikatea forest, Oliver in 1941 (POI\I, 
WELT) ; Wammometa, Wellington, collector not given (WELT, photo 



Vol. XXV] MUNZ : THE GEXUS FUCHSIA 73 

POM) ; Levin, Wellington, Oliver in 1941 (POM, WELT). South Island: 
Kaikoura, Kirk (GH) ; Pelorus, Kirk, (F) ; Otago : Port Chalmers, Petrie 
4109 (NY). This species is very near to F. excorticata and not surely distinct. 
With inadequate material I am following what seems to be a tendency among 
New Zealand botanists, namely to recognize these several Fuchsia segregates 
as species. 

(69) Fuchsia perscandens Cockayne and Allan 
(Plate 12, fig. 63) 
Fuchsia perscandens Cockayne and Allan, Trans. Proc. N. Z. Inst., 57:53, 1927. 

Climbing shrub, the main stems few-branched, the twigs appressed-puberu- 
lent, slender ; leaves orbicular-ovate, subcordate at base, acute at apex, sub- 
entire to sinuate-denticulate, subglabrous above except for possible minute 
puberulence along the impressed midrib, whiter and subglabrous beneath, 
short-ciliate, with about 3-6 main veins on each side of midrib ; blades 1-2.5 
cm. long, almost as wide; petioles almost filiform, 1-2 cm. long; pedicels fili- 
form, about 1 cm. long ; flower much as in F. Colensoi, the hypanthium about 
9 mm. long, inflated at base, etc. ; sepals about 7 mm. long ; petals 2.5 mm. long; 
berry dark purple, subcylindric, about 1 cm. long, 5 mm. thick. 

Type locality, "forests near Feilding," North Island, New Zealand. Material 

seen, NEW ZEALAND: North Island: Palmerston North, Zolro 3471 

(CAS) ; Feilding, forest margin, H. H. Allan, Aug. 1924 (NY); Sinclair 

Head, Wellington Harb., Bealy 1 and 2 (POM, WELT). A second sheet from 

Feilding also at New York is an intergrade with F. excorticata, with longer 

and more oblong leaves which are pilose beneath on the veins; it may be a 

hybrid. 

(70) Fuchsia procumbens R. Cunningham 

(Plate 12, fig. 64) 
Fuchsia procumbens E. Cunn., Ann. Nat. Hist., 3 :31, 1839. 
F. prostrata Baill., Bull. Soc. Linn., Par. 1:270, 1880; apparently a name for the above. 

Procumbent, woody, with very slender glabrous stems; leaves alternate, 
not crowded; petioles very slender, channeled and pubescent above, 1.5-3 cm. 
long ; stipules minute, ovate, deciduous ; blades suborbicular to round-ovate, 
subcordate at base, serrulate, subglabrous except on pubescent margin and 
veins of lower surface, somewhat paler beneath than above, 1-1.8 cm. long, 
sometimes somewhat wider, with about 5 principal veins on each side of mid- 
rib ; flowers solitary, axillary; pedicels slender, subglabrous, scarcely 1 cm. 
long ; ovary 3^ mm. long, fusif orm-clavate ; hypanthium apparently dark 
red, tubular-campanulate, 7-8 mm. long, constricted above ovary, then 
abruptly dilated to about 3 mm. wide, then more gradually until 5 mm. wide 
at summit, glabrous without and within ; sepals reflexed, green, lance-deltoid, 
acuminate, 6 mm. long, 2.5 mm. wide; petals lacking; stamens slightly un- 
equal, erect, ca. 4 and 5 mm. long; anthers 1 mm. or slightly longer; style 
glabrous, about as long as stamens, purplish; stigma 1.5-2 mm. thick; fruit 
not seen. 



74 CALIFOEXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Type locality, North Island, New Zealand, "around the village of ]\Iatauri 
on the east coast opposite the Cavallos Isles." Material seen, NEW ZEA- 
LAND : North Island, garden specimen, Khandallah, Sntherland in 1941 
(POM). South Island: Canterbury: Christchurch, Laing's Garden, Meehold 
25160 (BISH) ; Timaru, park gardens, Anderson 35 (NY). 

(71) Fuchsia Kirkii Hooker f . 

Fuchsia Kirkii Hook. f. ex T. Kirk, Trans. New Zealand Inst., 1 :92, 1868; Icon. PL, 11 :66, 
pi. 1083, 1871. 

"With prostrate habit and slender stems of F. procumhens ; petioles 1-1.5 
cm. long, blades 5-10 mm. long, orbicular-cordate; pedicels ca. 5 mm. long; 
ovary ca. 3 mm. long ; hypanthium campanulate, 6-7 mm. long, 5-6 mm. wide 
at summit ; sepals reflexed, oblong, obtuse, 5 mm. long, 2.5 mm. wide ; petals 
0; stamens 3-4 mm. long; style included in hypanthium ; stigma less than 0.5 
mm. wide. 

Type locality. Great Barrier Island, New Zealand. Specimens seen, NEW 
ZEALAND : North Island : Tryphena Bay, T. Kirk (F, GH) ; Port Tryphena, 
Kirk 528 (GH). Apparently distinct from F. procumhens in having a shorter 
wider hypanthium, sepals more oblong and obtuse ; style shorter and stigma 
smaller. 

Section 5. HEMSLE YELLA Munz, new section 

Nectary and hypanthium-base fused ; petals lacking or nearly so. Hypan- 
thium funnelform to cylindro-funnelform, enlarged at base. Sepals more or 
less connate at base. Leaves tending to be deciduous at anthesis. 

Nectarium basem hypanthii adnatum; petalis fere nullis; hypanthio infundibuliforme 
vel cylindro-infundibuliforme, base ampliato; sepalis base fere connatis. 

Type species. Fuchsia apetala Ruiz and Pavon. This section is dedicated to 
W. B. Hemsley who published a paper on the apetalous Fuchsias of South 
America in the Journ. of Bot. 14 :69-70, 1876. 

Key to Species of Section HEMSLEYELLA 

A. Hypanthium less than 2 cm. long; leaves with 13-16 principal lateral veins on each side 

of midrib 83. F. cestroides. 

AA. Hypanthium more than 2.5 cm. long; leaves with 5-10 principal lateral veins on each 
side of midrib. 
B. Hypanthium at least 4-5 times as long as the sepals (including the connate base of 
the latter). 

C. Leaves well developed at anthesis; petioles mostly 3-7 cm. long; pedicels 3-5 

cm. long; hypanthium 10-14 cm. long 81. F. Garlepiriana. 

CC. Leaves mostly deciduous at anthesis; petioles mostly 1-3 em. long; pedicels 

0.5-2.5 cm. long; hypanthium 6-12 cm. long 82. F. maciantha. 

BB. Hypanthium usually 1.5-3 times as long as sepals (including connate base). 
C. Hypanthium glabrous without. 

D. Leaves alternate, lance-ovate, the blades mostly 2.5-4 times as long as 
wide; hypanthium 8-12 mm. wide; sepals greenish 73. F. tuherosa. 

I 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 75 

DD. Leaves mostly subopposite, elliptic- or oblong-ovate, the blades not more 
than twice as long as wide ; hypanthium 5-8 mm. wide ; sepals red. 

E. Petals 2 mm. long ; sepals 8-12 mm. long. Mexico 72. F. decidua. 

EE. Petals none; sepals 18-22 mm. long. South America. 

F. L eaves largely deciduous at an thesis ; stamens exceeding sepals ; 
style 1-1.5 cm. longer than sepals. Peru to Bolivia. 

74. F. juntasensis. 

FE. Leaves well developed during anthesis ; stamens shorter than 
sepals ; style scarcely extending beyond sepals. Venezuela. 

75. F. memhranacea. 
CC. Hypanthium not glabrous under a lens. 

D. Leaves lance-ovate ; twigs glabrous ; hypanthium sparsely and finely 

glandular-pubescent 76. F. salicifolia. 

DD. Leaves elliptic- to oblong-ovate ; twigs not glabrous. 

E. Leaves subopposite ; pubescence very fine, short, velvety ; hypanthium 

5-8 mm. wide. Largely Bolivian 77. F. tunariensis. 

EE. Leaves alternate; pubescence longer; hypanthium 7-11 mm. wide. 

F. Stamens longer than sepals; pedicels 1.5-2.5 cm. long; inflores- 
cence short-pubescent. Venezuela to northern Peru. 

78. F. apetala. 

FF. Stamens shorter than sepals ; pedicels 1-1.5 cm. long ; inflores- 
cence shaggy-pilose. Central Peru to Bolivia. 

G. Leaves deciduous at anthesis; hairs white, 0.5-1 mm. long. 

79. F. liirsuta. 
GG. Leaves present at anthesis; hairs brown, 1-1.5 mm. long. 

80. F. unduavensis. 

(72) Fuchsia decidua Standley 
(Platel2, fig. 65) . 
Fuchsia decidua Standley, Pub. Field Mus. Nat. Hist., Bot. 4:248, 1929. 

Shrub with long snbterete or somewhat 4-angled. branches, the j^ounger ones 
3-9 mm. thick, brownish, glabrous, with internodes 1-3 cm. long; leaves oppo- 
site, unknown, deciduous at time of flowering ; inflorescence short racemose 
lateral panicles 2-6 cm. long, with glabrous, somewhat glaucous very slender 
branches; pedicels slender, recurved, 3-8 mm. long; hypanthium 2-3.5 cm. 
long, almost 2 mm. wide at base, then narrowed into an elongate tube which 
gradually flares upward until 5-6 mm. wide at mouth, somewhat glaucous, 
light vermilion, glabrous without and within ; sepals vermilion, oblong, ob- 
tuse, 8-12 mm. long, 4 mm. wide ; petals suborbicular, 2 mm. long ; stamens 
about 9 and 6 mm. long; filaments thickish; anthers 1.5-1.8 mm. long; style 
glabrous, about equal to sepals; stigma barely 1 mm. long; berry not seen, 
but probably longer than thick. 

Type locality, MEXICO : Jalisco : Real Alto, La Bufa, Sierra Madre Occi- 
dental, at 2500 m.; Mexia 1601 being the type no. (F, GH, CAS, MICH, NY, 
US). A second collection of apparently the same thing comes from the state 
of Guerrero; Sierra Madre near Chilpanengo (Chilpancingo?), Nelson 2195a 
(US) ; none of the flowers is fully matured, hence shorter than in the type 
collection. Not knowing the leaves, I find it difficult to determine a close rela- 



76 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

tionship for this species in the section Eufiuchsia, but it seems to be near to the 
apetalous species of Hemsleyella in its greatly reduced petals. Some of these 
species have also lateral reduced inflorescences. 

(73) Fuchsia tuberosa Krause 

Fuchsia Ulcerosa Krause, Fedde Rep., 1 :170, 1905. 

Shrub up to 1 m. tall, epiphytic or terrestrial; roots bearing clusters of 
roundish tubers; stems few-branched, reddish, glabrous, the branchlets 2-5 
mm. thick; leaves may shed at time of flowering, alternate, lance-ovate to 
lanceolate or ovate, subglabrous, round or acute, sometimes cordate at base, 
curved-acuminate at apex, entire to denticulate, paler beneath than above, 
with 6-8 principal lateral veins on each side of midrib ; blades 5-10 cm. long, 
2-3.5 cm. wide; petioles slender, 1-2 (3) cm. long; stipules deltoid, about 0.4 
mm. long, deciduous ; flowers in upper axils, sometimes crowded terminally ; 
pedicels slender, 2-6 cm. long, glabrous or nearly so; ovary cylindric, glabrous, 
8-12 mm. long, 1-2 mm. thick ; hypanthium red, 3.5-8 cm. long, 3-^ mm. wide 
at base, then narrowed to 1.5-2 mm. for less than 1 cm., then abruptly widened 
to 8-12 mm. for the rest of its length, glabrous without, villous within at base ; 
sepals green, erect-divergent, lance-deltoid, 12-20 mm. long, 5-8 mm. wide, 
acute, connate for one-fourth their length ; petals 0; episepalous stamens about 
as long as sepals, alternate ones three-fourths as long, yellow ; anthers 3-4 mm. 
long, yellow; style slender, pubescent near base, exserted beyond sepals for 
1-5 cm.; stigma elongate-capitate, 2-3 mm. long; fruit glabrous, 2-2.3 cm. 
long. 

Key to Varieties of Fuchsia tuberosa 

A. Hypanthium 3.5-5 cm. long 73a. var. typica. 

AA. Hypanthium 6-8 cm. long 73b. var. inflata. 

(73a) Fuchsia tuberosa Krause, var. typica Munz, new name 

(Plate 12, fig. 66) 

Fuchsia tuierosa Krause, Fedde Rep., 1 :170, 1905. 

Fuchsia chloroloha Johnston, Journ. Arnold Arboretum, 20:243, 1939. 

Hypanthium 3.5-5 cm. long. 

Type locality, between Sandia and Cuyocuyo, Puno, Peru, at 2700-2800 m., 
Dept. Cuzco: "Pillahuata," Cerro de Cusilluyoc, 2200-2400 m., Pennell 
based on Weherhauer 875 at Berlin (photo F, POM). Material seen, PERU : 
13973, type no. chloroloha (F, GH) ; Marcapata Valley, 1200 m., Herrera 1166 
(US) ; between Achirani and Medias-Mayu, 2700 m., Vargas 11116 (F) ; Rio 
Tambomayo, from Pillahuata bridge to head of Tambomayo Grade, 1600- 
2300 m., West 7094 (GH); between Tanamayo and Tambomayo, 2000 m., 
Vargas 70/7094 (F). The Krause description does not mention green sepals, 
as does Johnston's, but the region, the leaf- and flower-shape, glabrous hypan- 
thium, etc., certainly make ticberosa and chloroloha the same. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 77 

(73b) Fuchsia tuberosa Krause var. inflata (Schulze-Menz) Munz, new comb. 
Fuchsia inflata Schulze-Menz, Notizbl. Bot. Gart., Berlin, 15 : 136, 1940. 

Hypanthium 6-8 cm. long. 

Type from Cuzeo : prov Paucartambo, between the tambos Tres Cruces and 
Tambomayo, above Cosiiipata, at 3100 m., April, 1914, Weherhauer 6935 (F, 
GH, POM, US). This does not seem specifically distinct, and more material 
may show that it is not worth any recognition. 

(74) Fuchsia juntasensis 0. Kuntze 
(Plate 12, fig. 67) 

Fuchsia juntasensis O. Kuntze, Kev. Gen. 3, part 2 :97, 1898. 

F. Mattoana Krause, Engl. Jahrb., 37:599, 1906. 

F. Steinhachii Johnston, Contr. Gray Herb., 75:38, 1925. 

Vinelike, usually epiphytic shrub, 5 or more dm. tall, few-branched, the 
younger twigs usually purplish, subglabrous or sparsely pubescent, the older 
ones with exfoliating outer tissues ; leaves mostly subopposite, tending to fall 
during anthesis; the blades elliptic- to oblong-ovate, acute to obtuse at base, 
acuminate at apex ,subentire, 4-8 cm. long, 1.5-3.5 cm. wide, sparsely short- 
pubescent to giabrescent above and beneath, paler beneath than above, prin- 
cipal lateral veins 6-8 on each side of midrib ; petioles sparsely pubescent, 
5-12 mm. long, ca. 1 mm. wide; stipules subulate, 1-1.5 mm. long, caducous; 
flowers few, crowded in terminal racemose clusters, often on short lateral 
twigs ; pedicels slender, subglabrous, 1.5-5 cm. long ; ovary oblong-fusiform, 
puberulent to subglabrous, 5-10 mm. long, 2-2.5 mm. wide ; hypanthium rose 
to flesh-colored, tubular-funnelf orm, 2.5-4.5 cm. long, 2-2.5 mm. AVide at base, 
then slightly narrowed for one-third its length, then gradually ampliate until 
5-7 mm. wide at summit, glabrous without, pilose within near base ; sepals 4, 
reddish, ovate-lanceolate, 18-22 mm. long, 5-6 mm. wide, glabrous, sub- 
acuminate, connate for 2-3 mm. at base ; petals none ; episepalous stamens 
20-26 mm. long, alternate ones 16-20 mm. long ; anthers cream, 3-4 mm. long; 
style pubescent in lower portion, extending 1-1.5 cm. beyong the. sepals; 
stigma green, oblong-globose, 2-3 mm. long, scarcely lobed ; berry ellipsoid, 
about 12 mm. long. 

Type locality, east side of Cordillera, at 3000 m., between Cochabamba and 
Rio Juntas, Bolivia. I have not seen the type of F. juntasensis so labeled and 
have had some misgiving about taking up this name for this concept, especially 
since Kuntze's description sounds as if the calyx were short, giving sepals as 
12 mm. long, and even if one adds 3-4 mm. for the connate base, which Kuntze 
may well not have included as part of the sepals, the length is still rather 
inadequate. But all other features : glabrous flowers, ovate, acuminate leaves, 
shape of hypanthium, would point to this concept. Of F. Mattoana I have 
seen only the photograph; between it and F. Steinhachii I can see no difference 
except the somewhat shorter hypanthium (25-28 mm.) given by Krau^ 




LISRAR 



78 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

However, a series of specimens seems to break down the validity of this char- 
acter. For the species which I am calling F. jtintasensis I have seen the follow- 
ing collections: without locality Herb. 0. Kuntze (NY). PERU: between 
Cuzco and Santa Ana, 3400 m., Weherhmier 4976, type of Mattoana at Berlin 
(photo F, POM). BOLIVIA : Depto. Cochabamba : Ineachara, 2500 m., Stein- 
hach 5038, type Steinhachii (NY) ; Quebrada de Corani, prov. Chapare, 2400 
m., Steinlach 9862 (F, GH, NY). The species differs from F. apetala in its 
glabrous flowers and more southern range. 

(75) Fuchsia membranacea Hemsley 
Fuchsia membranacea Hemsley, Journ. Bot., 14:70, 1876. 

Shrub, glabrous throughout except ofr the minute pubescence on the upper 
surface of the young leaves which may persist along the veins ; branches few, 
younger ones purplish, 2.5-3.5 mm. thick, older ones exfoliating ; leaves sub- 
opposite, well developed at anthesis, the blades membranaceous, oblong-ovate, 
remotely denticulate, paler beneath, rounded at base, short-acuminate at apex, 
3-5.5 cm. long, 1.5-3.5 cm. wide, with 6-7 principal lateral veins on each side 
of midrib ; petioles puberulent, 6-15 mm. long, scarcely 1 mm. wide ; stipules 
subulate-deltoid, about 0.5 mm. long; flowers axillary ; pedicels slender, 2.5-4 
cm. long; ovary oblong, 5-7 mm. long, 2 mm. thick; hypanthium tubular- 
funnelform, 4-5 cm. long, slightly enlarged and 2.5 mm. wide at base, then 
1.5-2 mm. wide for about 1 cm., then gradually widened in upper half until 
7-9 mm. broad, glabrous without, pubescent within near base ; sepals diver- 
gent, ovate-lanceolate, 18-22 mm. long, 6-7 mm. wide, subacuminate, very 
little connate at base ; petals 0; episepalous stamens 14-20 mm. long, alternate 
ones 9-15 mm. long ; anthers 2-3 mm. ling ; style pubescent in lower portion, 
scarcely extended beyond sepals; stigma oblong-capitate, 2-3 mm. long ; berry 
short-ellipsoid, about 1 cm. long. 

Type locality, Caracas, Venezuela; type based on Linden 372, of which 
number I have seen fragment (F) and photo (F, POM). Other material seen, 
VENEZUELA : state of Trujillo : Paramo de la Cristalina, 2900 m., J aim 12 
(US). Merida: Paramo de Timotes, 3000 m., J aim 508 (GH, US). Like F. 
juntasensis, the species has glabrous flowers, but it is leafy at anthesis and has 
shorter stamens. 

(76) Fuchsia salicifolia Hemsley 

Fuchsia salicifolia Hemsley, Journ. Bot., 14:70, 1876. 

Epiphytic shrub, mostly glabrous except about the flowers ; branchlets few, 
purplish with persistent epidermis ; leaves fully developed during anthesis, 
alternate, lanceolate, obtuse at base, long-acuminate at apex, remotely den- 
ticulate, thickish, with about 10 principal lateral veins on each side of midrib, 
the blades 6-10 cm. long, 2-2.6 cm. wide ; petioles ca. 1 cm. long, 1.5 mm. wide ; 
stipules not seen; flowers few, axillary; pedicels 2-2.5 cm. long, slender, 



Vol. XXV] MUNZ: THE GENUS FUCHSIA 79 

pubescent, with some gland-tipped hairs; ovary linear-oblong, 1 cm. long, 
2-2.5 mm. wide; hypanthium tubular-funnelform, 4-5.5 cm. long, sparsely 
glandular-pubescent without, 2.5 mm .thick at base, then narrower for about 
1 cm., then gradually widened to 7 mm. ; sepals lanceolate, acuminate, spread- 
ing, ca. 3 cm. long, 1 cm. wide, connate at base for 2-3 mm. ; petals ; epi- 
sepalous stamens about 27 mm. long ; alternate ones 20 mm. long ; anthers 3-4 
mm. long; style exserted; stigma ''clavate"; berry about 2 cm. long. 

Type locality, Sandillani ; I am not sure whether this is in southern Peru or 
in Bolivia. The only specimen I have seen which I can refer here is from 
Unduavi, North Yungas, Bolivia at 2450 m.. Bushy 1803 (NY) ; it is a speci- 
men with 1 leaf and 1 flower. The species seems distinct in its narrow leaf, 
subglabrous condition and long sepals. In hypanthium-shape it seems near 
F. apetala. 

(77) Fuchsia tunariensis 0. Kuntze 

(Plate 13, fig. 68) 
Fuchsia tunariensis O. Kuntze, Eev. Gen. PI., 3, part 2 :98, 1898. 

Epiphytic or prostrate to scandent shrub with tuberous roots ; stems up to 
8 dm. long, few-branched, the young growth pallid with densely velvet pubes- 
cence ; twigs often knotty and irregular, purplish, densely pubescent ; leaves 
largely subopposite, tending to be deciduous at anthesis ; blades elliptic- to 
oblong-ovate, 4-15 cm. long, 2-6 cm. wide, membranaceous, densely fine- 
pubescent, velvety when young, entire, rounded to obtuse at base, subacumi- 
nate at apex; petioles 5-20 mm. long, densely pubescent, about 1 mm. wide; 
stipules subulate, gland-like, scarcely 1 mm. long, deciduous ; flowers few, 
solitary in uppermost axils ; pedicels densely pubescent, 1-1.5 cm. long ; ovary 
densely short-pubescent, oblong, 5-8 mm. long ; hypanthium red to pinkish, 
tubular-funnelform, 4.5-5.5 cm. long, densely short-pubescent without, pilose 
within lower portion, 2.5-3 mm. wide at base, then 1.5-2 mm. wide for about 
1.5 cm., then gradually ampliate until 5-8 mm. wide at mouth; sepals red, 
lance-ovate, acute, 18-22 mm. long, connate for about 3 mm. at base ; petals 
0; episepalous stamens 13-18 mm. long, alternate ones 10-13 mm. long; 
anthers about 3 mm. long ; style pilose at base, scarcely exserted or as much as 
1 cm. longer than sepals ; stigma ca. 2 mm. long, subglobose ; fruit hairy, 
ellipsoid, almost 2 cm. long. 

Type locality, "Tunarigebirge," Cochabamba, Bolivia, at 3000 m. The type 
in Herb. Otto Kuntze (NY) is badly broken, the solitary flower has pedicel, 
ovary and lower half of hypanthium, but the pubescence is very character- 
istic. In fact, it is the close, fine, dense pubescence which distinguishes this 
species, of which I have seen the following representation: PERU: Cuzco, 
prov. Paucartambo, below "Pillahuata," 2300 m.. West 7106 (GH) with 
rather large sepals, but typical pubescence. BOLIVIA : Cochabamba : Tunari, 
Kuntze in 1892, type (NY) ; Ayopaya, Sailapata, 3500 m., Cardenas 3393 
(US), 3369 (US). Pongo, Tate 188 (NY). 



80 CALIFORNIA ACADEMY OF SCIENCES [Peoc. 4th Ser. 

(78) Fuchsia apetala Ruiz and Pavon 
(Platel3, fig. 69) 
Fuchsia apetala Ruiz and Pav6n, F1. Peruv., 3 :89, pi. 322, fig. b, 1802. 

F. msig7iis Hemsley, Journ. Bot., 14:69, 1^76; three collections were cited from Ecuador, 
2 by Jameson and 1 by Spruce. The last whicli cannot be taken as the type, is the only one 
I have seen, but I am unable to separate insignis from apetala. 

Vine-like shrub, often epiphytic, up to about 1 m. high, the younger 
branches reddish, more or less pubescent, with some gland-tipped hairs, rather 
coarse, commonly 3-5 mm. thick, with exfoliating epidermis and frequently 
M'ith adventitious roots; leaves largely deciduous at anthesis, alternate, the 
blades elliptic-ovate, subentire, rounded to subcordate at base, acute to 
acuminate at apex, somewhat coriaceous, finely pubescent to glabrescent 
above, villous beneath, especially beneath on midrib and veins, principal 
lateral veins 6-8 on each side of midrib ; leaf -blades 5-9 cm. long, 3-4.5 cm. 
wide; petioles 1.5-2.5 cm. long, somewliat villous, 1.5-2 mm. wide; stipules 
subulate, 1.5-2 mm. long, caducous; flowers few, crowded near apex of branch- 
lets; pedicels slender, 1.5-2.5 cm. long, villous, with some gland-tippecl hairs; 
ovary cylindrical, 8-15 mm. long, villous, 2-A mm. thick ; hypanthium orange- 
scarlet, 4-5.5 cm. long, tubular-funnelform, 3-5 mm. wide at bulbous base, 
then abruptly narrowed to 1.5-3 mm. for about 1 cm., then gradually ampliate 
until 7-11 mm. wide, sparsely glandular-villous without, villous within lower 
portion; sepals reddish-orange, more or less spreading, lance-ovate, 22-25 
mm. long, 8-10 mm. wide, acute to obtusish, pubescent without, connate for 
2-3 mm. at base; petals 0; episepalous stamens 25-30 mm. long, alternate 
18-20 mm. long; anthers green, 3-4 mm. long; style pubescent at base, ex- 
serted 1-2 cm. beyond the sepals ; stigma subglobose, 2-3 mm. long, usually 
scarcely lobed ; fruit oblong, red. 

Type locality ''Habitat copiose in Huassahuassi et Muna," Peru. At Field 
]\Iuseum are fragments of what I here consider to be apetala mixed with F. 
niacrantha Hook., collected by Ruiz and Pavon ; my judgment as to which is 
to be taken as apetala is based on the plate by Ruiz and Pavon. I prefer, since 
this Ruiz and Pavon material in its present condition contains both entities 
and since I believe these are separable on the basis of geographical and 
morphological characters, to keep the name apetala for the Ruiz and Pavon 
plant that was illustrated, rather than to treat all as one thing as does Mac- 
bride (Field Mus. Nat. Hist., Bot. 13 :part 4, no. 1 :546, 1941). I would refer 
the following material to apetala, VENEZUELA : Merida : Paramo de Aric- 
gua, 2000 m., Jahn 1030 (US) ; Paramo de San Jose, 2400 m., Jalin 969 (US) ; 
Paramo de Mucuchachi, 1800 m., Jahn 981 (US). ECUADOR: between 
Cuenca and Huigra, Chimborazo, 2700-3000 m., Hitchcock 21701 (GH, NY, 
US) ; Huigra, Rose and Rose 22230 (GH, NY, US) ; Mt. Azuay, 8pruce 5976 
(GH, NY, photo of Geneva specimen at F, POM). PERU?: Churupullana, 
Domhey ex Herb. Mus. Paris (F). I can see no difference between the Vene- 
zuela and Ecuador specimens in spite of the discontinuous distribution. 



Vol. XXV] MUNZ: TEE GENUS FUCHSIA 81 

(79) Fuchsia hirsuta Hemsley 
(Plate 13, fig. 70) 
Fuchsia hirsuta Hemsley, Journ. Bot., 14 :69, 1876. 

Tuber-bearing, climbing, epiphytic shrub or scanclent over faces of cliffs, 
few-branched, mostly leafless at anthesis, the younger twigs 3-8 mm. thick, 
knotty, tortuose, densely pilose with white hairs not over 1 mm. long, more or 
less purplish underneath the hairs; leaves alternate, ovate to oblong-ovate, 
membranous, subentire to denticulate, rounded at base, acute to almost acumi- 
nate at apex, densely pilose on both surfaces when young, more or less glabres- 
cent except for veins of under surface, principal lateral veins 5-8 on each side 
of midrib ; leaf -blades 6-12 cm. long, 3.5-6 cm. wide ; petioles pilose, 1-4 cm. 
long, 1.5-2 mm. wide; stipules pilose, lanceolate, ca. 1 mm. long; flowers few 
to many, crowded near tips especially of lateral twigs ; pedicels slender, pilose, 
1-1.5 cm. long; ovary cylindric, densely shaggy-pilose and pubescent, 8-12 
mm. long, 1.5-2.5 mm. thick ; hypanthium tubular-funnelform, densely pilose 
and pubescent, with some hairs gland-tipped, fiery to purplish red, 3-5.5 cm. 
long, 2-3 mm. thick at base, then slightly narrowed for one-third its length 
and then rather abruptly widened into the cylindrical upper part which is 
8-10 mm. wide, pilose within the lower portion ; sepals reddish, ovate to lance- 
ovate, obtuse to acute, pilose on backs, 15-20 mm. long, 7-10 mm. wide, con- 
nate only a short distance at base ; petals ; episepalous stamens 15-18 mm. 
long, alternate ones two-thirds as long ; anthers about 3 mm. long ; style pilose 
above the base, extending to as much as 1.5 cm. beyond the sepals; stigma 
about 2 mm. long ; berry narrowly ellipsoid, almost 2 cm. long. 

Type locality, "Peruvia" ; I have not seen the type, LecJiler 1989. The species 
is included in F. apetala by Macbride (Field Mus. Nat. Hist., Bot., 13, part 4, 
no. 1 :546, 1941), but to me it is a more southern entity with short sepals and 
more included stamens, gray-pilose and shaggy ovaries and hypanthia. I have 
seen from PERU: Ayacucho : Punaccuanca Pass, 4100 m.. West 3669 (GH). 
Cuzco : prov. Paucartambo, disto. Marcachea, forest of Pucara, 3400 m., 
Vargas 11170 (F) ; below Tres Cruces, Weherlauer 6975 (F, GH) ; prov. 
Urubamba, disto. Chincheros, Chincheros, 3700 m., Vargas 9602 (F) ; Yucay, 
3000 m., Soukup 715 (F) ; Cuzco, 2500 m., Herrera2189 (GH) ; Saxaihuaman, 
Herrera 2189 (F) ; Combapata, Canchis, 3700 m., Vargas 1900 (GH). Puno, 
Soukup 525 (F), BOLIVIA: La Paz: Ingenio, between Sorata and Mapiri, 
3700 m., Cardenas 1142 (NY) ; La Paz : prov. Larecaja, near Sorata, Mandon 
629 (F, GH) ; Pongo, White 248 (GH, US). 

(80) Fuchsia unduavensis Munz, new species 
(Plate 13, fig. 71) 

Much like F. hirsuta, but with leaves well developed at time of flowering, 
blades elliptic-ovate, 7-10 cm. long, 2-5 cm. wide. Math 6-7 principal lateral 



82 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

veins on each side of midrib ; young stems, ovaries, pedicels, hypanthia with 
spreading, brownish hairs about 1.5 mm. long; hypanthium 4.5^5.5 cm. long, 
7-8 mm. wide in upper part ; sepals 2 cm. long, lance-ovate, acute to obtusish. 

F. hirsutae similis, foliis ad anthesin praesentibus, elliptico-ovatis, laminis 7-10 cm. 
longis, 2-5 cm. latis; caulibus junioribus, ovariis, pedicelis, hypanthiis capillis fuscis et 1.5 
mm. longis vestitis; hypanthio 4.5-5.5 cm. longo, supra 7-8 mm. lato; sepalis 2 cm. longis, 
lanceolato-ovatis, acutis vel subobtusis. 

Type: Unduavi, North Yungas, Bolivia, at 3600 m., Feb. 13, 1907, Buchtien 
2925, United States Nat. Herb. 1177335. Other collections from same place : 
at 3200 m., Nov., 1910, Buchtien (F), at 3600 m., Nov., 1910, Buchtien 2925 

(US). 

(81) Fuchsia Garleppiana Kuntze and Wittmack 

(Plate 14, fig. 72) 
Fuchsia Garleppiana Kuntze and Wittm., Gartenfl., 47 : 461, fig. 96, 1893. 

Tuberous, the thickened roots said to be 20-90 cm. long ; slender-stemmed, 
few-branched, up to 1 m. high, epiphytic or on wet slopes, the ultimate branch- 
lets reddish, soft-pubescent, 2-4 mm. thick, tardily exfoliating ; leaves alter- 
nate, well developed at anthesis, ovate to oblong-ovate, the blades thin, 
rounded to subcordate at base, subacuminate at apex, subentire to remotely 
denticulate, finely pubescent above and beneath, with 8-9 principal veins on 
each side of midrib ; blades 6-10 cm. long, 4-6 cm. wide ; petioles pubescent, 
2.5-7 cm. long, about 1 mm. wide; stipules lanceolate, scarcely 1 mm. long, 
deciduous ; flowers in axils on main growing branches ; pedicels slender, soft- 
pubescent, 3-5 cm. long; ovary 5-10 mm. long at anthesis, soft-pubescent, 2-3 
mm. thick; hypanthium 10-14 cm. long, subeylindric, finely pubescent with- 
out and within, pink, 5-6 mm. thick at base, then narrowed to 2-3 mm. and 
then gradually wider until 8-10 mm. broad below the somewhat contracted 
mouth ;^epals pink, lance-ovate, acute, somewhat divergent, 13-18 mm. long, 
5-7 mm. wide, connate almost one-half their length, fine-pubescent on both 
surfaces; petals 0; episepalous stamens about three-fourths, alternate ones 
about two-thirds the sepals; anthers 2.5-3.5 mm. long; style pubescent near 
the base, slender, scarcely extending beyond the sepals; stigma subglobose, 
2-3 mm. thick ; fruit not seen. 

Type locality, Cocapata, Cochabamba, Bolivia, at 3000 m. in the Tunari 
Mts. The species is near to F. macrantha in its long hypanthium, but differs 
in many respects : flowers not on spur-like branchlets, plant leafy at anthesis, 
petioles longer and more slender, pubescence finer and shorter, sepals hairy 
on both surfaces, filaments pubescent; hypanthium narrowed at end. Material 
seen, PERU: Dept. du Cuzco, Aout., 1847, Weddcll, from Herb. Mus. Paris 
(F). BOLIVIA: Cochabamba: Incachaca, 2700 m., Cardenas 688 (US); 
Incacorral-Aduana, Chapare, 2700 m., Steinbach 9538 (GH, NY) ; Tunarige- 
birge, 3000 m.. Herbarium Otto Kuntze, type coll. (F). 



Vol. XXV] MUNZ : THE GEXUS FUCHSIA 83 

(82) Fuchsia macrantha Hooker 
(Platel4, fig. 73) 
Fuchsia macrantha Hook., Bot. Mag., 72 :pl. 4233, 1846. 

Low, trailing or straggling, or epiphytic, few-branched, the twigs 2.5-6 
mm. thick, pubescent to villous, with some hairs gland-tipped, freely ex- 
foliating; leaves largely lacking at anthesis, alternate, elliptic-ovate, acute, 
subentire, rounded or subcordate at base, pubescent above, pilose beneath 
especially on veins and when young, with 6-7 principal lateral veins on each 
side of midrib, blades 4-9 cm. long, 3-6 cm. wide ; petioles 1-2 (3.5) cm. long, 
more or less pilose, 1-2 mm. wide ; stipules lance-deltoid, 0.5 mm. long, decidu- 
ous ; flowers crowded on short branchlets ; pedicels slender, pubescent, 1-2 cm. 
long; ovary linear, pubescent, 6-12 mm. long, 1.5-3.5 mm. thick; hypanthium 
rose-red. to coral or scarlet, subcylindric to cylindro-fusiform, 6-12 cm. long, 
3-8 mm. wide at base, then narrower and very gradually ampliate until 5-12 
mm. wide below the mouth, 8-ribbed, finely pubescent, with some hairs gland- 
tipped, villous within near base ; sepals ovate to lance-ovate, divergent, 1.5-2.5 
cm. long, 0.7-1.2 cm. wide, tips greenish yellow, somewhat pubescent on both 
surfaces, obtusish, connate about half their length; petals 0; episepalous 
stamens about three-fourths the sepals, alternate ones slightly shorter; fila- 
ments glabrous ; anthers yellow, 2.5-4.5 mm. long ; style pubescent just above 
the ovary, slender, exserted 5-20 mm. beyond the calyx; stigma conico- 
spherical, greenish, 2-A mm. long, scarcely lobed. 

Type locality, Cundimarca, Peru. Material seen, PERU: locality not given, 
Pavon 1826, as apetala, Geneva specimen (photo F, POM) ; Peru, mixed with 
apetala, Buiz and Pavon, Madrid Herb. (F). Huanuco: Villcabamba, haci- 
enda on Rio Chinchao, 1800 m., Machride 4970 (F, GH) ; Tambo de Vaca, 
4000 m., Machride 4901 (F, GH, US). Apurimac : prov. de Abancay, bosques 
del Ampay, 3200-3500 m., C. Vargas C. 1000 (GH). Dept. not known : Lucu- 
mayo Valley, 1800-3600 m.. Cook and Gilbert 1312 (US). Machride 4970 and 
Vargas 1000 tend to have the hypanthium somewhat fusiform and for a time 
seemed to me worth recognizing as a separate entity, but the material is 
insufficient. 

(83) Fuchsia cestroides Schulze-Menz 

(Plate 14, fig. 74) 

Fuchsia cestroides Schulzb-Menz, Notizbl. Bot. Gart., Berlin, 15:317, 1940; Macbride, 
Field Mus. Nat. Hist., Bot. 13, part 4, no. 1 :550, 1941. 

Erect shrub, 3 m. tall, the young twigs densely puberulent to subpilose, 
1.5-3 mm. thick, somewhat reddish in color; leaves present at anthesis, mostly 
ternate, not crowded, firm-membranaceous, elliptic-obovate to oblong-ovate, 
obtuse to acute at base, acuminate to acute at apex, plainly and rather closely 
serrulate, subglabrous to hispidulous-puberulent above, paler and short- 
pilose beneath especially on the veins, principal lateral veins 14-16 on each 



84 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

side of midrib, submarginal scarcely developed ; leaf -blades 4-8 (10) em. long, 
2-3.5 (4) cm. wide ; petioles 3-6 mm. long, pubescent ; stipules subulate, often 
reflexed, about 1 mm. long, deciduous; flowers numerous in short lateral 
racemes or terminal panicles; bracts lanceolate to linear, pilose, 3-15 mm. 
long; pedicels pilose, slender, 3-5 mm. long; ovary fusiform, pilose, 2-A mm. 
long; hj-pantliium dark red, 17-18 mm. long, 1-1.5 mm. wide at base, then 
somewhat narrowed until gradually widened to 4 mm. at summit, pilose with- 
out, glabrous within ; sepals lanceolate, 6-7 mm. long, 2.5 mm. wide, pilose on 
backs, somewhat divergent, acute; petals 0; stamens 7-8 and 5-6 mm. long, 
exserted; anthers 1 mm. long; style glabrous, exceeding sepals by 6-7 mm. ; 
stigma narrow, about 1 mm. long ; berry apparently ellipsoid, at least 6 mm. 
long. 

Type locality, below Frias, prov. Ayabaca, Piura, Peru, at 1100-1200 m., 
the type collection, Weherhauer 6423 (F, GH, US). This species is not at all 
close to the other apetalous ones, and probably should not be placed with 
them. It differs in its many veins, smaller flowers, etc., and suggests in general 
appearance F. Lehmannii. 

Section 6. SCHUFIA (Spach) Munz new section 

Schufia, as genus, Spach, Ann. Sci. Nat., (2), 4:177, 1835. 
Schufia, as subsection, Endlicher, Gen. PI., 1193, 1840. 

Flowers erect in a terminal cymose panicle, arranged subtrichotomously. 
Leaves opposite or verticillate. Nectary fused with base of hypanthium. 
One species, F. arhorescens Sims. 

(84) Fuchsia aborescens Sims 
Fuchsia arborescens Sims, Curtis Bot. Mag., 53:2620, 1826. 

Low shrub to small tree up to 8 m. tall, sometimes epiphytic, glabrous or 
nearly so ; branchlets, petioles and veins sometimes tinged with red ; leaves 
opposite or ternate, oblong-oblanceolate to oblong-elliptic, acute or acuminate 
at both ends, paler beneath than above, entire to serrulate, blades 3-20 cm. 
long, 1-7 cm. wide, with 9-13 principal lateral veins; flowers numerous, 
crowded in corymbose panicles 5-25 cm. long and equally wide, lower branches 
of inflorescence with few reduced leaves, upper floriferous ones with much 
reduced, deciduous, subulate bracts; pedicels reddish, slender, erect, 5-15 
(25) mm. long ; hypanthium tubular to funnelform, rose to magenta, 3-5 mm. 
long, 1-3 mm. wide at summit in pressed specimens ; sepals reddish to wine- 
purple, linear to lance-oblong, not apiculate in bud, 3-7 mm. long, 1-2.5 mm. 
wide, spreading-reflexed at anthesis; petals lavender or lilac, lance-ovate, 
two-thirds as long as sepals, spreading; episepalous stamens exceeding sepals, 
4-8 mm. long, epipetalous two-thirds as long; filaments pinkish lavender, 
anthers purplish, 1-1.6 mm. long; style just longer than stamens, filiform, 
glabrous ; stigma purplish, conic, about 0.5 mm. long ; berry purplish, covered 
with blue bloom, subglobose, 7-10 mm. thick. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 85 

Key to Forms of Fuchsia arborescens 

A. Sepals 4.5-7 mm. long; liypanthium 4.5-8 mm. long. 

B. Sepals 1.5-2.5 mm. wide; liy2)antliium 2-3 mm. wide 84a. f. typica. 

BB. Sepals 1-1.5 mm. Avide; hypanthium 1 mm. wide 84b. f. tenuis. 

A A. Sepals 3-4.5 mm. long ; hypanthium 3-4 mm. long , 84c. f . parva. 

(84a) Fuchsia arborescens Sims forma typica new name 
(Plate 14, fig. 75) 

F. arborescens Sims, Curtis Bot. Mag., 2G20, 1826; Lindl., Bot. Eeg., 11:943, 1826. 

Schufia arborescens Spach, Ann. Sci. Nat., (2), 4:177, 1835. 

F. hamellioides Moc. and SessiS ex. G. Don, Gen. Syst., 2 :677, 1832, in synon. Have seen no 

material. 
F. syringae flora Care., Eev. Hortic, 1873 : 311, 1873 ; judging from plate. 
F. arborea SESst and Moc, PI. N. Hispan., ed. 1, 58, 1887-'90 ; type seen. 
F. arborescens var. (?) megalantha Donnell Smith, Bot. Gaz., 18:2, 1893. Type seen; 

agrees Avell in broad sepals, etc., with Sims' plate. 
F. Liebmanni L6vL., Bull. Georgr. Bot., 22 :24, 1912. Photograph of type seen. 

Hypanthium 4.5-8 mm. long, 2-3 mm. wide at summit ; sepals 4.5-7 mm. 
long, mostly 2-2.5 mm. wide. 

Type locality, Mexico. Ranging at altitudes of 1000-2500 m., from Morelos 
and Guerrero to Panama ; the most common form of the species. In this treat- 
ment of F. arborescens, forms rather than varieties are being recognized, 
since there is no geographical correlation and the intergradation is so com- 
plete. Representative material of f . typica is, MEXICO, without definite local- 
ity, '^F. arliorca," Sesse, Mocino, Castillo y Maldonado 5216 (F). Morelos, 
mts. above Cuernavaca, Pringle 6825 (F, GH, MICH, NY, POM, US). Guer- 
rero, Sierra Madre del Sur, Distrito Mina, 2d ridge west of Petlacala, Mexia 
9052 (GH, NY) . Vera Cruz, Xalapa, Galeotti 3035 (NY) . Puebla, St. Vincent, 
Nicolas 11 (F, NY) ; Puebla, Bro. Arscne in 1909 (US) . Oaxaca, Sierra de San 
Felipe, Pringle 6242 (F, NY, GH, CAS, POM, US) ; Barranca de San Luis, 
Conzatti 1538 (US). GUATEMALA: Alta Vera Paz, near San Jose, s.e. of 
Tactic, Standley 69660 (F), Tactic, von Tuerckheim under J. D. Smith 8395 
(F, NY, US). Quezaltenango, Volcan de Santa Maria, Steyermark 33616 
(F) ; Cumbre de Tuilacan, Standley 67808 (F) ; Volcan de Zunil, Skutch 877 
(F). Chimaltenango, Chichavac Tecpan, Skutch 255 (US) ; Chichavac, Skutch 
59 (US) ; Las Calderas, Standley 60074 (F). Sacatepequez, Volcan de Agua, 
Maxon and Hay 3745 (US), Standley 65066 (F) ; Volcan de Acatenango, 
J. D. Smith 2469, type no. var. megalaiitha (GH, US). COSTA RICA: Ala- 
juela, Volcan de Poas, Cufodonti 601 (F). San Jose, La Hondura, Standley 
36587 (US); between Guayabillos and Cabeza de Vaca, Dodge and Thomas 
4936 (F, MICH, US) ; La Palma, Tonduz under J. D. Smith 7410 (F, GH, 
US). Cartago, Volcan de Turrialba, Pittier under J. D. Smith 7508 (F, US) ; 
Volcan de Irazu, Oersted, type F. Liebmannii at Copenhagen (photographs 
F, POM). PANAMA : Chiriqui, el Boquete, Pittier 2974 (US) ; Casita Alta, 
Volcan de Chiriqui, Woodson, Allen and Seihert 797 (MO, POM). 



86 CALIFOEXIA ACADEMY OF SCIEXCES [Proc. 4th Ser. 

(84b) Fuchsia arborescens Sims forma tenuis new form 

Hypanthium tubular, 5-6 mm. long, 1 mm. wide; sepals linear, 5-6 mm. 
long, 1-1.5 mm. wide. 

Hypanthium tubulosum, 5-6 mm. longum, 1 mm. latum ; sepalis linearibus, 5-6 mm. longis, 
1-1.5 mm. latis. 

Type : Vara Blanca de Sarapiqui, north slope of Central Cordillera, be- 
tween Poas and Barba Volcanoes, Costa Rica, alt. 1740 m., Oct., 1937, A. F. 
Skutch 3357 (US 1643234); isotype (NY). With about some range as f. 
typica. Representative material, MEXICO : Michoacan and Guerrero, Sierra 
Madre, Langlasse 872 (F, GH, US). Orizaba, Botteri (F, GH). Chiapas, San 
Cristobal,^. W. Nelson 3236a (US); Mt. Pasitar, Matuda 1695 (MICH). 
GUATEMALA: Alta Vera Paz, Coban, von Tuerckhcim II 713 (F, US). 
Quezaltenango : Fuentes Georginas, w. slope Volcan de Zunil, Standley 67493 
(F). Solola, Volcan de Atitlan, Hatch and Wilson in 1936 (F). Chimalte- 
nango : Chichavac, Skutch 750 (F). PANAMA : Chiriqui : Casita Alta, Volcan 
de Chiriqui, Woodson, Allen and Seihert 911 (MO, POM); El Boquete, 
Maxon 5015 (NY, GH, US). COLOMBIA: Dept. Boyaca : Labranzgrande, 
1150 m., B. Guevara Amortegui 389, in 1932 (F, US). Since this is the only 
collection I have seen from south of Panama, I cannot help wonder whether 
it represents an introduction or not. 

Intergrades with f . typica are numerous ; among such may be cited, Oaxaca : 
Coyula a Cuyamecalco, Cancim 2438 (US) ; Chiapas, Mt. Orlando, Matuda 
0437 (F, MICH, US) ; Vera Paz in Guatemala: Chucaneb, /. D. Smith 182 
(F, GH, NY, US) ; Huehuetenango, Yalambofroch, Seler 2869 (GH, US) ; 
San Jose in Costa Rica : Between Aserri and Tarbaca, Standley 34100 (US). 

(84c) Fuchsia arborescens Sims forma parva Munz, new form 

F. paniculata Lindl., Gard. Chron., 1856:301, 1856. Type not seen, but the "much smaller 
flowers" of the description would probably mean this. 

HjT)anthium funnelform, 3-4 mm. long, 1.5 mm. wide; sepals 3-4 mm. 
long, 1 mm. wide. 

Hypanthium 3-4 mm. longum, 1.5 mm. latum ; sepalis 3-4 mm. longis, 1 mm. latis. 

Type : Finca Pireneos, below Santa Maria de Jesus, Dept. Quezaltenango, 
Guatemala, March 11, 1939, Standley 68287, Pomona College Herb. No. 
254648; isotype Field Museum 988800. With much the same range as the other 
forms; for example, MEXICO : Oaxaca, Tonaguia, Galcotti 3035 (US). Vera 
Cruz, Jalapa, Walters (NY) ; Barranca de Tenampa, Zaeuapan, Purpus 7647 
(NY, US). Chiapas, Volcan de Tacana, Matuda 2924 (MICH) ; Cerro del 
Boqueron, Purpus 6970 (NY, US). GAUTEMALA: Quezaltenango, Fuentes 
Georginas, Volcan de Zunil, Standley 67356 (F, POM) ; Pirenos above San 
Felipe, Standley 68287 (F, POM). Quiche, San Miguel Uspantan, Heyde and 
Lux under J. D. Smith 3000 (US) . Alta Vera Paz : Coban, von Tuerckheim in 
1908 (NY). Sacatepequez, Volcan de Agua, Kellerman in 1905 (US). Zacapa, 



1L 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 87 

Sierra cle las Minas, Steyermark 29996 (F). Chiquimula, Montana Norte to 
El Jntal, Steyermark 31042 (F). COSTA RICA: Alajuela, La Yentolera, 
Volcan de Poas, Standley 34580 (US). PANAMA: Chiriqui, Rio Chiriqui 
Viejo, Allen 1351 (MO, NY, GH, POM) , Vi^liUe 179 (MO, POM) . 

Intergrades with f. tenuis are, MEXICO, Chiapas, Mt. Pasitar, Matuda 
1695 (MICH, NY). COSTA RICA : San Jose, Laguna de la Chonta, Standley 
42260 (US) ; Cartago, Volcan de Irazu, La Esperanza, Standley 35374 (US). 

Section 7. ENCLIANDRA (Zucc.) Endlicher 

EncUandra (Zucc.) Endlicher, Gen PL, 1183, 1840; Benth. and Hook., Gen. PL, 1, part 

3:791,1862. 
As genus, Zuccarini, Abh. Akad. Muenchen, 11:335, 1837; spelled Eucliandra, Steud., 

Nom.ed., 2, 1:600, 1840. 
Brebissonia Spach, Hist. Nat. Veg. Phan., 4:401, 1835; Ann. Sci. Nat., (2), 4:175, 1835; 

as subsect., Endlicher, Lc. 
Lyciopsis Spach, I.e., 176; as subsect., Endlicher, I.e. 
Myrinia Lilja, FL sverig. Vexter., Suppl. 1 :25, 1840. 

Flowers polygamous. Petals flat. Stamens short, equal, the epipetalous re- 
flexed into the hypanthium. Berry few-seeded. Slender shrubs with opposite 
leaves, small solitary axillary flowers. 

Type species. Fuchsia EncUandra Steud. 

Key to Species of Section ENCLIANDEA 
A. Hypanthium obconic; sepals more than half as long as hypanthium, generally reflexed 
at anthesis. 
B. Leaf -blades mostly less than 2 em. long, rounded to obtuse at base, obtuse to acute 
at apex ; petioles not more than 1 em. long. 

C. Hypanthium 4-6 mm. long ; sepals 4-5 mm. long 85. F. thymifolia. 

CO. Hypanthium 1-3 mm. long; sepals 1.5-3 mm. long. 

D. Sepal-tips eonnivent in bud ; hypanthium 2-3 mm. long. 

E. Petals 2-3 mm. long; sepals 2-3 mm. long. Hidalgo to Miehoacan. 

86. F. Pringlei. 
' EE, Petals 1 mm. long ; sepals 1.5 mm. long, Volcan de Colima. 

87. F. colimae. 
DD. Sepal-tips spreading in bud; hypanthium 1-2 mm. long; petals 1-1.5 mm, 

long. Mexico to Chiapas 88, i^, minimiflora. 

BB. Leaf -blades mostly 2-5 (7) em. long, acute at base, subaeuminate at apex; petioles 
1-3 cm. long. 
C. Hypanthium longer than wide; stamens included; young growth subglabrous 

to pubescent 89. i^. tacanensis. 

CC. Hypanthium not longer than wide in pressed flowers ; stamens evident ; young 

growth fine-strigulose 90. F. STcutchiana. 

AA. Hypanthium subcylindric ; sepals usually not more than half the hypanthium, ascending 
to somewhat spreading at anthesis. 
B. Leaves subeoriaceous, frequently conspicuously serrulate. 

C, Plant quite glabrous ; hypanthium 5-6 mm. long, 1,5-2,5 mm. wide at summit ; 

sepals 3-5 mm. long. Guerrero to Chiapas 91. F. iacillaris. 

CC. Plant puberulent on young twigs, hypanthia, etc. 

D. Leaf-blades 3-8 (10) mm. long; hypanthium 3-4 mm. long; sepals 2-2.5 
mm. long ; petals 1.5-2 mm. long. Hidalgo, Vera Cruz, Mexico, 

92. F. minutiflora. 



88 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

DD. Leaf -blades 7-20 or more mm. long. 

E. Hypanthium 5-9 (11) mm. long; sepals 2.5-5 mm. long. 

F. Ilypantliium 7-9 (11) mm. long, 2.5-5 (6) mm. wide in the 

perfect flowers. Hidalgo to Guatemala 93. F. microphylla. 

FF. Hypanthium 5-7 (9) mm. long, 1-2 mm. wide. Costa Eica to 

Panama 94. F. Hemsleyana. 

EE. Hypanthium 3-4 mm. long, 1 mm. wide; sepals 2 mm. long. Chiapas. 

95. F. striolata. 
BB. Leaves membranaceous, often subentire. 

C. Leaf -blades mostly less than 3 cm. long, usually pilose beneath ; sepals 2-3 
mm. long; petals 0.6-2 mm. long. Michoacan to Costa Eica. 

96. F. michoacanensis. 
CC. Leaf-blades not pilose beneath, or if so, larger. 

D. The leaf-blades 8-15 (20) mm. long; hypanthium of staminate flowers 

6-8 mm. long, 2-3 mm. wide. Hidalgo to Oaxaca 97. F. Encliandra. 

DD. Leaf -blades 20-60 (100) mm. long; hypanthium of staminate flowers 
mostly 8-10 mm. long, 2.5-3.5 mm. wide. 
E. Hypanthium of pistillate flowers 1-3 mm. wide; petals red; leaves 
pubescent or pilose beneath. 
F. Petioles 3-10 (12) mm. long; branchlets pubescent; leaves sub- 
glabrous to strigose above, pilose beneath. State of Mexico to 

Oaxaca 98. F. cylindracea. 

FF. Petioles 15-30 (40) mm. long; branchlets puberulent; leaves 
pubescent on both surfaces. Oaxaca to Guatemala. 

99. F. tetradactyla. 
EE. Hypanthium of pistillate flowers barely 1 mm. wide ; petals appar- 
ently white; leaves subglabrous to minutely puberulent on both 
surfaces. Jalisco to Morelos 100. F. Mexiae. 

(85) Fuchsia thymifolia H.B.K. 

(Plate 15, fig. 76) 

Fuchsia fhijmifolia H.B.K., Nov. Gen. et Sp., 6:104, pi. 535, 1823. F. ihymifolia DC, Prodr., 
3:37, 1828; Lindl., Bot. Eeg., 15:1284, 1829; Sweet, Brit. Fl. Gard., 4:25, 1829; Hem- 
SLEY, Bot. Biol. Centr. Amer., 1:460, 1880; Standley, Contr. U. S. Nat. Herb., 23:1079, 
1924. 

Lopezia thymifolia "Willd., herb. Link, Jahrb., I. 3:25; ex Link in Schult., Mant., 1:50, 
according to H.B.K., I.e., 

Lyciopsis thymifolia Spach, Ann. Sci. Nat., (2) 4:176, 1835. 

Encliandra thymfolia Lilja, Linnaea, 15:262, 1841. 

Fuchsia alternans Moc. and SESsfi, Fl. Mex. Icon, ined., ex DC, Prodr., 3 :37, 1828, in synon. 

F. parviflora Lindl., Bot. Eeg., 13:1048, 1827. Type not seen, but the figure given is good. 

F. ovata Sess£ and MociNO ex DC, Prodr., 3 :37, 1828, in synon. Not seen. 

Shrub about 1 m. tall, openly branched, with slender puberulent somewhat 
reddish twigs, the ultimate ones scarcely 1 mm. thick; leaves opposite to alter- 
nate, not crowded, elliptic-ovate to round-ovate, subentire, rounded to obtuse 
at base, obtuse to acute at apex, membranaceous, paler beneath than above, 
minutely pubescent to subglabrous above, minutely pubescent beneath, some- 
what ciliate, the blades 0.7-2 (3) cm. long, 0.5-1.6 cm. wide, with 3-5 pairs 
lateral veins ; petioles very slender, puberulent, 3-10 mm. long, often reddish ; 
stipules subulate, somewhat glandular, scarcely 1 mm. long ; flowers solitary, 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 89 

axillary, white to pink, darker in age ; pedicels puberulent, filiform, 5-10 (12) 
mm. long ; hypanthium puberulent without and within, ob conic, 4-6 mm. long, 
2-3 mm. wide at summit in pressed specimens, whitish to reddish (especially 
in age) ; sepals ovate, puberulent, about 4 mm. long, white and spreading- 
reflexed at anthesis, later red, the subulate tips connivent in bud, 1 mm. long; 
petals ovate-oblong, spreading, 4 mm. long, white becoming purplish red, ob- 
tuse, often undulate ; stamens subsessile, the episepalous ones exserted from 
hypanthium, 1.5 mm. long, the epipetalous included, 1 mm. long ; style white, 
glabrous, exserted, 6-9 mm. long from base of hypanthium ; stigma greenish, 
the lobes erect or spreading, barely 0.5 mm. long ; berry globose, glabrate to 
puberulent, 4-6 mm. long. 

Type locality, Patzcuaro, Michoacan, Mexico. Kanging through several 
states of south central Mexico; representative material, without locality, 
fragment Bonpland 4106 (F) ; ex horto bot. petropolitano, seeds from Mexico, 
in 1828 (NY); Coulter 176 (NY). Hidalgo, Sierra de Pachuca, Rose and 
Painter 8023 (US), 6737 (GH, NY, US), PnngrZe 9472 (G, US in part), 5754 
(F, GH, MICH, NY, US), Eose, Painter and Rose 8859 (GH, NY, US) ; El 
Chico, Lyonnet 534 (US). Michoacan, Quinceo near Morelia, Arsene 3228 
(GH, US) ; Cerro Azul, Arsene 2453 (US) . Mexico, Salazar, Rose and Painter 
7035 (US) ; Federal District, Cerro Xitle, Russell and Souviron 175 (US) ; 
Eslava, Rushij 342 (NY, US) ; Pedregal, Pringle 7206 (GH, US) ; Contreras 
Canyon, Munz 15001 (POM). Vera Cruz, Pedregal Naolingo, Purpus 296 
(US). Sinaloa, El Carrizo, Ortega 6009 (US). 

(86) Fuchsia Pringlei Robinson and Seaton 

(Plate 15, fig. 77) 

Fuchsia Pringlei Robins and Seat., Proe. Arner. Acad., 28:106, 1893; Standley, Contr. 
U. S. Nat. Herb., 23:1079, 1924. 

Habit and pubescence of F. thymifolia, but leaves closer together, blades 
mostly less than 1 cm. long, 0.3-0.7 (1) cm. wide; petioles 2-5 (7) mm. long; 
hypanthium 2-3 mm. long ; sepals 2-3 mm. long ; petals 2-3 mm. long ; style 
4r-o mm. long. 

Type locality, Patzcuaro, Michoacan. In the original description were 
cited : Mts. near Patzcuaro, Pringle 5063 ; barranca near Guadalajara, Pringle 
5002, and Mts. near Patzcuaro, Pringle 4140. Since the first two cited were in 
fruit and only the last had flowers, it should be taken as the type number. 
Fruiting specimens alone can be distinguished from F. thymifolia only with 
difficulty. The species ranges with F. thymifolia. Specimens seen, MEXICO : 
"Fuchsia microphylla N. ic." Sesse, Mocino, Castillo, y Maldoyiado 5210 (F) ; 
Salto de Agua, Purpus 1780 (POM, US) . Hidalgo, Pachuca, Rose and Hough 
4452 (GH, US). Mexico, Comunidad, Temascaltepec, Hinton 1814 (GH, US) ; 
Popocatepetl, Purpus 3059 (NY, US) ; Ixtaccihuatl, Purpus 370 (US) ; Cima, 
Rose and Painter 7189 (GH, US) ; Federal district, Sierra de Ajusco, Pringle 
6412 (CAS in part, F in part, MO, NY, US in part), 11988 (F, GH, MICH. 



90 CALIFOEXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

US). Vera Cruz, Acultzinco, Matuda 1097 (F, MICH, NY); Maltarata, 
Matilda 1218 (F, MICH) . Morelos, Serrania de Ajusco, Pringle 7570 (POIM) ; 
Ti^s Marias, Orcutt 3754 (F, US) . Michoacan, Morelia, Cerro Azul, Arscne in 
1909 (F, GH) ; Patzcuaro, Pringle 4140, type no. (F, GH, CAS, NY, US) ; 
Tarascon, Pringle 13463 (US). Tepie, between* Santa Gertrudis and Santa 
Teresa, Rose 2117 (US). Jalisco, west of Bolafios, Rose 2964 (US), not typi- 
cal ; flowers white. 

(87) Fuchsia minimiflora Hemsley 

(Plate 15, fig. 79) 

Fuchsia minimiflora Hemsl., Bot. Biol. Centr. Amer., 1:458, 1880; 5: pi. 28, fig. 1, 1880; 
Standley, Contr. U. S. Nat. Herb., 23 : 1079, 1924. 

Shrub 1-4 m, tall, loosely branched with long very slender reddish crisp- 
puberulent twigs; leaves oposite or alternate, ovate to elliptic-ovate, acute at 
apex, i^ounded to obtuse at base, subentire to serrulate, paler beneath, sub- 
giabrous to minutely crisp-puberulent, the blades mostly 1-2 cm. long, 0.5- 
1.2 cm. wide ; petioles very slender, puberulent, 2-5 mm. long ; flowers solitary, 
axillary ; pedicels filiform, puberulent, 4-12 mm. long ; hypanthium obconic, 
1-2 mm. long, whitish to reddish, puberulent without; sepals lanceolate, 
reflexed-spreading, whitish to reddish, 1.5-2.5 mm. long, the tips apiculate, 
spreading in bud ; petals oblong-elliptic, white to red, less than 1.5 mm. long; 
stmens subsessile, less than 1 mm. long ; style glabrous, 2-3 mm. long ; stigma 
4-lobed, the lobes spreading, 0.5-1 mm. long ; berry subglobose, glabrate to 
puberulent, 4-5 mm. thick. 

Type locality, Chiapas, Mexico. A species of south-central Mexico, known 
from but few collections, of which I have seen : MEXICO, state not known to 
me, Salto de Agua, Purpus 1780 in part (F) . Mexico, Cima, Rose and Painter 
7189 (US) ; Federal District, Sierra de Ajusco, Rose and Hay 5521 (US, 
Pringle 6412 in part( F, GH, MO, US) ; District of Temascaltepec, Hint on 
1351 (F, NY) . Puebla, Boca del Monte, Piirpus 5828 (F, GH, NY, US) . Chia- 
pas, without definite locality. Linden 680, type coll., photo of Geneva specimen 
(F). 

(88) Fuchsia colimae Munz, new species 
(Plate 15, fig. 78) 

Habit and puberulence of F. minimiflora; leaves orbicular-ovate, the blades 
5-12 mm. long, 4-11 mm. wide, obscurely serrulate, obtuse to rounded at both 
ends; petioles 3-5 mm. long; hypanthium whitish-green, 2-2.5 mm. long; 
sepals white, reflexed-spreading, 1.5 mm. long, the tips not free in the bud ; 
petals white, elliptic-oblong, 1 mm. long ; stigma-lobes spreading. 

Habitus et puberulencia Fuchsiae minimiflorae, foliis orbiculare-ovatis, laminis 5-12 
mm. longis, 4-11 mm. latis, obscure serrulatis, obtusis vel rotundatis in extremis utrisque; 
petiolis 3-5 mm. longis; hypanthio albido, 2-2.5 mm. longo ; sepalis albidis, reflexo-expansis, 
1.5 mm. longis; petalis albis, elliptico-oblongis, 1 mm. longis; lobis stigmatis expansis. 

Type: from Volcano of Colima, Jalisco, Mexico, July 13, 1892, M. E. Jones 



Vol. XXV] MUNZ : TEE GENUS FUCHSIA 91 

224, Pomona College Herbarium 83711 ; isotypes United States National 
Herbarium 237441 and at New York Botanical Garden. This species is near 
F. minimiflora, but the sepals are shorter than the hypanthium, the tips are 
not free in the bud, and the leaves are more rounded. 

(89) Fuchsia tacanensis Lundell 

(Plate 15, fig. 80) 
Fuchsia tacanensis Lundei^l, Contr. Univ. Mich. Herb., 4:20, 1940, 

Shrub up to 4 m. high, open-branched, the ultimate branchlets slender, 
greenish, subglabrous to pubescent; leaves opposite, the blades membranace- 
ous, rhombic-ovate, abruptly cuneate at base, acuminate at apex, rather evenly 
and coarsely sinuate-serrulate, paler beneath than above, somewhat strigose 
above, subglabrous to pilose beneath, 2.5-5 (7) cm. long, 1.2-3.5 (4.2) cm. 
wide, with 7-9 pairs of lateral veins ; petioles slender, subglabrous to pubes- 
cent, 1-2 cm. long; pedicels filiform, glabrous to pubescent, mostly 1-2 cm. 
long; flowers perfect or imperfect; hypanthium obconic, 3-5.5 mm. long, 
2-3.5 mm. wide at summit, glabrous to puberulent, greenish to reddish ; sepals 
reflexed-spreading at anthesis, lanceolate, acuminate, usually somewhat pu- 
bescent, whitish or greenish, usually shorter than hypanthium, 2.5-6 mm. 
long, apiculate, the tips 1-2 mm. long, connivent in bud ; petals oblong-ovate, 
white to pink, red in age, 2-3.5 mm. long, more or less emarginate; stamens 
included, anthers about 1 mm. long ; style in perfect flowers 4-7 mm. long ; 
stigma 4-lobed, scarcely 1 mm. long ; fruit subglobose, black, 4-5 mm. thick. 

Type locality, Volcan de Tacana, Chiapas, Mexico ; the species is known 
from the following collections, all from MEXICO : without locality, Schiede 
(US) ; Morelos, barranca above Cuernavaca, Pringle 6380 (F, GH, CAS, 
MICH, NY, US) ; Chiapas, Volcan de Tacana, Matuda 2399 (MICH, type) ; 
Mt. Pasitar, Matuda 1686 (MO, NY, MICH, US). F. tacanensis is variable in 
flower-size, pubescence, length of hypanthium, etc., but is distinguished by the 
large thin toothed leaves, wide hypanthium, long sepal-tips, light-colored 
flowers, sepals shorter than hypanthium. Pringle 6380 has staminate more 
crowded flowers, pedicels less than 1 cm. long. 

(90) Fuchsia Skutchiana Munz, new species 

(Platel5, fig. 81) 

Much like F. tacanensis in habit, foliage, etc., but finely strigulose on young 
branchlets, hypanthia, sepals; leaves more finely strigose; petioles 2-3 cm. 
long; pedicels 1.5-3 cm. long, strigulose; hypanthium not longer than wide 
in pressed specimens, 2-3 mm. long, slightly wider ; sepals at least as long as 
hypanthium, reflexed, 2.5-4 mm. long, white to pink, aging red, the tips 1-1.5 
mm. long; petals about as long as sepals, white to pink, aging red, emarginate ; 
stamens not included, about 1 mm. long, anthers 0.5 mm. long; style 4—5 mm. 
long ; fruit black, 4-5 mm. thick. 



92 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Habitus et folia Fuchsiae tacanensis; ramulis, hypanthiis sepalisque strigulosis; foliis 
strigulosis; petiolis 2-3 cm. longis; pedicellis 1.5-3 cm. longis, strigulosis; hypanthio 2-3 
mm. longo, latiore; sepalis hypanthio aequalibus, reflexis, 2.5-4 mm. longis, albis vel carneis, 
in aetate roseis, cum apicibus 1-1.5 mm. longis; petalis sepalis aequalibus, emarginatis, 
albis vel carneis, in aetate roseis; staminibus non inclusis, 1 mm. longis; antheris 0.5 mm. 
longis; stylo 4-5 mm. longo; bacca nigra, 4^5 mm, diametro. 

Type: from near Quezaltenang-o, Quezaltenango, Guatemala, at 2500 m., 
July 24, 1934, A. F. Skutch 812 (NY) ; isotype (F) . Other material from same 
place and on same date, Skutch 813 (F) ; Chiapas, Mexico, Ghieshreght 730, 
in 1864-70 (F, Gil, MO). 

(91) Fuchsia bacillaris Lindley 

(PlatelS, fig. 82) 

Fuchsia bacillaris Lindl., Bot. Eeg., 18:1480, 1832; F. bacillaris in Curtis, Bot. Mag., 76: 
4506, 1850; Hemsley, Biol. Centr. Amer., Bot., 1:456, 1880; Standley, Contr. U. S. Nat. 
Herb., 23:1079, 1924. 

Shrub 1-2 m. tall, practically glabrous throughout, openly and finely 
branched ; leaves opposite, subcoriaceous, sinuate-denticulate, paler beneath 
than above, lance-ovate to elliptic-ovate, more or less cuneate at base, acute to 
obtuse at apex, sometimes with line of minute hairs on upper surface of mid- 
rib ; blades 1-3 (4.5) cm. long, 0.5-1.2 (1.5) cm. wide; petioles 3-6 mm. long, 
sometimes with lines of puberulence ; flowers solitary, axillary, apparently 
perfect or pistillate ; pedicels glabrous, filiform, 5-15 mm. long ; hypanthium 
subcylindric, glabrous, red, 5-6 mm. long, 1.5-2.5 mm. wide at summit ; sepals 
spreading-reflexed at anthesis, rose, glabrous, lanceolate, 3-5 mm. long, apicu- 
late, the tips connivent in the bud ; petals rose, round-obovate, refuse, 2.5-5 
mm. long; stamens included, the anthers 1-1.5 mm. long ; style 8-13 mm. long ; 
stigma reddish, the 4 lobes tending to spread ; fruit spherical, 4-5 mm. thick. 

Type locality, Mexico, seeds sent by J. S. Mill, Esq. This species is an un- 
certain entity, the exact location of its origin being unknown. It has always 
been keyed out as being glabrous. The plate by Lindley shows it as having 
relatively large flowers and is pretty well matched by Purpus 6708. Material 
seen, MEXICO : "seed from Mexico, 1829," Ex horto bot. Petropolitano (NY) , 
matching the Lindley plate and description and is likely authentic material. 
Guerrero: Mts. near Chilpancingo, E. W. Nelson 2258 (US), with leaves 
rather small; Sierra Madre, near Chilpancingo, Nelson 2232 (US) ; crest of 
Sierra Madre, Langlasse 890 (F, GH, US). Chiapas: Cerro del Boqueron, 
Purinis 6708 (F, GH, NY, US). GUATEMALA: Iluehuetenango, Sierra 
Cuchumatanes, Skutch 1247 (F), twigs slightly pubescent. 

(92) Fuchsia minutiflora Hemsley 

Fuchsia minutiflora Hemsl., Diag. PI. Nov., 1:15, 1878; Bot. Biol. Centr. Amer., 1:458, 
1880; 5: pi. 28, 1880; Standley, Contr. U. S. Nat. Herb., 23:1079, 1924. 

Shrub 1-2 m. tall, with very slender rather numerous branches, the ultimate 
ones scarcely 1 mm. thick and covered with a loose brownish puberulence, the 



Vol. XXV] MUNZ: TEE GENUS FUCHSIA 93 

hairs more or less spreading, often with incurved tips ; leaves mostly opposite, 
numerous, mostly longer than the internodes, the blades thickish, lance-ovate 
to -elliptic, conspicuously serrulate with 4-5 teeth on each edge, glabrous, 
somewhat paler beneath, 3-8 (10) mm. long, 2-5 (7) mm. wide, obtuse to acute 
at both ends; petioles 1-2 (4) mm. long; flowers solitary, axillary; pedicels 
filiform, sparsely puberulent, 2-8 mm. long; hypanthium tubular-funnel- 
form, 3-4 mm. long, 1.5 mm. thick at base, gradually ampliate, subglabrous 
to puberulent, apparently reddish ; sepals lanceolate, acuminate, apiculate, 
somewhat spreading in anthesis, 2-2.5 mm. long, the tips connivent in the bud ; 
petals oblong, obtuse, entire, 1.5-2 mm. long, apparently white ; stamens very 
small, almost included ; style glabrous, scarcely as long as petals, stigma deeply 
4-lobed ; berry sub-globose, subglabrous, 3-4.5 mm. thick. 

Ket to Varieties of Fuchsia minutiflora 

A. Hypanthium and sepals white; hypanthium pubescent 92a. var. hidalgensis. 

AA. Hypanthium red, subglabrous 92b. var. typica. 

(92a) Fuchsia minutiflora Hemsley, var. hidalgensis Munz, new variety 

(Plate 15, fig. 83) 

Hypanthium pubescente, album ; sepalis albis. 

Type: from Trinidad, Hidalgo, Mexico, at 1850 m., April 30, 1904 Pringle 
8808, Pomona College Herbarium 59867 ; isotypes (F, GH, CAS, MICH, NY, 
US), the label on the Michigan specimen reads "Honey Station, Puebla." 
Material seen, Trinidad, Pringle 8807 (F, GH, CAS, NY, US) ; Zacualtipan, 
Selerl50 (GH, US). 

(92b) Fuchsia minutiflora Hemsley, var. typica Munz, new name 
Fuchsia minutiflora Hemsl., Diag. PI. Nov., 1 :15, 1878. 

Hypanthium red, subglabrous. 

Type locality, "in monte Orizaba, alt. 5000 ped.," based on Linden 631. 
Material seen, MEXICO : without locality, Mueller 1339 (NY) ; garden of 
Mrs. Nuttall, Mexico City, Safford 1511 (US) ; Vera Cruz, Chinantla, Lieh- 
mann 3231 (US) ; Prov. Jalapa, com. Schlechtendal in 1840, coll. Schiede 525 
(GH, NY) ; Aseradero de Santa Cruz, Mueller 1550 (NY) ; Orizaba, Mueller 
1702 (GH, NY), 640 (NY) ; Maltrata, Matuda 1358 (MICH, NY). Mexico, 
Ixtaccihuatl, Purpus 160 (POM). 

(93) Fuchsia microphylla H.B.K. 

Fuchsia microphylla H.B.K., Nov. Gen. et Sp., 6 :103, pi. 534, 1823. 

Subshrub or shrub, 3-20 dm. tall, rather densely branched, the ultimate 
branchlets slender, barely 1 mm. thick, with a dense subappressed puberu- 
lence; leaves crowded, opposite, subcoriaceous, lance-oblong, to elliptic-obo- 
vate, usually sharply serrulate, sometimes subentire, deep green above, paler 
beneath, acute to obtuse at both ends, subglabrous except sometimes for line 
of puberulence along upper surface of midrib, the blades 5-20 (25) mm. long. 



94 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

3-8 (10) mm. wide; petioles glabrous to puberulent, 2-8 (15) mm. long; 
flowers solitary, axillary, perfect or pistillate ; pedicels appressed-puberu- 
leut, 3-6 (10) mm. long; hypanthiiim deep red, subcylindric, constricted 
above the ovary, then usually 7-9 (11) mm. long, 2.5-5 (6) mm. wide in 
pressed specimens, sometimes shorter and narrower in pistillate flowers, gla- 
brous to minutely strigose ; sepals deep red, mostly 4, sometimes 3, lanceolate 
to ovate, 2.5-5 (7) mm. long, acuminate, usually apiculate, the tips up to 1 
mm. long and connivent in the bud, glabrous to strigulose ; petals rose, broadly 
elliptic to almost round, usually toothed, 3-5 (7) mm. long; stamens nearly 
or quite included ; anthers 1.5-2 mm. long, sometimes lacking ; style 8-15 (20) 
mm. long; stigma reddish, deeply 4-lobed; fruit subglobose, glabrous to 
strigulose, purple-black when ripe, up to 8 mm. thick. 

Characterized by its coriaceous glabrous usually serrulate and smallish 
crowded leaves and rather large flowers, but variable; flowers small if pistil- 
late only. 

Key to VAiiiETiEs of Fuchsia microphylla 

A. Ovary and hypantliium glabrous 93a. var. typica. 

A A. Ovary and lower part of hypantliium minutely strigulose 93b. var. aprica. 

(93a) Fuchsia microphylla H.B.K. var. typica Munz, new name 

(Plate 15, fig. 84) 

F. microphylla H.B.K., Nov. Gen. et Sp., 6:103, pi. 534, 1823; Lindl., Bot. Eeg., 15:1269, 
1829; Sweet, Brit. PI. Gard., ser. 2, pi. 16, 1829?; Hemsley, Bot. Biol. Centr, Amer., 
1 : 458, 1880 in part ; Standley, Contr. U. S. Nat. Herb., 23 : 1079, 1924 in part. 

Brebissonia microphylla Spach, Ann. Sci. Nat., ser. II, 4:175, 1835. 

Myrinia microphylla Lilja, F1. Sverig. Suppl., 1:25, 1840. 

F. gracilis Sess£ and Mocino ex DC, Prodr., 3:37, 1828 in sjTion. 

F. mixta Hemsley, Diag. PI. Nov., 1:14, 1878; Biol. Centr. Amer. Bot., 1:459, 1880; 5:pl. 
28, fig. 2, 1880. Based on Galeotti 3025 from Orizaba, which I have not seen, but which 
is in the range of the typical form if referred to F. microphylla. I agree with Standi., I.e., 
that such must be the case. 

Ovary and hypantliium glabrous. 

Type locality, Volcan de Jorullo, which I believe is in Michoacan, Mexico. 
Ranging from Hidalgo to states of Mexico, Michoacan and Jalisco. Material 
seen, MEXICO : without definite locality, Sesse, Mocino, Castillo y Maldonado 
5197 (F) ; Lithrum vulnerarium, N., Sesse et at 1842 (F) ; ex horto bot. Petro- 
politano, introd. 1828 from Mex. (NY). Hidalgo, El Chico, Lyonnet 2204 
(US) ; between Real del Monte and Chico, Hartweg 285 (NY) ; Lagunas 
Zempoala, Lyonnet 2510 (US). Vera Cruz, Pico de Orizaba, Vacqueria de 
Jacal, Lielmann 3230 (US, GH). Federal District, El Desierto, Conviento, 
Russell and Souviron 133 (CAS, US) ; La Cima, Barnes and Land 395 (F) ; 
San Nicolas, Valle de Mexico, Bourgeau 974 (GH, US) ; Pass near Desierto 
de los Leones, Munz 15062 (POM) intergrading toward aprica; Contreras, 
Lyonnet 1637 (US). Mexico, Salazar, Pringle 13462 (US), Rose and Painter 
7036 (US) ; Montes de Gavia, Asphmd 356 (NY) ; Nevada de Toluca, Rose 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 95 

and Painter 7935 (NY, US) ; Sierra de las Cruces, Pringle 4314 (F, GH, 
MICH, NY, US) ; Ixtaccihuatl, Punms 1779 (F, GH, NY, POM, US) ; Meson 
Viejo, Temascaltepec, Hinton 1902 (US) ; San Rafael, Lyonnet and Elcoro 
1923 (US). Morelos, Popocatepetl, Purpus 3060 (F, NY, GH, US), Rose and 
Hay 6256 (US). Michoacan, Mt. Patamban, Nelson 6566 (NY, US); "in 
monte ignivomo JoruUo, alt. 550 hex.," Humboldt et al, type coll. (scrap at 
F). Jalisco, volcano of Colima, Jones 225 (POM, US). 

(93b) Fuchsia micropliylla H.B.K. var. aprica (Lundell), new comb. 
F. aprica Lundell, Contr. Univ. Mich. Herb., 4:18-19, 1940. 

Ovary and lower part of hypanthium minutely appressed-puberulent. 

Type locality, Volcan de Tacana, Chiapas, Mexico. Ranging from Oaxaca 
and Chiapas to Guatemala. Representative material seen, MEXICO, Puebla, 
Tezuitlan, Orcutt 4001 (F, MO, US). Oaxaca, Yavezca, Galeotti 3039 (US) ; 
Cerro de los Frailes, between Teotitlan del Camino and Huantla de Jimenez, 
Schultes and Reko 370 (F, GH) . Chiapas, San Cristobal, Goldman 985 (US) ; 
Mt. Tacana, Matuda 2378, type coll. aprica (MICH, NY). GUATEMALA: 
Alta Vera Paz, Mts. east of Tactic, Standley 71428 (F). Huehuetenango, San 
Juan Ixcoy, Skutch 1069 (F). Quiche, San Miguel Uspantan, Heyde and Lux 
under J. D. Smith 2999B (NY, US) ; Nebaj, Skutch 1711 (F) pistillate. San 
Marcos, Volcan de Tajumulco, Schmidt 1934 (F) ; Volcan de Tacana, Steyer- 
mark 36159 (F). Quezaltenango, Volcan de Santa Maria, Steyermark 34187 
(F). Chimaltenango, Volcan de Acatenango, Standley 61954 (F) ; Volcan de 
Agua, Johnston 896 (F). Guatemala, between San Martin and Todos Santos, 
Nelson 3627, pistillate (F, US). Sacatepequez, Antigua, Kellerman 6535 (F) ; 
Volcan de Agua, Kellerman 7577 (NY). 

This pubescent plant intergr^des freely with F. michoacanensis which has 
thinner more membranaceous leaves with coarser longer hairs on their lower 
surface, for example: GUATEMALA: Quezaltenango, Hartweg 527 (NY) ; 
Sacatepequez, Volcan de Agua, Shanyion 3655 (US) ; Alta Vera Paz, Coban, 
von Tuerckheim II 1295 (F, MICH, NY) ; Mts. east of Tactic, Standley 71357 
(F). Oaxaca, Cerro de la Virgen, Conzatti 4352 (US). In these and many 
others, one gets varying combinations of thick leaves and pilosity. 

(94) Fuchsia Hemsleyana Woodson and Seibert 
(Plate 15, fig. 85) 

Fuchsia Hemsleyana Woodson and Seibert, Ann. Mo. Bot. Gard., 24:194, 1937. 

F. pulcliella "Woodson and Seibert, I.e., 195-196. Based on Seibert 183, which I have seen. 

Shrub 1-3 m. tall, rather openly branched, the ultimate branchlets slender, 
ca. 1 mm. thick, minutely pubescent, sometimes strigulose; leaves mostly 
opposite, rather crowded, subcoriaceous, rhomboid-elliptic to -obovate, cuneate 
at base, acute to obtusish at apex, serrulate to sinuate-dentate, paler beneath 
than above, subgiabrous except on upper surface of midrib and along margin, 
the blades 7-15 (25) mm. long, 4-8 (12) mm. wide ; petioles sparsely puberu- 



96 CALIFORNIA ACADEMY OF SCIENCES [Proc. iTH Ser. 

lent, 3-8 (12) mm. long; flowers solitary, axillaiy, perfect or pistillate; pedi- 
cels filiform, more or less puberulent, 4-6 (8) mm. long; hypanthium rose, 
tubular, constricted above ovary, then 5-7 (9) mm. long, 1-2 mm. wide at 
summit, glabrous to puberulent; sepals deltoid-lanceolate, rose, spreading- 
reflexed at anthesis, 3.5-4 mm. long, apiculate, the tips connivent in the bud ; 
petals rose or with purplish tinge, oblong-obovate, 2.5-4 mm. long, undulate 
and rounded at apex; stamens usually included; anthers 1-1.2 mm. long; 
style 8-10 mm. long; stigma deeply 4-lobed, exserted, 1 mm. long; ovary sub- 
glabrous to strigulose ; fruit subglobose, black-purple, 4-5 mm. thick. 

Type locality, Rio Chiriqui Viejo, near Monte Lirio, Chiriqui, Panama- 
Found in the mountain of Costa Rica and Panama. Representative material 
may be cited, COSTA RICA : Vara Blanca de Sarapiqui, Skiitch 3225 (NY). 
Alajuela, Volcan de Poas, Standley 34629 (VQ), 34620 (US), Allen 600 (F). 
Cartago, Cerros de Irazu, Pittier 13048 (US) ; Volcan de Turrialba, Standley 
34949 (US). San Jose, La Guaba, Valerio 1526 (F) ; Laguna de la Clionta, 
Standley 42275, strigulose (F, US) ; Cerro de Piedra Blanca, above Escasu, 
Standley 32598 (US), Solis 154 (F) strigulose; Finca La Cima above Los 
Lotes, Standley 42758 (US); Cerro de Las Vueltas, Standley and Valerio 
43523 (F, \^^), 43499 (US) ; Laguna de la Escuadra, n.e. of El Copey, Stand- 
ley 42078 (F, US), 42057 (US) ; between Guayabillos and Cabeza de Vaca, 
Dodge and Thomas 4940 (F). PANAMA: Chiriqui, upper Rio Chiriqui 
Viejo, Seibert 162, type no. Hemsleyana (MO, NY) ; near Monte Lirio, Scibert 
182, type pulchella (MO) ; Rio Chiriqui Viejo valley. White 2 (GH, POM) ; 
trail Paso Ancho to Monte Lirio, Allen 1497 (MO, NY, POM) ; Volcan de 
Chiriqui, Terry 1331 (F, MO), Davidson 876 (F, MO) ; Casita Alta, ^yoodson, 
Allen and Seibert 873 (MO, POM) ; Bajo Chorro, Boquete District, Davidson 
319 (F, MO.) ; Cuesta de Cerro Quemado, Pittier 3114 (US, GH). 

This species is near F. microphylla in its coriaceous, smallish, usually 
toothed leaves, but has smaller flowers and a more southern range. It often is 
much like F. michoacanensis, but the leaves are thicker, smaller, not pilose ; 
however, it intergrades with it (see under michoacanensis) . Plants that I 
refer to Hemsleyana have largely been determined as F. parviflora (F. 
Encliandra of this paper), but that species belongs in the thinner or mem- 
branaceous-leaved group. 

(95) Fuchsia striolata Lundell 
(Plate 15, fig. 86) 
Fuchsia striolata Lundell, Contr. Univ. Mich. Herb., 4:20, 1940. 

Shrub, openly branched, more or less scandent, the ultimate branchlets 
somewliat strigulose ; leaves opposite, subcoriaceous, ovate to lance-ovate, the 
blades 1.5-9.0 cm. long, 1-5 cm. wide, cuneate at base, acuminate to acutish at 
apex, minutely strigulose above, subglabrous and paler beneath, subentire 
to remotely denticulate in upper half, ciliolate, with 6-7 pairs of lateral veins ; 
petioles strigulose, 1-2.5 cm. long; flowers solitary, red; pedicels strigulose. 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 97 

6-12 mm. long; hypanthium tubular, strigulose, 3-4 mm. long, barely 1 mm. 
wide ; sepals red, deltoid, ea. 2 mm. long, apieulate ; petals red, 1 mm. long, 
rounded ; anthers included, 0.6 mm. long ; style 4-5 mm. long; stigma exserted, 
4-lobed, less than 1 mm. long; fruit subglobose, ca. 7 mm. thick, strigulose. 
T.^'pe locality, Volcan de Tacana, Chiapas, Mexico, the type number being 
Matilda 2765 (F, ]\IICH, NY) entirely without flowers, although the sheet 
at New York has one young bud. On the whole, though, Matuda 2765 seems to 
match Matuda 4004, from Mt. Ovando, Chiapas (MICH, NY) which does 
have flowers. From GUATEMALA : San Marcos, Volcan Tacana, Quebrada 
Canjula, between Sibinal and Canjula, SteyermarJc 3608 (F) also without 
flowers, seems to be this species. It is characterized by the large thickish leaves 
which are quite glabrous beneath and with short pedicels and small flowers. 
A collection from Jalisco, San Sebastian, Canon El Ranchito, Mexia 1465 (F, 
MICH, NY, US) has certain characters of F. striolata (leaf -shape, size, pubes- 
cence; pedicel -length ; flower-size, shape, color; fruit-size), but the leaves are 
very much thinner than in the Chiapas specimens. 

(96) Fuchsia michoacanensis Sesse and Moeino 

(Plate 15, fig. 87) 

Fuchsia michoacanensis Sess6 and MociNO, PI. N. Hispan., ed. 1, 57, 1887-90. 

Fuchsia iiflora Sess6 and MociNO, PI. Mex., ed. 2, 93, 1894 given as same as F. michoacan- 
ensis. Based on specimen no. 5209, which I have seen. 

F. chiapensis T. S. Brandg., Univ. Calif. Pub. Bot., 6:59, 1914; Standley, Contr. U. S. Nat. 
Herb., 23 : 1079, 1924. Based on Purpus 6707, which I have seen. 

F. heterotricha Lundell, Contr. Mich. Herb., 4:19, 1940. Based on Matuda S457, which I 
have seen. It may have somewhat finer pubescence than usual. 

Shrub 1-3 m. tall, with slender branches, the ultimate ones about 1 mm. 
thick, adpressed-puberulent to densely spreading-pubescent or pilose, usually 
with both kinds of hairs at least on the petioles and veins of lower leaf- 
surfaces; leaves opposite, the blades orbicular-ovate or elliptic-ovate, mem- 
branaceous, subentire, to denticulate, bieolored, rounded to cuneate at base, 
rounded to acute at apex, subglabrous to puberulent or sparingly strigose 
above, sparsely to densely pilose beneath, 7-25 (40) mm. long, 5-20 (25) mm. 
wide ; petioles 3-25 mm. long, usually pilose and strigulose ; flowers axillary, 
perfect or pistillate; pedicels pilose to puberulent, filiform, 5-15 (25) mm. 
long ; hypanthium red, cylindrical in pistillate flowers to tubular-f unnelf orm 
in perfect ones, often more or less pilose, pubescent, 4-6 (10) mm. long, 1-3 
mm. wide at apex in pressed specimens, constricted below the rounded base ; 
sepals red, usually pilose, sometimes only puberulent, more or less spreading, 
deltoid, acuminate, 2-3 mm. long, the apieulate tips connivent in bud ; petals 
coral red, round-ovate to -elliptic, erect, 0.6-2 mm. long; stamens subsessile, 
almost or quite included; style 6-8 (12) mm. long, the stigma somewhat ex- 
serted, deeply 4-lobed, more or less red ; berry subglobose, black-purple, 4-7 
mm. thick. 



98 CALIFORNIA ACADEMY OF SCIENCES [Paoc. 4th Ser. 

Type locality, not given, probably southern Mexico. Ranging from southern 
Mexico to Costa Rica. Representative material, MEXICO : without definite 
locality, ''Ftichsia hiflora N. t. 2. f. 159," Sesse, MociTio, Castillo and Maldon- 
ado 5209 (fragment at F). Mexico, district of Temascaltepec, Pantoja, Hinton 
2846 (NY) . Michoacan, Tarascon, Pringle 11989 (F, GH, US) . Chiapas, Cerro 
del Boqueron, Piirpus 6707, type no. chiapensis (F, GH, NY, US), 7168 (F, 
NY, US) ; Volean de Tacana, Matuda 2457, type heterotricha (MICH). Jal- 
isco, Real Alto, trail to El Tajo de Santiago, Mexia 1744 (F, CAS, MICH, NY, 
US). Morelos, El Parque, Rose and Rose 11118 (NY, US) very smooth. 
GUATEMALA : San Marcos, Quebrada Conjula, Volean Tacana, Steyer- 
mark 36047 (F) ; Rio Vega, Volean Tacana, Steyermark 36233 (F) ; El 
Boqueron, Standley 66305 (F, POM) . Quiche, so. of Chiehicastenango, Stand- 
ley 62408 (F). Chimaltenango, Chichivac, Skutch 43 (US), 710 {¥)'; Cerro 
de Tecpam, Standley 58716 (F). Guatemala, Finca La Aurora, Aguilar 80 
(F); Volean de Pacaya, above Las Calderas, Standley 58400 (F, POM). 
Quezaltenango, Volean Santa Maria, Steyermark 33915 (F) ; above Santa 
Maria de Jesus, Standley 67264 (F), 67188 (F) ; Volean Zunil, Steyermark 
34929 (F). Jalapa, Volean Jumay, Steyermark 32353 (F), Amatitlan, be- 
tween San Vicente Pacaya and El Cedro, Tonduz 428 (US). Sacatepequez, 
Volean de Fuego, J. D. Smith 1460 (GH, US) ; Volean de Agua, so. of Santa 
Maria de Jesus, Standley 59494 (F, POM) ; Finca Carmona, s.e. of Antigua, 
Standley 63805 (F, POM). Sacapa, Sierra de las Minas, Steyermark 29741 
(F). EL SALVADOR : San Vicente, Volean de San Vicente, Standley 21549 
(NY, GH, US). COSTA RICA: San Jose, Las Nubes, Standley 38705 (US). 

Material which I refer to this species has largely passed as F. minutiflora 
which is a smaller-leaved and smaller-flowered plant of more northern range. 
F. michoacanensis intergrades freely with F. microphylla var. aprica espe- 
cially in Guatemala, where much of the material is intermediate. In Costa 
Rica, it intergrades likewise with F. Hemsleyanu, but in general, F. michoa- 
canensis has thin leaves while the other two have more coriaceous ones. Ex- 
amples of intergradation with microphylla are, Chiapas, Ohiesbraght 697 
(F) ; Barranca Eminencia, San Marcos, Guatemala, Standley 68586 (F) ; 
Cerro Quemado, Quezaltenango, Kellerman 5939 (US); San Juan de Dios, 
Sonsonate, El Salvador, Pittier 1995 (US) ; Volean de San Salvador, Stand- 
ley 22810 (NY, GH, US). In these there is a tendency toward the pubescence 
of michoacanensis and toward the small thickish leaves of microphylla. With 
the pubescence of F. michoacanensis, but the leaf -thickness of F. Hemsleyana 
may be cited, COSTA RICA: Cartago, Torres Rojas 48 (US) ; Volean de 
Turrialba, Pittier 13191 (GH, NY, US) ; Irazu, Knntze 2343 (NY). 

(97) Fuchsia Encliandra Steudel 

(Plate 15, fig. 88) 

Fuchsia Encliandra Steud., Nomen. Bot., 1:649, 1840 (pul)lished as Enclinndra) . 
Encliandra parvi flora Zucc, Abli. Akad. Wiss. Muench., 2:335, 1831-1836. 



Vol. XXV] MVNZ: THE GENUS FUCHSIA 99 

Fuchsia parvi flora Zucc, I.e., 337; Hemsley, Biol. Centr. Amer., Bot., 1:460, 1880; Stand- 
ley, Contr., U. S. Nat. Herb., 23:1080, 1924. Not F. parviflora Lindl., Bot. Eeg., 13: 
1048, 1827 (see F. thymi folia). 

Eierschlegeria Lindleyi Spach, Ann. Sci. Nat., (2), 4:176, 1835. 

Shrub 1-4 ni. tall, densely branched, with ultimate branchlets very slender, 
puberuleut, somewhat strigulose and dark colored under the whitish puberu- 
lence ; leaves mostly opposite, membranaceous, lance-ovate to round-elliptic, 
entire to inconspicuously serrulate, minutely stri^lose to almost glabrous, 
bicoloi-ed, obtusish to acute at both ends, blades 8-15 (20) mm. long, 4-10 
(15) mm. wide; petioles puberuleut, 4-8 (12) mm. long; flowers solitary, 
axillary, dioecious or polygamous; pedicels filiform, puberuleut, 5-15 mm. 
long ; hypanthium cylindrical, strigulose, red, not constricted at base in stam- 
inate flowers, 6-8 (10) mm. long, 2-3 mm. wide, in pistillate flowers con- 
stricted above ovary, then 5-7 mm. long, 1-2 mm. wide ; sepals deltoid-ovate, 
ascending in anthesis, puberuleut, reddish, 2.5-3.5 mm. long, apiculate, the 
short tips connivent in the bud ; petals sub orbicular, 2-3 mm. long, reddish, 
more or less undulate ; stamens included, anthers ca. 1.5 mm. long ; style 7-9 
mm. long; stigma exserted, deeply 4-lobed; berry dark, 7-8 mm. thick. 

Type locality, Mexico. Ranging from Hidalgo to Oaxaca, and character- 
ized by its small entire, thin leaves and cylindrical flowers. As here under- 
stood this species is not nearly so inclusive as determinations on herbarium 
material would indicate as to previous treatment. Material seen, MEXICO, 
Hidalgo, Zimapan, Coulter 175 (NY, GH), possible mistake on label. Mexico, 
base of Ixtaccihuatl, Arsene 5539 (MO, US). Morelos, Valle del Tepeite, 
Lyonnet 1358h (US) ; Huitzilac, Lyonnet 1994 (US) ; Valle de Cuernavaca, 
Lyonnei 735 (NY, US) ; kilometer 61, road, Mexico City to Cuernavaca, 
Mexia 2718 (F, MICH, GH, CAS, NY, POM, US) ; El Parque, Orcutt 4406 
(F, GH). Oaxaca, Sierra de San Felipe, Pringle 7485 (US), Nelson 1179 
(US) ; San Felipe, Rose and Hough 4605 (US) ; Oaxaca, Conzatti and Gon- 
zales 20 (MO, US) ; Reyes, Nelson 1803 (GH, US) ; Yotado, Galeotti 3068 H 
(fragment at F). 

(98) Fuchsia cylindracea Lindley 
(Plate 15, fig. 89) 
Fuchsia cylindracea Lindl., Bot. Keg., 24:66, 1838. 

Shrub, rather openly branched, up to 5 m. high, the ultimate branchlets 
purplish, subglabrous to sparingly pubescent, the youngest tips more densely 
but rather coarsely pubescent, rather slender ; leaves membranaceous, oppo- 
site, obovate to elliptic, subentire, cuneate at base, obtuse to acutish at apex, 
strongly bicolored, subglabrous to somewhat strigose above, more or less pilose 
beneath, the blades 2-5 (6.5) cm. long, 1-2 (2.5) cm. wide ; petioles somewhat 
pilose or pubescent, 3-10 (12) mm. long; plants apparently dioecious; flowers 
solitary axillary, dioecious ; pedicels filiform, pubescent, 10-25 mm. long ; 
staminate flowers with narrow basal portion, then widened into hypanthium 
8-10 mm. wide, cylindrical, deep red, somewhat pubescent ; sepals ovate- 
deltoid, 3-3.5 mm. long, red, somewhat ascending in anthesis, short-apiculate, 



100 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

the tips connivent in the bud ; petals red, suborbieular, more or less apiculate 
and notched, 3-4 mm. long ; stamens included, anthers 1.2-1.4 mm. long; style 
about 1 mm. long ; stigma 4-lobed, about 0.5 mm. long ; pistillate flowers con- 
stricted above the ovary, then expanded into a cylindric hj^anthium 4-5 mm. 
long and 2-3 mm. wide ; sepals and petals as above, but somewhat smaller ; 
stamens present, the anthers about 0.5 mm. long, not functional ? ; stigma ex- 
serted, 4-lobed, ca. 1 mm. long; fruit subglobose, dark, 6-8 mm. thick. 

Type locality, not known; seed of original plants from Mexico. I have seen 
none of Lindley's herbarium material, but his plate fits well the specimens I 
refer to this species, with large leaves, long flowers (cylindrical, % inch long 
ace. to Lindley), pubescence, etc. The species is near F. Enclianclra (parvi- 
flora) but has larger leaves and coarser hair ; there is some intergradation, as 
Ghieshreght 696 (MO) from Chiapas with leaves of cylindracea and flowers of 
Encliaoidra; "F. puhescens," SessS and Mocino 5215 (F). Kepresentative of 
cylindracea may be cited, MEXICO, without locality, Coulter 174 (GH, NY) . 
Mexico, District of Temascaltepec, Timbres, Hint on 3055 (US) ; Cajones, 
Hinton 4044 (US). Michoacan, Campanario near Morelia, Arsene 5665 (NY, 
US), 6523 (US), Arsene in 1910 (F) ; Patzcuaro, Pringle 4266 (F, GH, 
]\riCH, NY, US) , 3583 (F) . Oaxaca, 18 miles s.w. of Oaxaca City, Nelson 1391 
(US). 

(99) Fuchsia tetradactyla Lindley 
(Platel5, fig. 90) 

Fuchsia tetradactyla Lindl., Joum. Hort. Soc, 1:304, 1846. 

F. Seleriana Loesener, Verb. Bot. Ver. Brandenb., 55:179, 1913; Standley and Steyeb.- 
MARK, Field Mus. Nat. Hist., Bot., 22 : 363, 1940. 

Shrub 6-15 (24) dm. tall, openly branched, the ultimate branchlets red- 
dish, 1-2 mm. thick, puberulent, more or less strigulose ; leaves opposite or 
nearly so, broadly ovate, somewhat rhomboid, membranaceous, not crowded, 
subentire or remotely denticulate, soft pubescent on both surfaces, bicolored, 
rounded to broad-cuneate at base, abruptly acute to obtuse at apex, the blades 
2-6 (10) cm. long, 1.5-5 cm. wide, with about 6 pairs of lateral veins; petioles 
slender, 1.5-3 (4) cm. long; flowers solitary, axillary, the plants dioecious; 
pedicels filiform, strigulose to pubescent, 1-1.8 cm. long; staminate flowers 
with hypanthium subcylindric, 7-10 mm. long, 2.5-3.5 mm. wide, puberulent, 
rose-orchid to deep red ; sepals red to orchid, puberulent, 2.5-3 (4) mm. long, 
ascending, deltoid, apiculate, the tips about 0.5 mm. long, somewhat separate 
in bud; petals rose-scarlet or paler, suborbieular, ca. 3 mm. long; stamens 
included, anthers 2 mm. long ; stigma included ; pistillate flowers smaller, 
constricted above ovary; hypanthium 5-6 mm. long, 1-2 mm. wide; anthers 
less than 1 mm. long ; style 8-9 mm. long, stigma exserted, 4-lobed, 1 mm. long ; 
fruit purplish or black, up to 1 cm. thick. 

Type locality "Guatemala," type collected by G. W. Skinner, April 4, 1846. 
I have not seen the type, but am taking the name tetradactyla for the plant 
recently called Seleriana. The large leaves (2.7 inches long, including peti- 



Vol. XXV] MUNZ : THE GENUS FUCHSIA 101 

oles), the calyx (^ inch long, that is hypanthium plus sepals) , pedicels equal- 
ing petioles, and origin Guatemala, all these points mentioned in the original 
description can indicate only this species. Material seen, MEXICO, without 
definite locality, '^Fuchsia tomentosa N.," Sesse, Mociiio, et al. 5207 (F). 
Oaxaca, between Suchistepec and Miahuatlan, Nelson 2515 (US) ; between 
Plunia and San Miguel Suchistepec, Nelson 2505 (US). Chiapas, Teopisca, 
Goldman 970 (US) . GUATEMALA : Baja Vera Paz, Santa Rosa, von Tuerck- 
heim under J. D. Smith 1315 (US). Quezaltenango, Volcan Zunil, Steyermark 
34741 (F) ; near Quezaltenango, Seler 2894, type of Seleriana (photos F, 
POM, specimen CAS) ; Quezaltenango, Skiitch 810 (F, NY) ; Volcan de 
Santa 'NLaria,8teijermark 34031 (F, POM). Chimaltenango, Tecpam, John- 
ston 761a (F), Skutch 489 (F, GH, MICH, US). Sacatepequez, Santa Lucia, 
Popenoe 689 (US) ; Volcan de Acatenango, Shannon under J. D. Smith 3656 
(US) ; Finca El Hato, ne. of Antigua, Standley 61202 (F) ; Santiago, Gomez 
under J. D. Smith 872 (US). Guatemala, Carmela, Morales 1100 (F). Jalapa, 
Cerro Alcoba, Steyermark 32512 (F, POM). Zacapa, Sierra de las Minas, 
Steyermark 29704 (F). 

(100) Fuchsia Mexiae Munz,, new species 
(Plate 15, fig. 91) 

Leafy shrub apparently rather large, ultimate twigs very slender, some- 
what reddish, somewhat fine-pubescent; leaves opposite, petioles slender, 
pubescent, 7-20 (25) mm. long; leaf -blades membranaceous, rhomboid-ovate 
to -obovate, bicolored, acute to acuminate at both ends, remotely sinuate- 
serrulate, subglabrous to minutely puberulent on both surfaces, especially 
on the veins and margins, 1.5^ (4.5) cm. long, 1-2 cm. wide ; flowers reddish, 
axillary, dioecious ; pedicels filiform, strigulose, about 1 cm. long; hypanthium 
of pistillate flowers tubular, strigulose, 3-4.5 mm. long, barely 1 mm. wide ; 
sepals ovate, 1.5-2 mm. long ; petals white, rounded, ca. 1 mm. long ; style 5-6 
mm. long; stigma deeply 4-lobed, well exserted, ca. 1 mm. long; staminate 
flowers not seen, but undoubtedly larger; fruit dark, 5-7 mm. thick. 

Frutex f oliosus ; ramulis tenuibus, rubescentibus, puberulentis ; f oliis oppositis ; petiolis 
tenuibus, pubescentibus, 7-20 (25) mm. longis; laminis membranaceis, rhomboideo-ovatis 
vel -obovatis, acutis vel acuminatis in extremis utrisque, remote sinuato-serrulatis, sub- 
glabris vel puberulentis in superficiebus utrisque, 1.5-4 cm. longis, 1-2 cm. latis; floribus 
rubescentibus, in axillis solitariis; pedicelis filiformibus, strigulosis, 1 cm. longis; hypan- 
thio florium pistillatorum tubuloso, striguloso, 3-4.5 mm. longo, 1 mm. lato ; sepalis ovatis, 
1.5-2 mm. longis; petalis albis, orbiculatis, 1 mm. longis; stylo 5-6 mm. longo; stigmate 
4-lobato, exserto, 1 mm. longo ; floribus staminatis inconnatis ; f ructu nigro, 5-7 mm. crassi. 

Type: from open dry pine forest, at 1500 m., Canon El Ranchito, San 
Sebastian, Jalisco, Mexico, Jan. 12, 1927, Ynez Mexia 1465, U. S. Herbarium 
no. 1319484 ; isotypes F, GH, CAS, MICH, NY. Another collection is from 
Morelos, El Parque, Rose and Rose 11118 (NY, US). The proposed species is 
much like F. striolata, but has membranaceous leaves, more rhomboid leaf- 
blades, white petals, and more western range. 



INDEX TO NAMES 
The number given after each name refers to the species number used in this paper. 



Brebissonia Spach, see Sect. 7, Encliandra. 

micropliylla Spach, see 93a. 

thymifolia Steud., see 85. 
Dorvalla eucharis Comm., see 3a. 
EUobium Lilja, see Sect. 2, Eufuchsia. 

fulgens Lilja, see 47. 
Encliandra Zucc, sect. 7. 

parvi flora Zucc, see 97. 

thymifolia Lilja, see 85. 
Fuchsia abrupta Johnston, no. 43. 

acynifoUa Scheidw. in Otto and Dietr., 
Allg. Gartenz., 15:226, 1847. I have 
seen no material so labeled; no type 
number was designated. It was said to 
have come from Mexico and the char- 
acters mentioned (small ovate leaves 
dentate toward the tips; calyx cylin- 
drical, 6 lines long; leaves with some 
puberulence above, glabrous beneath ; 
pedicels short) suggest F. microphylla 
H.B.K., no. 93. 

affinis Cambess., see 6c. 

alpestris Gardn., see 6d. 

alternans Moc. and Sesse, see 85. 

amoena Hort., mentioned in DC. Prodr., 
3:37, 1828 a ssynonym of F. arbores- 
cens. 

ampliata Bouth., see 21. 

Andrei Johnston, no. 54. 

apetala Kuiz and Pavon, no. 78. 

apiculata Johnston, see 16. 

aprica Lundell, see 93b. 

araucana F. Pliil., see 3a. 

arborea Sess6 and Moc, see 84a. 

arborescens Sims, no. 84. 
Fuchsia arborescens var. megalantha Donn. 
Smith, see 84a 
forma parva Munz, no. 84c. 
forma tenuis Munz, no. 84b. 
forma typica Munz, no. 84a. 

asperifnlia Krause, no. 59. 

Aspiazui Macbr., no. 33. 

Asplundii Macbr., no. 26. 

atrorubra Johnston, see 64. 

austromontana Johnston, no. 9. 

ayavacensis H.B.K., no. 21. 

bacillaris Lindl., no. 91. 

biflora Sesse and Moc, see 96. 

boliviano Britton, see 52. 



boliviana Carr., no. 46. 

f . puberulenta Munz, no. 46b. 

var. luzurians Johnston, no. 46c. 

var. typica Munz, no. 46a. 
Bracelinae Munz, no. 2. 
Brittoni Johnston, see 52. 
Campos-Portoi Pilger and Schulze, no. 1. 
canesccns Benth., no. 14. 
caracasana Field, and Gardn., see 40. 
cestroides Schulze-Menz, no. 83. 
chiapensis Brandg., see 96. 
chloroloba Johnston, see 73a. 
chonotica Phil., see 3a. 
coccinea Curtis, see 3b. 
coccinea Soland. in Ait., no. 4. 

var. chronotica Eeiche, see 3a. 
Colensoi Hook, f ., no. 68. 
colimae Munz, no. 88. 
confertifolia Field, and Gardn., no. 32. 
conica Lindl., see 3a. 
corallina Hort. ex Lynch, Gard. Chron., 

2:604, 1883; said by Lynch to be syn-j 

onym of F. exoniensis. 
Fuclisia cordi folia Benth., no. 8. 
cordifolia /3 Hook., see 7. 
corollata Benth., see 14. 
corymbi flora R. and P., no. 42. 
corymbosa Pritz., see 42. 
Cuatrecasasii Munz, no. 44. 
curviflora Benth., see 24a. 
cuspidata Fawcett and Eendle, see 46c. 
cylindracea Lindl., no. 98. 
cyrtandroides Moore, no. 67. 
decidua Standley, no. 72. 
decussata R. Grah., see 3b. 
decnssata R. and P., no. 48. 
denticulata R. and P., no. 10. 
dependens Hook., see 42. 
discolor Lindl., see 3a. 
dolichantha Krause, see 32. 
Bominiana Hort. ex. Fl. des Serres, ser. 1, 

95, pi. 1004, 1854-5 ; said to be a horti- 
cultural hybrid. 
elegans Paxt. ex. Steud.; see globosa var. 

elegans. 
elegans Salisb., see 4. 
Encliandra (spelled Encliandra) Steud., 

no. 97. 
excorticata L. f., no. 66. 



[102] 



Vol. XXV] 



MUNZ: THE GENUS FUCHSIA 



103 



exoniensis Paxt., Mag. Bot., 10 : 153, 1843 ; 
hybrid between F. cordifolia and F. 
globosa. 

filipes Kusby, see 52. 

Fischeri Macbr., no. 36. 

fontinalis Macbr., see 48. 

fulgens Moc. and Sesse, no. 47. 

furfuracea Johnston, no. 38. 

fusca Krause, see 48. 

Garleppiana Kuntze and Wittm., no. 81. 
Fuchsia Gelirigeri Munz, no. 30. 

glaierrima Johnston, no. 61. 

Glazioviana Taub., see 4. 

gloiosa Lindl., Bot. Beg., 18: pi. 1556, 
1833 ; apparently a plant of horticul- 
tural origin. I have seen no wild speci- 
mens like it; it apparently has arisen 
from F. magellanica. 
var. elegans Paxton, Mag. Bot., 4:75, 
1838 ; apparently horticultural. 

gracilis Lindl., see 3b. 
var. multiflora Lindl., see 3a. var. ten- 
ella Lindl., see 3a. 

gracilis Moc and Sesse, see 93a. 

grandiflora Euiz ex Dahlgren, see 10. 

hamellioides Moc. and Sesse, see 84a. 

Hartwegii Benth., no. 57. 

Hemsleyana Woodson and Seibert, no, 94. 

heterotricha Lundell, see 96. 

hirsuta Hemsl., no. 79. 

hirtella H.B.K., no., 40. 

Hitchcoclcii Johnston, see 21. 

liortensis Bergmans, Vaste PI. Eotshee- 
sters, 227, 1924 ; horticultural. 

hyhrida Hort., no. 5. 

hypoleuca Johnston, no. 49. 

inflata Schulze-Menz, see 7.3b. 

insignis Hemsl,, see 78. 

integrifolia Cambess., see 6a. 

intermedia Hemsl., see 7. 

involucrata Sw. Prodr. Veg. Ind. Occ, 62, 
1788, is Schradera Cephalotes of Eubi- 
aceae. 

involucrata K. and P., Field Mus. Nat. 
Hist., Bot. 21 : 212, 1940 ; nomen. nudum 

Jahnii Munz, no. 29, 

juntasensis Kuntze, no. 74. 

Killipii Johnston, no. 45. 

Kirhii Hook, f., no. 71. 

Lehmannii Munz, no. 55. 

Lenneana Warcz. in Otto and Dietr., Allg. 
Gartenz., 20:293, 1852; novien sub- 
nudum. 



Fuchsia leptopoda Krause, no. 11. 
Liebmanni Levi., see 84a. 
Llewelynii Macbr., no. 27. 
longiflora Benth., see 20a. 
Lowei Hort. ex L. H. Bailey, Standard 

Cycl. Hort., 1301, 1915, where given in 

synonymy under F. magellanica var. 

discolor, 
loxensis H.B.K., no. 16. 
lycioides Andr., no. 65. 
macrantha Hook., no. 82. 
macropetala Presl., see 42. 
macrophylla Johnston, no. 62. 
macrostema E. and P., see 3b. 

var. conica Sweet, see 3a. 

var. gracilis Sweet, see 3b. 

var. recurvata Hook., see 3b. 

var. tenella DC, see 3a. 
macrostigma Benth., no. 20. 

var. longiflora Munz, no. 20a. 

var. typica Munz., no. 20b. 
magdalenae Munz, no. 13. 
magellanica Lam., no. 3. 

var. conica Bailey, see 3a. 

var. discolor Bailey, see 3a. 

var. globosa Bailey, see F. globosa. 

var. gracilis Bailey, see 3b. 

var. inacrostema Munz, no. 3b. 

var. Molinae Espinosa, no. 3c. 

var. Biccartoni Bailey, see F. Riccar- 
toni. 

var. typica Munz, no. 3a. 
Mathewsii Macbr., no. 35. 
Mattoana Krause, see 74. 
membranacea Hemsl., no. 75. 
Mexiae Munz, no. 100. 
michoacanensis Sesse and Moc, no. 96. 
microphylla H.B.K., no. 93. 

var. aprica Munz, no. 93b. 

var. typica Munz, no. 93a. 
Fuchsia miniata Planch and Linden, see 40. 
minimiflora Hemsl., no. 87. 
minutiflora Hemsl., no. 92. 

var. hidalgensis Munz, no. 92a. 

var. typica Munz, no. 92b. 
mitifolia (mutifolia in index) E. and P., 

in Field Mus. Pub,, Bot., 21:80, 1940; 

nomen nudum, 
mixta Hemsl., see 93a. 
mollis Krause, see 6d. 
montana Cambess. ,see 4. 
multiflora Lodd., see 3a. 



104 



CALIFORNIA ACADEMY OF SCIENCES 



[Proc. 4th Ser. 



multiflora Murr., Syst. ed. 13, 299, 1774; 
nomen nudum. 

Munzii, Macbr., no. 39. 

myrtifolia Hort. ex Koehne, Deutsche 
Dendrol., 429, 1893 ; said to be near F. 
discolor. 

nigricans Linden, see 64. 

Notarisii Lehm., Hamb. Gartenz,, 8:459, 
1852; I have not seen authentic mate- 
rial; description suggests a member of 
section Encliandra. 

Osgoodii Macbr., no. 53. 

ovalis E. and P., no, 58. 

var. dberrans Macbr., see 53. 

ovata Moc. and Sesse, see 85. 

pallescens Diels, no. 17. 

paniculata Lindl., see 84c. 

parviflora Lindl., see 85. 

parviflora Zucc, see 97. 

pendula Salisb., see 4. 

pendulae flora Hort., Gard. Chron., 1:521, 
1877; hybrid. 

perirevis Johnston, see 51. 

persicandens Cockayne and Allan, no. 69. 

peruviana Carr. in Johnston, Contr. Gray 
Herb., 75:39, 1925, apparently a slip 
for F. boliviana. 
Fuchsia petiolaris H.B.K., no. 24. 
var. bolivarensis Munz, no. 24b. 
var. typica Munz, no. 24a. 

pilosa Field, and Gardn., no. 60. 

platypetala Johnston, no. 19. 

polyantJia Killip, no. 41. 

polyanthella Johnston, see 58. 

Pringlei Eobinson and Seaton, no. 86. 

Pringsheimii Urb., no. 22. 

procumbens Cunn., no. 70. 

prostrata Baill., see 70. 

pubescens Cambess., see 4. 

pubescens R. and P., in Field Mus. Nat. 
Hist., Bot., 21:212, 1940, nomen nudum. 

pulchella Woodson and Seibert, see 94. 

pumila Hort. ex. Muenissier, Rev. Hort., 
n.s. 20:173, 1926; horticultural. 

pumila Hort. ex Vilmorin, Blumeng., ed. 
3, Sieb. and Voss, 1 : 332, 1894, in synon- 
ymy under F. gracilis f. pumila Voss. 

punicea R. and P., Field Mus. Nat. Hist., 
Bot., 21:212, 1940, nomen nudum. 

putumayensis Munz, no. 56. 

pyrifolia Presl, see 6b. 

quinduensis H.B.K., see 24a. 



racemosa Lam., Encye. 2:565, 1788; pi. 
282, fig. 1, 1793. Given with F. triphylla 
as synonym, but as coming from Carta- 
gena, Colombia. It may be an earlier 
name for F. petiolaris, but I have not 
had access to a specimen since I dis- 
covered that fact. 

racemosa Sesse and Moc, PI. N. Hispan., 
ed. 1, 58, 1887-'90. Description given 
suggests F. fulgens, although it was 
referred to F. arborescens by G. Don, 
Gen. Syst., 2:677, 1832. Leaf at Field 
(ex herbario Musei Parisiensis) labeled 
"F. racemosa Lam." can well be F. ful- 
gens in size, veins, pubescence. 

radicans Miers, see 6b. 
Fuchsia recurvata Niven, see 3b. 

regia (Veil.) Munz, no. 6. 
var. affinis Munz, no. 6c. 
var. alpestris Munz, no. 6d. 
var. radicans Munz, no. 6b. 
var. typica Munz, no. 6a. 

Biccartonensis Bergmans, Vaste PI. Rot- 
sheesters, 227, 1924; see F. Riccartonii. 

Biccartoni Hort. ex Tillery, in Flor. and 
Pom., 217, 1871; horticultural name 
with inadequate description. 

Biccartoniana Hort. ex Koehne, Deutsche 
Dendrol., 429, 1893 ; horticultural name. 

rivularis Macbr., no. 15. 

robusta Tengbergen, Tuinb. FL, 2:345, 
1855; hybrid. 

rosea R. and P., see 65. 
var. spinosa Eeiche, see 65. 

salicifolia Hemsl., no. 76. 

sanctae-rosae Kuntze, no. 52. 

scabriuscula Benth., no. 50. 

scandens Krause, see 48. 

Scherffiana Andre, Rev. Hort., 1888:233 
and 268, from Ecuador ; would seem 
nearest to F. Townsendii Johnston, over 
which it has priority, but the description 
sounds as if it had narrower petals. I 
have not seen type material. 

Seleriana Loes., see 99. 

semperfiorens Rozain, Wien. Illustr. Gar- 
tenz., 1888:244; hybrid. 

serratifolia R. and P. ,see 10. 

serratifolia Hook., see 9. 

sessilifolia Benth., no. 63. 

simplicicaulis R. and P., no. 31. '^ 

siphonantha Krause, see 11. 



Vol. XXV] 



MUNZ: TEE GENUS FUCHSIA 



105 



STcutchiana Munz, no. 90. 
Smithii Munz, no. 25. 
Fuchsia speciosa Hort., see 5. 
spectaiilis Hook., see 20a. 

var. pubens Johnston, see 20b. 
spinosa Presl, see 65. 
splendens Zucc, no. 7. 
Standishii Paxton, Bot. Mag., 11 : 33, 1844 ; 

garden hybrid. 
Steinbachii Johnston, see 74. 
Storlcii Munz, no. 37. 
striolata Lundell, no. 95. 
sylvatica Benth., no. 64. 
syringae flora Carr., see 84a. 
tacanensis Lundell, no. 89, 
tacsoniiflora Krause, see 10. 
tenella Hort., see 3a. 
tetradactyla Lindl., no. 99. 
Thompsoni Hort. ex Koehne, Deutsche 

Dendrol., 430, 1893 ; nomen nudum. No 

description, 3 varieties figured. 
thymifoUa H.B.K., no. 85. 
tincta Johnston, no. 34. 
Townsendii Johnston, no. 18. 
tripliylla L., no. 23. 
tuierosa Krause, no. 73. 

var. inflata Munz, no. 73b. 

var. typica Munz, no. 73a. 
tunariensis Kuntze, no. 77. 
umhrosa Benth., see 16. 
unduavensis Munz, no. 80. 
uniflora Sesse and Moc, PI. N. Hispan., 

ed. 1, 58, 1887-'90. Unknown to me. 
velutina Johnston, see 42. 
venusta H.B.K., no. 28. 

var. huilensis Munz, no. 28b. 

var. typica Munz, no. 28a. 



Fuchsia verrucosa Hartw., no. 51. 

verticillata E. and P., in Field Mus. Nat. 

Hist., Bot., 21:46, 1940; nomen nudum. 

violacea E. and P., in Field Mus. Nat. 

Hist., Bot., 21 : 128, 1940 ; nomen nudum. 

virgata Sweet, Hort. Brit., ed. 1, 485, 1827 ; 

nomen nudum, 
vulcanica Andre, see 14. 
Weberbaueri Krause, see 52. 
WoytTcowsMi Maebr., no. 12. 
Kierschlegeria Spach, section no. 3. 
Lindleyi Spach, see 97. 
lycioides Spach, see ^^5. 
Lopesia thymifoUa Willd., see 85. 
Lyciopsis Spach, see Encliandra, sect. 7. 

thymifoUa Spach, see 85. 
Myrinia Lilja, see EncUandra, sect. 7. 

microphylla Lilja, see 93a. 
Nahusia Schneev., see Quelusia, sect. 1. 

coccinea Schneev., see 4. 
Quelusia Vandelli, section no. 1. 

regia Vand., see 6. 
Schufla Spach, section no. 6. 

arborescens Spach, see 84a. 
SJcinnera Forst., section no. 4. 

antartica Forst. said to be BlacTcweUia 

antartica Gaertn. 
coccinea Moench, see 4. 
exeorticata Forst, see 66. 
Spachia Lilja, in Traedgardtidning, 62, 
1840; ex Linnaea, 15:262, 1841. No 
species named; synonym of Fuchsia. 
Thilcum (or Thileum'?) Molina, see Nahusia, 
sect. 1. 
tinctorium Molina, see 3b. 
Tilco Adans., Fam. 2:498, 1763; no species 
named; synonym of Fuchsia, ace. to 
Kew Index. 



EXPLANATION OF PLATES 

Measurements given for flower are from base of ovary 
to tip of stigma, for leaf include petiole and blade. 

PLATE 1 
Fig. 1. F. Campos-Fortoi, drawn from Brade 15676 ; flower 32 mm. ; leaf 27 mm. 
Fig. 2. 7*'. Bracelinae, drawn from Mexia 4013 ; flower 42 mm. ; leaf 45 mm. 
Fig. 3. F. magellanica, from "Werdermann 61 ; flower 56 mm. ; leaf 53 mm. 
Fig. 4. F. coccinea, from Hoehne 6278; flower 41 mm.; leaf 41 mm. 
Fig. 5. F. regia, from Eegnell 794 ; flower 62 mm. ; leaf 66 mm. 
Fig. 6. F. splendens, from Standley 61808 ; flower 60 mm. ; leaf 80 mm. 
Fig. 7. F. cordi folia, from Skutcli 867 ; flower 83 mm. ; leaf 120 mm. 
Fig. 8. F. auiitromontana, from Pennell 14110 ; flower 56 mm. ; leaf 104 mm. 



[106] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



Cmunz] plate I 




[107] 



PLATE 2 

Fig. 9. F. denticulata, from Weberbauer 7587 ; flower 72 mm. ; leaf 103 mm. 
Fig. 10. F. leptopoda, from Weberbauer 6934; flower 80 mm.; leaf 107 mm. 
Fig. 11. F. W oyiTcoivskii, from Woytkowski coll. ; flower 58 mm. ; leaf 73 mm. 
Fig. 12. F. magdalenae, from Seifriz 392; flower 95 mm.; leaf 65 mm. 
Fig. 13. F. canescens, from Penland 623 ; flower 80 mm. ; leaf 55 mm. 
Fig. 14. F. loxensis, from Penland 959 ; flower 30 mm. ; leaf 68 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 2 




[109] 



PLATE 3 

Fig. 15. F. Townsendii, from Townsend A93 ; flower 75 mm.; leaf 128 mm. 

Fig. 16. F. platypetala, from West 3705 ; flower 78 mm. ; leaf 91 mm. 

Fig. 17. F. macrostigma, from Eose 22479 ; flower 83 mm. ; leaf 145 mm. 

Fig. 18. F. ayavacensis, do not have record of specimen ; flower 56 mm. ; leaf 
95 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 3 




[111] 



PLATE 4 

Fig. 19. F. Prm^rs/ietrnw, from von Tuerckheim 3151 ; flower 65 mm. ; leaf 25 mm. 

Fig. 20. F. triphylla, from Leonard 3681; flower 50 mm.; leaf 70 mm. 

Fig. 21. F. petiolaris, from Pennell 2379; flower 69 mm.; leaf 57 mm. 

Fig. 22. F. petiolaris var. iolivarensis, from Pennell 4324; flower 80 mm.; leaf 
76 mm. 

Fig. 23. F. Smithii, from Killip and Smith 17300 ; flower 75 mm. ; leaf 57 mm. 

Fig. 24. F. Asplundii, from Weberbauer 6054 ; flower 63 mm. ; leaf 75 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 4 







[ 113 ] 



PLATE 5 

Fig. 25. F. LZcwdi/nM, from Llewelyn Williams 7594; flower 68mm.; leaf 87mm. 

Fig. 26. F. venusta, no record of specimen from which drawn; flower 62 mm.; 
leaf 125 mm. 

Fig. 27. F. Jahnii, from Jahn 506 ; flower 60 mm. ; leaf 71 mm. 

Fig. 28. F. Gehrigeri, from Gehriger 322; flower 62 mm.; leaf 80 mm. 

Fig. 29. F. simplicicaulis, from Macbride 4014; flower 68 mm.; leaf 141 mm. 



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PROC, CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 5 




[1]5] 



I 



PLATE 6 

Fig. 30. F. con ferti folia, from Weberbauer 7153 ; flower 74 mm. ; leaf 17 mm. 
Fig. 31. F. Aspiazui, from Weberbauer 7042; flower 78 mm.; leaf 133 mm. 
Fig. 32. F. tincta, from West 7092 ; flower 44 mm. ; leaf 134 mm. 
Fig. 33. F. StorTcii, from Stork 10073 ; flower 41 mm. ; leaf 75 mm. 
Fig. 34. F. furfuracea, from Bang 674 ; flower 68 mm. ; leaf 100 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



CMUNZ]«PLATE 6 




[117] 



PLATE 7 

Fig. 35. F. Munzii, from Weberbauer 6648; flower 52 mm.; leaf 115 mm. 
Fig. 36. F. hirtella, from Cuatrecasas 6657 ; flower 64 mm.; leaf 125 mm. 
Fig. 37. F. polyantha, from Mexia 7571 ; flower 55 mm. ; leaf 69 mm. 
Fig. 38. F. corymbiflora, from Mexia 7765 ; flower 68 mm. ; leaf 148 mm. 
Fig. 39. F. dbrupta, from Macbride 4541 ; flower 58 mm. ; leaf 116 mm. 
Fig. 40. F. Cuatrecasasii, from Cuatrecasas 8738; flower 47 mm.; leaf 84 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 7 




[119] 



PLATE 8 

Fig. 41. F. Killipii, from Killip 16695; flower 62 mm.; leaf 102 mm. 

Fig. 42. F. boUviana, from Balls 6715 ; flower 84 mm. ; leaf 140 mm. 

Fig. 43. F. fulgens, from Pringle 4123 ; flower 150 mm. ; leaf 180 mm. 

Fig. 44. F. decussata, from Macbride and Featherstone 2124; flower 28 mm.; 
leaf 65 mm. 

Fig. 45. F. hypoleuca, from Hitchcock 21440 ; flower 42 mm. ; leaf 70 mm. 

Fig. 46. F. scabriuscula, from unknown source; flower 45 mm.; leaf 106 mm. 

Fig. 47. F. verrucosa, from Cuatrecasas 8465 ; flower 24 mm. ; leaf 65 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV, NO. 1 



[MUNZ] PLATE 8 




[121] 



i 



PLATE 9 
rig. 48. F. sanctae-rosae, from West 6414 ; flower 40 mm. ; leaf 96 mm. 
Fig. 49. F. Osgoodii, from Osgood and Anderson 47 ; flower 38 mm. ; leaf 99 mm. 
Fig. 50. F. Andrei, from Andre K 820 ; flower 45 mm. ; leaf 122 mm. 
Fig. 51. F. Lehmannii, from Lehmann 5498 ; flower 35 mm. ; leaf 140 mm. 
Fig. 52. F. putumayensis, from Garcia 4639; flower 28 mm. ; leaf 91 mm. 
Y\g.5i.F.Bartwegu, from Killip 6829; flower 37 mm.; leaf 130 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES, VOL. XXV. NO. 1 



[MUNZ] PLATE 9 




[123] 



PLATE 10 

Fig. 54. F. ovalis, from Webcrbauer 6721 ; flower 24 mm. ; leaf 189 mm. 
Fig. 55. F. glaberrima, from Hitchcock 21750; flower 42 mm.; leaf 161 mm. 
Fig. 56. F. viacrophylla, from Maebride 5616; flower 31 mm.; leaf 225 mm. 



[124] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL, XXV. NO. I 



[MUNZ] PLATE ID 




[125] 



PLATE 11 

Fig. 57. F. sessilifolia, from Mexia 7597; flower 30 mm.; leaf 138 mm. 

Fig. 58. F. sylvatica, from Dawe 769; flower 42 mm. ; leaf 131 mm. 

Fig. 59. F. lycioides, from Garaventa 1641 ; flower 14 mm. ; leaf 26 mm. 

Fig. 60. F. excortieata, from Petrie 4110 ; flower 40 mm. ; leaf 95 mm. 

Fig. 61. F. ciirtandroidea, from MacDaniels in 1927 ; flower 34 mm. ; leaf 130 mm. 



[126] 



PROC. CALIF. ACAD. SCI., 4TH SERrES. VOL. XXV. NO. 1 



[MUNZ] PLATE 1 1 




[127] 



PLATE 12 

Fig, 62. F. Colensoi, from Petrie 4109 ; flower 28 mm. ; leaf 48 mm. 
Fig. 63. F. perscandens, from Allan in 1924; flower 32 mm.; leaf 38 mm. 
Fig. 64. F. procumiens, from Meebold 25160 ; flower 17 mm.; leaf 40 mm. 
Fig. 65. F. decidua, from Mexia 1601 ; flower 45 mm. 
Fig. 66. F. tuberosa, from Pennell 13973 ; flower 82 mm. ; leaf 135 mm. 
Fig. 67. F. juntasensis, from Steinbach 5038 ; flower 82 mm. ; leaf 91 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 12 





[129] 



PLATE 13 

Fig. 68. F. tiinariensis, from Cardenas 3369; flower 91 mm.; leaf 76 mm. 

Fig. 69. F. apetala, from Hitchcock 21701; flower 72 mm.; leaf 108 mm. and  
from Eose 22230. 

Fig. 70. F. hirsuta, from Weberbauer 6975; flower 67 mm.; leaf 131 mm. 

Fig. 71. F. unduavensis, from Buehtien 2925; flower 80 mm.; leaf 88 mm. 



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PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 



[MUNZ] PLATE 13 




[131] 



PLATE 14 

Fig. 72. F. Garleppiana, from Cardenas 688; flower 152 mm,; leaf 153 mm. 
Fig. 73. F. macrantha, from Cook and Gilbert 1312; flower 136 mm.; leaf 87 mm. 
Fig. 74. F. cestroides, from Weberbauer 6423 ; flower 35 mm. ; leaf 111 mm. 
Fig. 75. F. arhorescensva.T.typica, from Skutch59; flower 15mm.; leaf 125mm. 



[132] 



=ROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. I 



[MUNZ] PLATE 14 




[133] 



PLATE 15 

Fig. 76. F. thymifoUa, from Pringle 9794; flower 11 mm.; leaf 21 mm. 

Fig. 77. F. Pringlei, from Pringle 4140 ; flower 7 mm. ; leaf 20 mm. 

Fig. 78. F. colimae, from Jones 224; flower 6 mm.; leaf 20 mm. 

Fig. 79. F. minimiflora, from Rose and Hay 5521 ; flower 5 mm. ; leaf 14 mm. 

Fig. 80. F. tacanensis, from Matuda 1686; flower 13 mm. ; leaf 80 mm. 

Fig. 81. F. STcutchiana, from Skutch 812; flower 7 mm.; leaf 75 mm. 

Fig. 82. F. bacillaris, from Hort. Bot. Petrop., 1829; flower 16 mm. ; leaf 30 mm. 

Fig. 83. F. minutiflora var. hidalgensis, from Pringle 8807 ; flower 11 mm. ; leaf 
14 mm. 

Fig. 84. F. microphylla, from Pringle 4314; perfect flower 18 mm., leaf 17 mm. ; 
pistillate flower 10 mm., from Hartweg 285. 

Fig. 85. F. Hemsleyana, from Davidson 876 ; flower 11 mm. ; leaf 15 mm. 

Fig. 86. F. striolata, from Matuda 4004 ; flower 7 mm. ; leaf 95 mm. 

Fig. 87. F. michoacanensis, from Standley 67188; perfect flower 11 mm., leaf 
40 mm. ; pistillate flower from Standley 58934, 8 mm. 

Fig. 88. F. Encliandra, from Mexia 2718 ; staminate flower 13 mm., leaf 25mm. ; 
pistillate flower from Nelson 1803, 12 mm. long. 

Fig. 89. F. cylindracea, from Arsene 6523 ; flower 15 mm. ; leaf 65 mm. 

Fig. 90. F. tetradactyla, from Popenoe 689 ; staminate flower 13 mm., leaf 40 
mm.; pistillate flower from Skutch 810, 15 mm. 

Fig. 91. F. Mexiae, from Mexia 1465 ; flower 7 mm. ; leaf 75 mm. 



[ 134 1 



PROC. CALIF. ACAD. SCI.. 4TH SERIES, VOL. XXV. NO. 1 



[MUNZ] PLATE 15 




[ 135 ] 



PLATE 16 
(Photographs by Dr. G. Dallas Hanua) 

Fig. 92. Photograph of title-page of Plumier's "Nova Plantarum americanarum 
Genera." 

Fig. 93. Photograph of Plate 14 of same with first illustrations of the Genus 
Fuchsia. 

Figs. 94 and 95. Photographs of pages 14 and 15 of same containing the descrip- 
tion of Fuchsia by Plumier. 



[136] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 1 

92 
NOVA 

PLANTARUM 

A M E R I C A N A R U M 

GENERA. 

Authore P. Carolo PluMIER Oidinis 
Minimorum in Fiovincia Francix, &C apud Infulas 
AiTiericanasBotanico Regio. 



[MUNZ] PLATE 16 

93 




PARISIIS, 

Apud JOANNEM BOUDOT, Regis Si Rcgi* Sclcnnirunv 
Aadema- Typographum > via Jacobxa.ad SoIldi A.iicjm. 

M. D C C 1 1 1."""" 

CVM r RIV 1 LE G I O REGIS. 




la*. 11. 



ViUi'ir 
Mm. 



Vt. M. 



94 

14 NovxPlant. 

mus Viofioridis ^ fatlidjfmtts inda^aior fUntdritfH nort dnees 
co^nttarvm fuit. yid qHoa pater exemplo Inflammavu , tjm eum 
ab inndcun^thulis inter iffu hcrbas 0" fores zoiutt educart. Rome 
moritur I544. £tat. 19. Edidit HiJ^oridftirpiuni Lib. ^. Extant 
Argentine ijtfj. lal. 

T R A G I A. 

TRagia eft plantar genus flore A monopecilo , infundibu- 
li toimi , pIcru!Tiqiietnfa,riamdiviro, fed fterili. Embryoncs 

cnim C a floribus fcjunguntur in cadem plsnta &:abcunt dcin- 

de in frutftum D tricoccum , ex tribus fcilicet capfulis E compo- 

Ctum , femine foccis fphartico F. 
Tragii fpecles func. 

Tragia fcandens longo Becontcx folio. 

Tragia alia fcandens uriicx folio. 

Hitronimus Tragus alias Hifronimus Bo^ius vemdcuJe Sock, dc 
Meidejpach , na:us cnim eji HeideJpachUfub an. 1 498. B:ponti w- 
xic per alitjuot artnos uhl hcrrum Ducis Ludovid Palatini Jtheni 
varji generis plantis injlruxit. Inde Harnhatum^ poj^ea Saraponturrt 
fc confer em Htjloriam hcrbarum condidit, in e^ua in tres Ltbros divi- 
fa de Jiirpium dijferenriis & facutiatibus fufe dijferir. Mortuits^ 
eft autem an. 1554. 21 Febr. £tat. ^6. txtar o^us jirgentorad 
IJJI. in ^u^rr. 

Fuchsia. 

FUchfiaeft plmtx genus flore A monoperalo , infundibuli- 
forrci & mulu-fido ; cujus calyx B abic delnde in fruftuiu 

C fubrorundum , mollem , carnofum , in quacuoi loculamenta 

divifum D, fcmimbjlque factum fubrocundis E. 
Fuchfix unlearn fpccietn agnovi. 

Fuchfia iriphylla , flore coccineo. 

leonartuj Furhfus Vembding£ , Rhxda Oppido in dicione Vucnrri 
Baviridt natui 1501. Mcdlcin£ Do6lor renuncia/us Monachum 
adiit t deinde Tubir.^m 1 ubi 35. annis prxcUre docuit. Vir 
fwt d^idul (abo/is , piantarum Cermanit dili^ens explordtor. 
Magna litrbdriorHm comrnodo hdrum iconcs jio. ampUorit for- 



A M E R r c. G E N E a. A. ty 

mt exhibttlt. Tiibing£ mortuus eft anno ij65. lo. Maji , dta- M*Uhm 
tis 6j. Scripfit de Hiftoria ftirpium Commtntari^sinfignss \ Bd"'^^^ 
files i|4i< in folio* 

RONDELETIA. 

ROndeletia eft p!antar genus fiore A raonopetalo , hypocra-r**. m 
tcri-formi , cubulaco 8c calyci C infidcnce. Is autem ca- 
lyx abit deinde in fruiium D fubrotundum > coronatum , bicap* 
fularem E > & feminibus factum exiguis F. 

Rondeletis unicam fpeciem novi. 
Rondcletia arborefcens , tini facie. 

Cuillflmus Rondelctius MonfpeJJiiU natus > dnno 1^07. die 17. 
Sept. Medicind arte prtceilem , de pijcibus &• de pifcium natu- 
Yd pr£cUrum Opus condidit 1 fed etiam fmplicibus medicament 
tis dignofcendis ^ inveniendis fumtnam operam coUocavit , at- TamffAn^, 
que in eo multum excelluit , Diofcoridrm primus MonfpeUt enar-* ' '*'" 
rans. Plura in Diofcoridem Jcripffte apparet , ex Epiftch's GeJ^ 
neri ad I. Bauhinum , a C. Bauhino edltts. Cancellarius ScnO' 
Id Monfpelienfts mortuus eft anno 1^66. 

T U R N E R A. 

TUrnera eft plants genus flore A monopctalo , inftindibu- Tab. n: 
H-formi fie multi-fido ; ex cujus calycc C bicoini furgjc 
piftillum infima: flciis parti B , ad inftar clavl intixum . quod 
deinde abic in frudum D fere globofum aut turbinatum , trila- 
riam dehifcentem E , feminibufquc fcctum F fubrotundis , te* 
Duibus capillamemis , ceu placenta: affixis. 

Turners unicam vidi fpecism. 
Turaera fi-utefccns ,- ulmi-folia. 

Cuiltelmus TumerHS yinglus y Medicine VoSlor , vir JoUdt eru- jtalut fr-fl 
ditionis (St- judtcli ,. emijn plantarum Hlftoriam jfngitt , anno f^'**- 
1^51. in qua Flgurds Fuchfii plerumque adhtbitit t nomlna expref 
fit latlne , grdce, dngUce » germankc > gdllicc , ordinem atplutm 
heticum JecHtus. 



95 



[137] 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXV, No. 2, pp. 139-146 Jiiiie 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 2 

RELATIONS OF THE TEMPERATE FLORAS OF 
NORTH AND SOUTH AMERICA 

BY 

DOUGLAS HOUGHTON CAMPBELL 

Professor of Botany, Emeritus 
Stanford University 

THE DiSTRiBUTiox of the existing land floras of the world, as well as their 
fossil record, have been studied by many botanists for a long time. These 
botanists have been, for the most part, Europeans or Americans, and have 
especially studied the floras of the Northern Hemisphere ; and the history of 
the Holarctic vegetation is fairly well known. There is much less information 
concerning the history of the floras of the Southern Hemisphere, especially 
the fossils. Since we now have as allies most of the South American countries, 
it might be well for us to become better acquainted with the botanists of South 
America and their studies of the floras of those countries. 

Great changes have occurred during past ages in the distribution of land 
and water of the globe; and great mountain ranges now stand where once the 
sea covered the land. These changes in the distribution of the main bodies of 
land and water have undoubtedly had much to do with fluctuations of climate 
and the distribution of many plants. 

Thus the invasion of the sea in the Cretaceous Age over much of the west- 
ern United States, and later, the elevation of the Rocky Mountains and the 
Sierra Nevada and Cascades, had a very marked effect on the climate of those 
regions and the distribution of the vegetation. During the Mid-Tertiary, for 
example, the redwood (Sequoia) now practically restricted to California, 
occurred over much of North iVmerica and Eurasia, and the southern cypress 
(Taxodium) now confined to the southeastern States was equally widespread. 

The great Pleistocene glaciation caused a great redistribution of the floras 

[ 139 ] 




140 CALIFOEXIA ACAD f: MY OF SCIENCES [Proc. 4th ser. 

of North America and Eurasia, and many American trees, like the tulip tree 
(Liriodendron), magnolia and walnut, before the Ice Age lived also in Eu- 
rope but were completely exterminated there at that time. 

Existing Factors ix Distribution 

Among the factors of prime importance in the distribution of the existing 
floras, the first is, perhaps, the relations of the great continental masses to 
each other, and to the surrounding oceans. The latter are effective barriers to 
the migration of all but a very few plants, and high mountains and extensive 
areas of desert and arid regions such as occur in the center of the continents, 
also are barriers to migration ; but mountains may also serve as liigliAvays — as 
may be seen in the occurrence of arctic and sub-arctic species in the Sierra 
Nevada and Rocky Mountains, far south of their normal habitat, and many 
plants are common to Chile and California. 

The major part of the land surface of the globe is in the Northern Hemi- 
sphere and comprises North America and Eurasia. It is generally recognized 
that these two continents have for a very long time been more or less inti- 
mately connected. This is indicated by the evident close relationship of their 
floras, especially in the higher latitudes. Throughout the Arctic and Sub- 
arctic Zones most of the trees, shrubs and herbaceous species are closely re- 
lated, or even identical. It has even been proposed to consider North America 
and Eurasia as parts of an ancient primary continent, "Laurasia." 

The southern continents. South America, Africa, Australia and Antarctica, 
are separated from each other by the width of the great oceans. Except for 
Antarctica which is destitute of terrestrial vegetation, the southern continents 
have varied and peculiar floras, indicating a long period of isolation. Never- 
theless they all have many obviously related genei-a, and even species, point- 
ing to some ancient land connections. 

It has been held by some geologists that in the late Palaeozoic there existed 
a great southern continent, "Gondwana," which included all the existing 
southern continents, and also India. Gondwana was separated from the north- 
ern land-masses by a broad oceanic belt, the "Tethys Sea" which persisted to 
near the end of the Mesozoic. This perhaps explains the radical differences 
between the present temperate floras of North and South America. 

How the existing southern continents became so completely isolated has 
caused much speculation. In a recent study of the subject by a South African 
geologist,' the author concludes that Gondwana split into segments which 
drifted apart and developed into the present continents. This theory of Con- 
tinental Drift, put forward by Wegener in 1912, is a plausible explanation of 
many puzzling facts in geographical distribution, both of animals and plants. 

The greater part of North America and Eurasia lies north of the Tropic of 
Cancer, and extends beyond the Arctic Circle. Almost the whole of the United 
States lies in the "Temperate" Zone, between 30° and 50° N. For the most 

^ Our Wandering Continents. .\. T.. r>n Toit : London, 1937. 



Vol. XXV] CAMPBELL: FLORAS OF NORTH AND SOUTE AMERICA 141 

part the climate is distinctly "continental," with great annual range of tem- 
perature. In general, winter is a season of complete cessation of plant activity. 
In the United States milder climates prevail in the Southern States and espe- 
cially on the Pacific Coast, where weather conditions are very different from 
those over most of the country. From Sitka to San Diego, the climate is 
remarkably equable. The January isotherm of 0°C, which on the Atlantic 
Coast is in the region of New York and Philadelphia, on the Pacific Coast is 
pushed north to Sitka (lat. 57° N.). In San Francisco, there is only a differ- 
ence of ten degrees Fahrenheit between the coldest and warmest months (50° 
to 60°) ; in Washington, D.C., about the same latitude (32° to 78°). 

The lofty barrier of the great Cordillera protects the coastal areas from the 
great temperature fluctuations of the interior regions, and the prevailing 
westerly winds from the Pacific have a great influence on both temperature 
and precipitation. 

In the strictly arctic regions, e.g., Greenland, northern Canada and Alaska, 
trees are absent and the vegetation is made up for the most part of perennials, 
either herbaceous species with subterranean roots or root-stocks which during 
the brief growing season of perhaps two months, complete their season's 
growth; or a few woody species, e.g., dwarf willows and birches, dwarf rhodo- 
dendrons, cranberries and a few other bog-plants may be found. In specially 
favorable places Iceland poppies, buttercups, saxifrage, with showy flowers,- 
as well as some grasses, quickly develop from their perennial roots during the 
very brief summer months. 

Further south in the Sub-arctic Zone, trees begin to play an important role. 
Thus in Canada, Scandinavia, Russia, there are extensive forests of spruce 
and pine, birches, willows and poplars and various deciduous shrubs, much 
the same throughout, from Alaska to Russia. 

The Sub-arctic merges gradually into the North Temperate which occupies 
much of North America and Eurasia. In the United States, especiallj^ in the 
eastern portion, this zone is characterized by an extraordinary variety of 
deciduous trees — oaks, beech, ash, elm, walnut, hickory, chestnut, and many 
others, which occur over most of the region east of the Mississippi. 

About two-thirds of the United States lie between the Rocky Mountains and 
the Atlantic Ocean. This includes the great Mississippi Valley and the Great 
Lakes. The vast area between the Rocky Mountains and the Appalachians has 
no barriers to plant migration except climatic ones, e.g., temperature and 
rain-fall, and many species are distributed over most of the area. Thus several 
species of oak, the black walnut and American elm occur over most of the area 
eastward of the Rocky Mountains, except where the moisture is insuffieient. 

The rainfall decreases westward and the forest gives way to the prairies. 
West of the Mississippi trees usually occur only along streams. The prairies 
finally pass into the semi-arid steppes which occur at the base of the Rocky 
Mountains. These plains are characterized by scattered bunch grasses and low 
shrubs, together with some herbaceous species. 



LIBRARY a 



142 CALIFOBXIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

The "western tliird of North America is very different from the East. It is 
a region of mountains and elevated plateaus between the Rocky JMountains 
and the Pacific Coast. This region includes extensive plateaus and secondary 
mountain ranges. Much of the plateau region is arid and in some places, as 
in the Salt Lake region, and parts of Nevada, true deserts. 

The great Pacific Cordillera which, with few breaks, extends from Alaska 
to Patagonia, sharply sets off the Coastal Region from the rest of the Conti- 
nent. The whole of the western Mountain area is very different climatically 
and floristically from Atlantic North America, and is the only portion of the 
United States in which the highest mountains have perpetual snow and a true 
alpine flora. 

Southern California and western Mexico have very light rainfall and much 
of the country is desert, but the precipitation increases rapidly northward, 
and in northwestern California the rainfall is very heavy; and from central 
California northward through Oregon, and Washington to Alaska, there is 
heavy rainfall, resulting in the development of the greatest coniferous forest 
in the world. 

In the Gulf region, especially Florida, there are many tj'pes, like the palms, 
Ficus and Bromeliads, evidently derived from the tropical regions to the 
south. 

Central California is a meeting ground for the boreal floras of Noi'tli 
America and the subtropical floras of Mexico. There is also a considerable 
number of genera and even species common to California and central Chile. 

The coastal ranges of central California, with their cool, moist forests, con- 
tain many distinctly boreal types, like most of the conifers, as well as such 
deciduous trees as maples, oaks and alders, and deciduous shrubs, honey- 
suckle, roses, elder, spiraea, azalea, rhododendron, dogwood, and others, and 
mau}^ characteristic herbaceous flowers, e.g., Trillivm, Erythronium, Ranun- 
cnlns, Oxalis, saxifrage, AquiJegia, violets, and many others. 

With decreasing precipitation southward, these northern types are gradu- 
ally replaced by genera and species absent from the eastern States, or only 
occasionally found, e.g., Acacia, cacti, Yucca, Prosopis, Mimtihts, Penstemon, 
Castilleja, and others. 

The occurrence in California of a considerable number of species found also 
in Chile is remarkable. Among these may be noted Fragaria chilensis, Pro- 
sopis juliftora, Mesemhryanthemum aequilaterale, Calandrinia Menziesii. 

The temperate regions of South America include a small part of southern 
Brazil, Paraguay, Uruguay, Argentina, and Chile ; the two latter comprise 
the greater part of temperate South America. Between Chile and Argentina 
is the great barrier of the Andes, a condition similar to that separating the 
coastal region of California from Nevada and Arizona, and there is much the 
same difference between the coastal and inland climates in both cases, as well 
as in the vegetation. 

The central and southern parts of Chile have abundant rainfall, while in 



Vol. XXV] CAMPBELL: FLOBAS OF NOBTH AXB SOUTH AMERICA 143 

most of Argentina the precipitation is scanty, and much of the country is a 
desert. The continent diminishes in breadth southward, its southernmost 
point, Cape Horn, being less than 60°S. 

Owing to the proximity of the oceans, the climate is insular rather than 
continental, and there is nothing comparable to the Arctic Zone of the north- 
ern continents. Similar conditions prevail in temperate South Africa and 
Australia, where the southernmost regions are under 50°S. Excepting in the 
high mountains there is no such difference between winter and summer as ex- 
ists in corresponding latitudes in the Northern Hemisphere. Thus in Ushuaia, 
in Tierra del Fuego (lat. 55.5°S) the coldest monthly average is 28.9°F, the 
warmest 50° ,• at Nain (Labrador) in about the same latitude North, the range 
is -7°F to 46°; in Buenos Aires (lat. 35.5°S) the coldest month is 50°F, the 
hottest, 73.6°. St. Louis, about the same latitude North, has the range of 31° 
to 79.1°. 

As a whole the temperate floras east of the Andes are very poor compared 
with corresponding areas in North America. While the flora of Argentina is 
mainly related to the Brazilian regions to the north, there are also elements 
related to those of Chile, especially in the extreme south, where there is abun- 
dant rainfall, and where the so-called "Subantarctic" flora becomes evident. 
This flora is also related to that of New Zealand and South Australia. 

The transition from southern Brazil to Argentina is a gradual one with 
increasing aridity southward. The strip at the base of the Andes is very dry, 
with scanty vegetation, and might be compared to the most extreme deserts 
of Arizona and Nevada. It has been described as one of the most dismal and 
sterile regions that one could imagine. The precipitation increases somewhat 
toward the east, and much of central Argentina is occupied by the great 
pampas, recalling the great plains of the central United States, and like the 
prairies there is a great development of grasses and other herbaceous plants. 
The constituents of the pampas floras are for the most part quite different 
species from those in North America. In general the flora of northern Argen- 
tina is evidently related to that of southern Brazil. 

Southward the pampas merge into the arid, sterile plains of Patagonia, 
whete only such scanty vegetation is found as can survive the stony, exposed, 
and excessively dry conditions which prevail. 

The conditions in Chile, west of the Andes, are very different from those 
in Argentina. The coast of Chile extends from 16° 57' S to 55 S (the southern- 
most point in South America — Cape Horn). This corresponds to the north 
Pacific Coast from Lower California to Alaska. 

The northern coast of Chile is practically rainless, and much of the region 
is quite destitute of vegetation. Passing southward, the precipitation gradu- 
ally increases, and in the central region, e.g., Valparaiso, Santiago, conditions 
are very similar to those in Central California, and the general character of 
the vegetation, both native and cultivated, is very much the same. 

Trees and shrubs from California, e.g., Sequoia, Monterey Cypress, Ceano- 



144 CALTFOFXIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

thus, are planted in the parks and gardens, and the California poppy {Esch- 
scholtzia) is quite naturalized. In California the Chilean pepper tree {Schinus 
molle), and Araucaria imhricata, are familiar trees, and Escalloriia spp. and 
Buddleia glohosa are common ornamental shrubs. 

Central Chile, both in climate and topography, much resembles the coi-re- 
sponding regions in California, and the great central valley between the 
Andes and the Coast ranges may be compared with the great central valley 
of California. Santiago (lat. 35°S), the capital, resembles almost exactly in 
its rainfall and mean temperature, San Jose (lat. 37°N) in the Santa Clara 
Valley of California. 

The rainfall in southern Chile is very heavy and there is present a luxuri- 
ant rain forest very much as the heavy coniferous forest is developed on the 
northern Pacific Coast. The constituents of these forests are very different ; 
while the northern Pacific forests are composed almost exclusively of con- 
ifers — redwood, spruce, fir, hemlock, etc. ; conifers are relatively rare in the 
Chilean forests and are all different from the northern genera, and are most 
nearly related to those of New Zealand and Australia. The rain forest of Chile 
is mostly composed of broad-leaved evergreens, mostly lacking in the northern 
forests, and includes many genera common to New Zealand, e.g., evergreen 
beech (Xothofagus), Drimys (Magnoliaceae) , Laurelia and Weinmannia, both 
important genera in New Zealand, and also Fuchsia and Griselinea. Charac- 
teristic of the forests of the South Temperate Zone are many species of the 
Myrtle family, almost completely absent from the North Temperate Zone, 
Eucalyptus from Australia being perhaps the best known example. Another 
exclusively southern family, Proteaceae, especially developed in South Africa 
and Australia, is well represented in South America. Emhothrium and Lo- 
matia occur in Chile. 

While there is nothing corresponding to the Arctic Zone of the Northern 
Hemisphere, southern Chile has a flora which is unmistakably related to that 
of New Zealand and southern Australia. This flora has been called "Subant- 
arctic" and indicates some former land connections with New Zealand and 
southern Australia. 

While, in general, there is nearly a complete absence in South America of 
characteristic North American genera and species, in the Pacific Coastal Re- 
gion there is a remarkable number of genera and even species common to 
Chile and central and southern California, and the adjacent southwestern 
areas. These may be explained as due to migration along the great mountain 
system of the Pacific Coast, subsequent to the union of South America and 
Central America. 

Some of these are cosmopolitan tyi)es — like Caltha, Linnm, Geranium, 
Oxalis, but a considerable number are species common to California and Chile 
but unknown elsewhere, e.g., Fragaria Chilensis, Calandrinia Menziesii, Lupi- 
nus microcarpa, Madia sativa, Plantago magellanica, Gilia pusilla, Prosopis 
juH flora. 



Vol. XXV] CAMPBELL: F LOEAS OF NOETH AND SOLTR AMEEICA 145 

Fossil Evidence 

While the fossil record is necessarily incomplete it has added much to our 
knowledge of the history of the higher plants, especially in North America 
and Eurasia. 

The fossil remains of plants are restricted largely to such parts, e.g., stems, 
leaves, roots, as have firm tissues, resistant to decay. Occasionally the tissues, 
even the more delicate ones, are replaced by infiltration of mineral substances, 
lime or silica, which replace the organic matter of the cell-walls, and result 
in petrifactions ; but as a rule the more delicate organs, such as the flowers, 
are very seldom found as fossils. 

Much has been learned of the fossil floras of the geological eras, from the 
Devonian where the earliest certain remains of land plants have been found, 
to the Mesozoic and Tertiary formations, which contain many remains of 
plants, evidently closely related to living species. 

From a study of these ancient fossils it is evident that as a whole plants are 
very conservative, and a remarkably large number of forms have come down 
from a very remote past to the present with little change. The common "horse- 
tail" {Equisetum) and the "club-mosses" (Lycopodium) are very similar to 
some of their relations from the Carboniferous, and many fossils of the mod- 
ern flowering plants (Angiosperms) as far back as the Cretaceous, when they 
are first certainly recognizable, can be referred to such living genera as syca- 
more (Plat an us) , sassafras, oak, poplar, and many others. It is to be expected 
that many common herbaceous plants which now are associated with these 
woody species also existed, but owing to their delicate perishable tissues have 
not survived as fossils. 

Many of these fossils occur in regions now unsuited to their growth, showing 
that great climatic changes have taken place. Thus among the earliest Angio- 
sperms that have been found were some described from Greenland, from rocks 
supposed to be of Cretaceous age. These fossils represent genera now found 
in regions many degrees southward. Among these fossils were sycamores, 
magnolias, cinnamon, and others. Sycamore (Plafanus) is now represented 
in the United States by two species, one occurring from New England to the 
Mississippi, the other in central and southern California. It is hard to im- 
agine any possible conditions of climate in which these trees could have grown, 
within a few hundred miles of the North Pole, deprived of sunlight for many 
months. A similar problem is offered by fossils found near the Antarctic. 

While our knowledge of the fossil floras of South America is much less com- 
plete than that of North America, there have been some important contri- 
butions, especially Professor E. W. Berry's studies in the Tertiary floras of 
Argentina (Tertiary Flora from the Rio Pichileufu, Argentina, Proc. TJ. 8. 
Nat. Mu.se7(m, Vol. 72, 1928). 

This flora includes elements now belonging to the present subtropical floras 
of Brazil, Paraguay, and also genera now confined to the extreme south of 



140 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Patagonia, Tierra del Fuego, and Chile, representing the "Snbantarctic" 
regions. This indicates that the Tertiary elimate in Patagonia was both 
warmer and more humid than at present. There were no North American 
genera present, but the most remarkable case was Ginkgo, unknown else- 
where in South America. 

A recent study of fossil conifers from Chile (The Tertiary Fossil Conifers 
of South Chile and their Phytogeographical Significance) by Rudolph Florin, 
Stockholm, 1940, shows that the genera were the same as now exist, and that 
none of the characteristic North American genera, e.g., Pinus, Abies, Tsuya, 
Picea, were present. Florin states that the same complete separation of the 
northern and southern types has existed since the late Palaeozoic. 

From a study of the existing floras as well as the fossil forms, one may 
conclude that the floras of South America have developed independently,, 
except for the so-called "Subantarctic" floras, which probably originated in 
some common land-mass, perhaps the hj'pothetical Gondwana. This seems 
much more likely than that they originated in some northern extension of 
the present Antarctica. 

Acceptance of the recent hypothesis of Du Toit, that there were two pri- 
mordial continents, Laurasia in the Northern Hemisphere and Gondwana in 
tlie South, and from these primary continents, the existing continents were 
separated and shifted to their present positions, would, if true, remove most 
of the difficulties in explaining the present distribution of many existing 
l^lant families. 



I 



PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXV, No. 3, pp. 147-170, pi. 17 Ji^ii^e 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 3 
PUGILLUS ASTRAGALORUM ALTER 

BY 

R. C. BARNEBY 




THE OBSERVATIONS on Astragalus contained in the present paperf are based 
primarily on material gathered by Mr. H. D. Ripley and the writer in 
California, Nevada, Utah, and Arizona during the past two or three years. 
Through the kindness of Dr. P. A. Munz it has been possible to make critical 
comparisons with a number of types in the herbarium of Marcus E. Jones 
at Pomona College (PO), without which these notes could not have been pre- 
pared. In addition, the writer has had an opportunity to visit several herbaria 
in the East, and there to study type and other specimens not represented in 
California collections. To the gentlemen in charge of these herbaria, in par- 
ticular to Prof. M. L. Fernald of the Gray Herbarium at Harvard Universitj^ 
(G), Dr. H. A. Gleason of the New York Botanical Garden (NY) and Dr. 
P. C. Standley at the Field Museum (F), who generously placed at my dis- 
posal the collections in their care, I wish to express my lively appreciation. 
I am also especially grateful to Miss Alice Eastwood for the communication 
of the type specimen of Astragalus Bryantii, described below. The capital 
letters in parentheses have been used throughout the paper to denote the 
herbarium in which cited material has been seen. 

Types of all species and varieties described as new are deposited in the 
herbarium of the California Academy of Sciences together with duplicates 
of the collections discussed. 

t Mr. Barneby's earlier paper on Astragalus. "Pugillus Astragalorum Nevadensium," 
was published in" Leaflets of Western Botany 3 : 97-114, figs. A-F. 1942. 



[147 1 



148 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 



Astragalus diversif olius A. Gray 

Astragalus diversif olius A. Gray, Proc. Amer. Acad. 6:230. 1864. HomnJohus orthocarpns 
NuTT. ex T. and G., Fl. N. Amer. 1:351. 1838. A. convallarius Greene var. diversifolius 
(A. Gray) Tidestr., Proc. Biol. Soc. Wash. 50:20. 1937. 

Utah: clay slopes among junipers, 3 miles south of Enterprise, Washington Co., alt. 
6300 ft. Ripley and Barnehy 4967. 

This station extends the range of tlie species as known to the writer almost 
to the borders of Arizona, to which state it is accredited by Tidestrom (Fl. 
Ariz, and N. Mex. 197. 1941), thouyh not admitted by Kearney and Peebles, 
who record only A. jnnciformis A. Nels. of this immediate group as native 
to the state. Prof. Tidestrom's plant is unknown to me, but his combination 
is in any case untenable. If the plants originally named by Nuttall as Homa- 
lobns orthocarpus and H. campestris cannot be separated specifically, the 
oldest valid binomial for either element of the complex must be given the 
rank of species, not reduced to varietal status under a subsequently published 
name. A. diversifolius A. Gray, founded on H. ortJiocarpus, appeared 29 years 
earlier than A. convallarius Greene (1893), which was proposed to replace 
A. campestris (Nutt.) A. Gray (1864) non Linne (1753). Even if the com- 
bination A. diversifolius, often wrongly- applied, were discarded as a nomen 
confusum, A. ihapensis M. E. Jones, in all probability an exact synonym of 
H. orthocarpus and published in January of 1893, Avill take precedence over 
A. convallarius published in October of the same year. But the taxonomy and 
involved nomenclature of this group are still in need of critical revision. 

Astragalus Whitedii Piper fma. speirocarpoides Barneb}', forma nova 

(Platel7, fig. 31) 

Astragalus Whitedii Piper fnia. speirocarpoides Barneby fina. nov. a prole tvpica (seu 
A. speirocarpo var. falciformi A. Gray) legumine in annulum saltem conipletuni vel 
saepius sesquialterum contorto, ncc falcate tantum, diversa. 

Nevada: in deep sandy soil among sagebrush, 10 miles north of Winnemucca, Hum- 
boldt Co., alt. 4500 ft., 23 May, fl. and fr. Riphi/ ai\d Barnehy 45.',2. Type in Herb. 
Calif. Acad. Sci. No. 300414. Also at Austin, Lander Co., Jones, 16 June 1882 (PO, 
annotated as A. Gibbsii var.). 

With its fruit twisted into one or one and a half spiral coils, the proposed 
forma is very similar in aspect to the true A. speirocarjms A. Gray of Wash- 
ington and northern Oregon, but the long calyx (tube 8 mm.), rather coarse, 
procumbent stems and broad leaflets are all characteristic of A. Whitedii, 
in the typical form of which the pods are not more than strongly falcate. It 
should be noticed that the fma. speirocarpoides invalidates Rydberg's ke.y to 
Homalohus Ser. Collini (N. Amer. Fl. 24: 258. 1929), from which this plant 
would appear to belong to A. speirocarpus. 



Vol. XXV] BABNEBY: PUGILLUS ASTRAGALOEVM ALTEE 149 

Astragalus zionis M. E. Jones 

Astragalus zionis M. E. Jones. Xylophacos zionis (Jones) Eydb. 

Arizona: sandy declivities under the Echo Cliffs near Limestone Tanks, about 13 
miles south of Navajo Bridge, alt. 5200 ft., 7 June, fr. Eipley and Barneby 4S68. 
Also observed, but not collected, under the same cliffs 11 miles north of Cedar Kidge. 

An addition to the flora of Arizona. The specimens are slightly abnormal 
in having the peduncles much shorter than is usual in material from Zion 
and the Virgin Valley, while the whole plant is more copiously pubescent. 

Astragalus panguicensis M. E. Jones 

Astragalus panguicensis (M. E. Jones), M. E. Jones, Contrib. W. Bot. 10:62, tab. 4. 1902. 
A Chamaeleuce var. panguicensis M. E. Jones, Proc. Calif. Acad. Sci. (II), 5:671. 1895. 
A. argophyllus var. panguicensis M. E. Jones, Contrib. W. Bot. 8:5. 1898; Eev. Astrag. 
207, tab. 47. 1923. Xylophacos vespertinus Eydb., N. Amer. Fl. 24:299. 1929, pro parte, 
quoad syn. cit., non A. vespertinus Sheld. 

Utah: gravelly benches in yellow pine forest west of Alton, Kane Co., alt. 6800 ft., 
5 June, fl. and fr. Ripley and Barnehy 4822. 

It is not without sqme misgiving that the collection is referred to this 
obscure species, for the type, Jones 6023 f, and only specimen of A. Chamae- 
leuce var. panguicensis at Pomona is a very mature and imperfect plant with- 
out either flower or fruit. In an envelope attached to the sheet there is a single 
pod marked in Jones' hand "probably the type of A. 'panguicensis," but this 
creates a further difficulty, for the pod does not agree as closely with the 
figures or description published by Jones as do the fruits of our specimens, 
and it may never have belonged there. The plant itself differs from ours in 
being evidently caulescent and in having, for the most part, broader and more 
obtuse leaflets. Manj^ species of this group, however, even if quite stemless at 
first, tend to become caulescent late in the season, and we have one specimen 
which in shape of leaflets closely matches the type. In both collections the 
stipules and the characteristically dense, silvery, subappressed pubescence 
are identical, and it should be remembered that the type locality, Panguitch 
Lake, is in the same massif as our station and not more than twenty miles 
distant. Certainly our plant can be referred to no other described species. 

If our determination is accepted as correct it becomes evident that A. pan- 
guicensis has been universally misunderstood. Jones first described it as a 
variety of A. Chamaeleuce A. Gray, three years later transferred his variety 
to A. argophyllus Nutt., then raised it to specific rank, and finally, in the Re- 
vision of Astragalus, returned it as a variety to A. argophyllus. Rydberg, 
evidently without first-hand knowledge of the plant, reduced it to the very 
different A. vespertinus Sheld. Actually the small, pale purple or ochro- 
leucous flowers and the quality of the pubescence suggest an affinity with 
A. castaneiformis S. Wats., from which it is immediately distinguished by 
the curved, lanceolate and much longer pod. 



150 



CALIFOBKIA ACADEMY OF SCIEXCES [Proc. 4th ser. 



Astragalus uncialis Barneby 

Nevada: Currant (Callaway), Nye Co., 12 May, flor. Bipley and Barneby 4421, topo- 
type. Also 20 miles to the west, in the foothills of the Pancake Eange near Lockes. 
No. 4423. 

A. uncialh was described from fruiting material and the fresh flower was 
unknown. As surmised by the writer, the corolla is of a brilliant purple, simi- 
lar to that found in A. utahensis (Torr.) T. & G. though perhaps of a some- 
what more violet tinge. 

Astragalus marianus (Rydb.) Barneby comb. nov. 

Xylophacos marianus Eydb., Bull. Torr. CI. 52:233. 1925; N. Amer. Fl. 24:301. 1929. A. 
Purshii X Newberryi M. E. Jones, Eev. Astrag. 216. 1923. 

Nevada: frequent on dry benches under pines and aspens along Snake Creek, near 
Treasure Lake, Snake Range, White Pine Co., alt. ca. 8200 ft., July, fr. Ripley and\ 
Barneby 4036. 

This rather well-marked species has been reported only from the mountains 

of southern and central Utah, where it is of not uncommon occurrence inj 

sagebrush valleys between six and eight thousand feet. The plants from! 

Nevada are not identical with the type, but have the broader leaflets of I 

Fremont 406 (NY) from Utah Lake referred to X. marianus by Rydbergl 

{I. c, 1925), and without doubt represent a shade form of the species. Another] 

gathering from the state, our no. 3515 from talus slopes beneath limestone! 

cliffs in the canyon of the Muddy River near Caliente, Lincoln County, atj 

4500 feet may also belong here, but the determination is not certain. The] 

specimens resemble A. marianus closely in habit, in the slender, often well-j 

developed caudices and numerous, small, appressed-sericeous leaflets, but inj 

the dense floccose pubescence of the pods they recall A. Newherryi A. GrayJ 

or even A. Blyae (Rose) Tidestr. The locality is well below the normal alti- 

tudinal limit of A. marianus and it may be that the plants had seeded down! 

from a higher level of the Highland Range into a soil where lack of humidityj 

and greater temperature might account for the increased density of vesture.! 

Further investigation, particularly of the flowers, may reveal an undescribedj 

species. 

Astragalus sabinarum (Rydb.) Barneby comb. nov. 

Batidophaca sabinarum Eydb., N. Amer. Fl. 24:320. 1929. 

Only the type specimen from Iron County, Utah {Garrett B2660, NY), hasi 
been seen by the writer, but this clearlj^ represents a good species. The com-j 
bination in Astragahts is proposed in order to facilitate comparison with its] 
nearest ally, A. niusimnnitm nob., described below. 

Astragalus musimonum Barneby sp. nov. 
(Plate, 17, figs. 1-9) 

Astrar/ahis mv.'timninim Barnobv sp. nov. ex affinitate A. sabinarnm (Eydb.) nob. a quo! 
pendunculis validis elongatis folia multo supcrantibus, racemis pluritloris, dentibusj 
calycinis abbreviatis hand nigro-eiliatis, et praesertim legumine parcius strigoso ad 



VOL.XXV] BABNEBY: PUGILLUS ASIEAGALOEVM ALTER 15] 

apicem abrupte hamato-recurvo facillime diagnoscenda. Habitu A. amphioxyn A. Gray 
affinesque nonniillas species aliquantulum refert, sed calyce campanulato cariiiaque lata 
ab Argophyllis (sive Xylophacis Eydb.) omnibus graviter diserepat. 

Herba perennis depressa caespitosa breviter caulescens, pilis argenteis adscendentibus 
basifixis imprimis arete appressis paucisque patulis commixtis undique strigoso-cana: 
caulibus compluribus prostratis saepe flexuosis incanis e coUo radieis lignosae verticalis 
emissis, 2-7 cm. longis, fere ex ipsa basi racemigeris : stipulis priminn iml)ricatis demum in- 
ternodio 3-9 mm. longo separatis, laneeolatis vel ovato-lanceolatis acutis, 3-5 mm. longis, 
praeter nervum medianum viridem omnino hyalinis vel superne herbaceis, mox scariosis, 
inferne glabratis intus glabris longe ciliatis ad basin petiolo breviter adnatis : f oliis humi 
appressis 4-10 cm. longis, petiolo gracili elongato leviter canaliculato rachin saepissime 
superanti: foliolis 5-9-jugis parvis remotiusculis inferioribus plerumque alternis articulatis 
obovatis ellipticis oblanceolatisve, apice rotundatis acutis vel apiculatis 3-7 mm. longis, 
2-3 mm. latis, secus racliin sursum decrescentibus : pedunculis numerosis, praecocioribus 
subradicalibus, prostratis validis striatis parcius strigosis viridulis, iam ad anthesin folia 
superantibus fructiferis elongatis: racemis brevibus 7-12-floris primum subcapitatis ca. 
1.5 cm. longis, mox laxioribus, demum 4-6 cm. longis : bracteis ovato-acuminatis ea. 3 mm. 
longis hyalinis ciliatis: pedicellis firmulis, vix 1 mm. longis, adscendentibus: calycis pur- 
pureo-tincti extus strigoso-cani tubo campanulato postice valde gibbo 3.5 mm. longo, 2 mm. 
lato, legumine accreseenti nunquam rupto, dentibus subulatis acutis 1.5 mm. longis: petalis 
laete violaeeo-purpureis, vexillo macula magna elliptica alba superne pulchre pluristriata 
instructo: vexillo 12-14 mm. longo, supra unguiculum brevem et angustum in limbum 
plicatum late oblongum 4 mm. latum obtusum apice emarginatum abrupte expanso, infra 
medium plus minusve retro-arcuato, marginibus reflexis: alls ca. 10 mm. longis, carinam 
pauUulum superantibus, lamina anguste lanceolato-kmata ca. 1.5 mm. lata, auriculo parvo 
reflexo ineluso 7 mm. longa : petalorum carinalium lamina oblique ovato-lunata 6 mm. longa, 
saltern 2.5 mm. lata, marginibus superioribus leviter concavis, inferioribus connatis supra 
medium in apicem porrectum plerumque obtusissimum per angulum fere rectum sursum 
arcuatis: legumine patulo sessili uniloculari, ambitu anguste elliptico, falcato, 1.5-2.0 cm. 
longo, utrinque angustato atque compresso, medium versus admodum obcomresso vel saltern 
dorsaliter sulcato, in rostrum abrupte hamato-reeurvum stylo persistent! cuspidatum sur- 
sum attenuato, sutura ventrali per longitudinem totam acuta sensim concava, dorsali inferne 
subrecta, ad medium prominula sed ob leguminis obcompressionem ibi depressa, ad rostrum 
acutissima abrupte convexa, sectione transversali anguste obcordata 5-6 mm. lata, ca. 2 
mm. alta, valvulis coriaceis sparsiuscule albostrigosis transverse striatis immaculatis: 
seminibus maturis ignotis. 

Nevada: calcareous gravel slopes of the Sheep Mts., near the entrance to Deadman 
Canyon, Clark Co., alt 5700 ft., 5 May, tl. & fr. Ripley and Barnehy 3332. Type in_ 
Herb. Calif. Acad. Sci. No. 300411. 

A. musimonum, with its cespitose, very shortly caulescent, prostrate stems, 
silvery-strigose herbage and racemes of purple flowers, has at first sight the 
aspect of a slender form of A. amphioxys A. Gray, but the companulate calyx- 
tube, broad keel and thin-walled pod indicate a relationship with that group 
of Astragali segregated by Kydberg as the genus Batidophaca, and among 
those species it finds a natural place beside A. sahinariini in the series Villosae 
(cf. Rj'db., I. c). From the latter it differs in the short calyx-lobes, more 
numerous flowers and more copious pubescence of straight, appressed hairs, 
while the mature fruits, laxly racemose on the stout, elongate and at length 
reclining peduncles, are comparatively glabrate, attenuate at base and 




152 CALIFORXIA ylCADEMY OF SCIENCES [Proc. 4th ser. 

abruptly hooked at apex, in contrast with the canescent, regularly lunate pod 
of A. sahinarum. 

A. musimomim has been known in the type locality on the western slope 
of the Sheep ]\Iountains for several years, but adequate fruiting material has 
only recently been obtained. In this station it is local but abundant on cal- 
careous benches in the Upper Sonoran Life-zone and seems to belong to the 
calciphile foothill flora of which, in southern Nevada, Actinea Cooperi (A. 
Gray), 0. Kze., Senecio stygius Greene and Astragalus Tidest^^omii (Rydb.) 
Clokey are characteristic elements. Immature specimens from northern Ari- 
zona, found growing under similar conditions at 5300 feet in Mokiak Pass, 
Mohave County (No. 4321), are almost certainly to be referred to A. musi- 
momim, but in absence of the legume the determination remains in doubt. 

The proposed trivial, from the Latin 'musimon,' alludes to that magnificent 
race of the Mountain Sheep, now happily preserved from extinction, which 
is native to the Sheep ]\Iountains, and from which the name of the range itself 
is certainly derived. 

Astragfalus pinonis M. E. Jones 

Pisophaca pinonis (M. E. Jones) Eydb. 

Nevada: Highland Eange west of Caliente, Lincoln Co., alt. 5200 ft., 20 May, fl. & fr. 
Ripley and Barnrhy 34S7a. Currant Creek, N. E. Nye Co., alt. 6200 ft. No. 3640. 

This rare species has been known hitherto only from Beaver (Frisco, Jones, 
the type collection) and eastern Juab (Tintic, Jones) Counties, Utah. Our 
specimens are taller and better developed than Jones' evidently depauperate 
type, with pods up to 2.5 cm. in length, but they are essentially alike in struc- 
ture and pubescence. The plants were found only as isolated individuals and 
appeared very scarce. 

Astragalus oophorus S. "Wats. 

Nevada: Head of Lee Canyon, Charleston Mts., Clark Co., alt. 9100 ft., Ripley and 
Barneby 4979. 

Through the kindness of Mr. Ira W. Clokey it has been possible to compare 
.this collection with Train 2141 from the same locality which was cited in a 
recent paper on the Astragali of the Charlestons (Clokey, Madrono 6: 217. 
1942) as A. artipes A. Gray. Study of the more ample material now available 
makes it clear that the Charleston plant is more justly referred to A. oophorus, 
but the original determination, which was made by the Bureau of Plant In- 
dustry and based on meager and somewhat atypical specimens, is readily 
understandable when it is realized how variable and how closely allied the 
species are, and how incorrectly they have been interpreted in the past. 

In fact there seems to be little ground on which A. oophorus and A. artipes 
can be separated specifically, for, while the typical phases occupy distinct, 
contiguous areas, there occur, particularly toward the southern limit of A. 
oophorus, specimens which are to some degree intermediate. A. artipes of 



Vol. XXV] BAENEBY: PUGILLUS ASIBAGALORUM ALTER 153 

northern Arizona, middle Utah and adjacent Colorado is distinguished by 
the cream-colored flowers and by the oblique legume with nearly straight 
ventral and strongly convex dorsal sutures. A. uophorus, on the other hand, 
a common species throughout central Nevada which extends into transmon- 
tane California along Owens Valley to the Panamints, has flowers with pur- 
ple banner and keel, and a regularly ellipsoid pod with either suture about 
equally convex. In southern Nevada and in the Panamints, however, purple- 
flowered plants are sometimes found with somewhat oblique legumes, in which 
case the only absolutely reliable criterion is the color of the flowers. This raises 
a serious difficulty, since Watson, in the original publication, described the 
flowers of A. oophorus as ochroleucous. This I believe to have been an error. 
Not only is the pigment in the petals of A. oophorus extremely fugacious in 
the herbarium, but among the numerous collections of the species made in 
Nevada in recent times not one with ochroleucous flowers has been detected. 
I have myself collected the species to the north, south and east of the type- 
locality (at Austin, in the Toyabe Range and in Montgomery Pass), yet 
among thousands of individuals none was observed without purple keel and 
banner. Apart from the value of the flower color as a taxonomic criterion,* 
the matter is of special importance in so far as it has affected the interpreta- 
tion of A. oophorus by Jones and Rydberg. Evidently it was a factor in Jones' 
treatment of A. artipes from middle Utah as A. oophorus (and simultaneously 
of the purple-flowered A. oophorus as A. artipes), while it also prompted Ryd- 
berg to recognize in A. oophorus two species of Phaca, P. oophora and P. 
jucunda, separated mainly by this very character. 

For many years Jones seems to have considered these two names synony- 
mous, and in the Index to the Revision of Astragalus he actually stated that 
"A. artipes ... is oophorus." Nevertheless his real intentions at this time 
seem to have been otherwise, for on Plate 22 of the same work appears a figure 
under the name "var. artipes" and the combination is repeated in the Index 
to Plates. By a curious misprint, however, it was not validly published. On 
page 121 of the Revision, immediately following a detailed description of 
A. oophorus (in which the pod described is clearly that of A. artipes) are 
printed two short paragraphs, both under the heading A. oophorus var. 
caulescens. There seems to be no doubt that the first line of the upper para- 
graph should have read "A. oophorus var. artipes (Gray, Proc. Amer. Acad. 
13. 370" which would join on comprehensively with "(1878) as species)" of 
the second line. But Jones' confusion went deeper than a mere typographical 
lapsus, for the plant figured as var. artipes is plainly nothing more than 
A. oophorus in the strictest sense, while the true A. artipes of Gray is repre- 
sented by the A. oophorus var. caulescens of the second paragraph and of the 

* An exact parallel is found in the case of A. Bedcwithii T. and G. and its vicariant var. 
purpureus M. E. Jones (A. ariemisiarum M. E. Jones). The former, with ochroleucous 
floAvers, so common in western Utah, passes westward, almost exactly at the Nevada line, 
into a form with purple flowers. 



154 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

corresponding figure on Plate 22. This point has been verified by study of 
the typical material of A. megacai'pus. var. caulescens ]\I. E. Jones (PO). 
It should also be noted that the figures of what Jones regarded as typical 
A. oophorus on Plate 21 contain a mixture : true A. oopliorus is there rep- 
resented only by the upper pod and section, while the remainder (marked 
"a," from Mammoth, Utah) are quite characteristic of A. artipes and in no 
waj' separable from the figure of var. caulescens on Plate 22. A touch of fan- 
tasj^ is added to this labyrinth of confusion by the fact that the material of A. 
artipes gathered by Lemmon, Rusby and Tourney in the Flagstaff region of 
northern Arizona are referred by Jones (1. c. 120) to A. megacarpus var. 
Parryi A. Gray, although he noted that they were abnormal in having ochro- 
leueous flowers. 

Phaca jucunda Jeps. and Rydb. has since publication been rightly reduced 
by Dr. Jepson himself (Fl. Calif. 2: 348. 1936) to A. oophorus. Apart from 
the criterion of color, which I believe to be chimerical, Rydberg advanced 
only the shape of leaflets to distinguish the segregate P. jucunda from P. 
oophora. A mere glance at any considerable suite of specimens, however, 
reveals that the shape of leaflets is in no way correlated with flower-color, 
with shape of pods or with distribution, for they may vary throughout the 
range of the species from crowded, suborbicular and refuse through many 
intermediate stages to remote, narrowly elliptic and acute (as in the Charles- 
ton material) and exactly similar variation occurs in A. artipes. Considerable 
variation occurs also in the length of the stipe and in the size of the corolla. 

I am disposed, therefore, to regard the Avhole complex as a single species, 
of wlych A. artipes may be regarded as a well-marked variety, under the com- 
bination A. oophorus var. caidescens Jones. The pertinent literature is sum- 
marized in the following synonymy. 

Astragalus oophorus S. "Wats. var. typicus Barneby nom. nov. 

A. oophorus S. Wats., Bot. King 73. 1871. A. oophorus sensu M. E. Jones, Rev. Astrag. 
120, pro parte, PI. 21 (as to upper figures). A. oophorus var. caulescens M. E. Jones, 
op. cif. 121 lin. 32 (lapsu), non ibid. lin. 38. A. oophorus vnr. artipes M. E. Jones, op. 
cit., Index to Plates and PI. 23, nomen nudum, non A. artipes A. Gray. Phaca oophora 
(A. Gray) Rydb., N. Amer. Fl. 24:338. 1929. P. jucunda Jeps. and Rydb., in Rydb., 
op. cit. 339. Astragalus jucundus Jeps. and Rydb. ex Peck, Man. PI. Oreg. 446. 1940, 
nomen midum. 

Astragalus oophorus S. Wats. var. caulescens (M. E. Jones) M. E. Jones 

A. oophorus S. Wats. var. caulescens (M. E. Jones) M. E. Jones, Rev. Astrag. 121, lin. 38, 
PI. 22, 1923, non ibid., lin. 31. A. megacarpus var. caulescens M. E. Jones, Proo. Calif. 
Acad. Sci. II, 5:043. 1895. A. artipes A. Gray, Proc. Amer. Acad. 13:370. 1878. A. 
oophorus sensu M. E. Jones, Rev. Astrag. 120, pro ma.r. parte, PI. 21, fig. a, non S. Wats. 
riifira arlipes (A. Gray) Rydb., Bull. Toir. CI. 32:(i()4. 190G; N. Amer. Fl. I.e. 



VOL.XXV] BABNEBY: PUGILLUS ASTBAGALOEUM A LT E B 155 

Astragalus straturensis M. E. Jones 

Atelophragma straturense (M. E. Jones) Rydb. Hamosa atratiforniis Rydb. 

Nevada: dry slopes among junipers and pinyons, 12 miles east of Panaca, Lincoln 
Co., alt. 7050 ft., 9 May, flor. Bipley and Barneby 4386. Also ibid., 11 June, fruct. 
No. 4971. 

A new record for Nevada. The flowers are bright violet-blue when fresh, 
with a pale, striate diamond in the fold of the banner. 

This will almost certainly prove to be the same plant as that collected by 
Vernon Bailey {Death Valley Exped. No. 1975) "about 30 km. east of Panaca" 
in Utah and determined by Sheldon (ap. Gov., Contrib. U. S. Nat. Herb. 4 : 87. 
1893) as A. ohscurus S. Wats. The latter species is not known south of the 
valley of the Humboldt River and has ochroleucous, not "purplish," flowers. 

Astragalus eremiticus Sheld. var. typicus Barneby nom. no v. 

A. eremiticus Sheld., Minn. Bot. Stud. 1:161. 1896, sensu strict o. 

Utah: Gunlock, Washington Co., with ochroleucous flowers. BipJey and Barneby 4298. 
Arizona: Mokiak Pass, Mohave Co., with purple flowers. No. 4313. Nevada: Pah- 
ranagat Mts., west of Crystal Springs, Lincoln Co., flowers purjile. No. 4408. 

Astragalus eremiticus Sheld. var. spencianus M. E. Jones, 
Contrib. W. Bot. 10 : 60. 1902 
A. boiseanus A. Nels., Bot. Gaz. 53:223. 1912. A. arrectus var. Kelseyi M. E. Jones, Eev. 
Astrag. 161. 1923, non A. Kelseyi Rydb. Tium eremiticum (Sheld.) Rydb., N. Amer. Fl. 
24:390. 1929; Bull. Torr. CI. 57:400. 1930, pro parte. 

Nevada: 10 miles west of Elko, Elko Co., in sagebrush. Bipley and Barneby 4588. 

The type of this variety, though not specified by Jones, is evidently his 
collection from Spencemont ( ? Lander County), Nevada, in 1891 (PO), and 
is exactly' similar to authentic material of A. hoisea^ius from the neighborhood 
of Boise, Idaho. Later Jones referred it as a variety to A. arrectus A. Gray, but 
mistakenly identified it with A. Kelseyi Rydb. which is A. atropuhescens 
Coult. and Fish, and not certainly distinct from A. arrectus. Rydberg, in his 
review of Tium (Z. c, 1930), recognized the differences which exist between 
typical A. eremiticus and the var. spencianus, but was unwilling to separate 
them. Nonetheless the morphological criteria enumerated by him, when con- 
sidered in the light of factors of distribution and habitat, appear amply suf- 
ficient to support a variety. The var. typicus is endemic to the arid, subdesertic 
ranges about the common boundary-point of Nevada, Arizona and Utah, 
alwaj's in the drainage of the Colorado River or its immediate tributaries 
such as the Muddy River or the Virgin, where it occurs on dry, stony slopes 
in the juniper-pinyon association, sometimes in depauperate sagebrush. The 
var. spencianus, on the other hand, occupies the interior basin region of north- 
eastern Nevada and adjacent Idaho and Oregon, in the drainage of the 
Owyhee, Snake and Humboldt rivers, where it is a characteristic element of 
the flora of sagebrush plains and valleys. It is not known to extend south of 
White Pine County, Nevada, and no intergrades have been seen. 



156 CALIFOBNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

Astragalus Bryantii Barneby spec, no v. 
(Plate 17, figs. 10-18) 

Astra ffaliis Bryantii Barneby spec. nov. inter Palantia generis Tii Medic, sensi. Rydb. 
adnumeranda, habitu pube etc. A. mol-iacensi A. Gray forsan proxinie affinis, sed ab 
eo consociisque omnibus leguminis angnstioris valde compressi sutura ventrali acuta 
necnon pericarpio tenui diversissima et his notis Havinsae Tricarinatas Eydb., a quibus 
imprimis tubo calycino tubuloso nee campanulato longius distat, simulans. 

Herba verosimiliter pcrennis, 'tripedalis' teste cl. Bryant, pube e pilis brevibus adscenden- 
tibus candidis constituta praeter foliolorum paginam superiorem glabratam undique stri- 
gosa, partibus junioribus villosulo-eanescentibus : caulibus ut videtur adscendentibus (sed 
imis ignotis) plus minusve flexuosis striatis purpuraseentibus: stipulis deltoidco-acuminatis 
extus strigoais herbaceis vcl anguste scarioso-niarginatis ca. 5 mm. longis, mox reflexis: 
foliis patulis subsossilibus vel breviter petiolatis, 8-12 cm. longis, racM subtereti strigoso 
petiolum multoties superanti: foliolis 14-20, petiolulo 0.5-0.75 mm. longo gestis. ovatis, 
late ellipticis vel obovatis, 5-14 mm. longis, 3-8 mm. latis, inferne strigoso-eanescentibus: 
pedunculis firmis arcuato-adscendentibus striatis 4-5 cm. longis, in racemum laxum ca. 
15-florum fructiferum vix elongatum nee folio suffulcranti multo longiorem abeuntibus: 
bracteis lanceolato-attenuatis ca. 2 mm. longis: floribus violaceis 15-17 mm. longis pedicello 
arcuato 1.0-2.5 mm. longo saepissime atropiloso suffultis: ealycis membranacei pilis albis 
nigrisque commixtis strigosi tubo late cylindrico, 5 mm. longo, postice gibbo, ovario in- 
tumescenti mox rupto, dentibus anguste subulatis acutis 1.5-1.75 mm. longis coronato: 
vexillo obovato 16 mm. longo, ca. 8 mm. lato, ad medium retrorsus arcuato sed apice pro- 
funde emarginato declinato, marginibus reflexis: alls ca. 14 mm. longis, lamina lunata 
obtusa 1.5-1.75 mm. lata, auriculo basali conspicuo incluso 7 mm. longa : carinae petalis 
ca. 13 mm. longis laminis latius lunatis saltem 7 mm. longis, 2.5 mm. latis, marginibus 
superioribus profundiuscule concavis, inferioribus (connatis) inferne reetis ad medium 
abrupte in apicem latum obtusissimum sursum arcuatis: legumine horizontaliter patulo vel 
leviter adscendenti, arete sessili, 2-loculari, subreeto vel saepius paullum arcuato, ambitu 
lineari vel lineari-lanceolato, 2.0-2.5 (3) cm. longo, ad basin rotundato supra medium 
sensim attenuate, valde compresso, sulco 0.75-1.5 mm. alto impresso dorsaliter peicurso, 
sutura ventrali per longitudinem totam proniinula, dorsali ad ventralem usque introflexa 
septum subcompletum 2.0-2.5 mm. altum fere in leguminis apicem productum efformanti, 
valvulis chartaceis tenuiter reticulatis parce strigulosis vel demum glabratis nonnumquam 
minute purpureo-guttulatis, sectione transversali anguste deltoideo-obcordata, 3.0-4.5 mm. 
alta, 1.5-3.0 mm. lata: seminibus oblique reniformibus, 2 mm. latis, brunneis. M 

Arizona: at the head of Phantom Canyon in the Grand Canyon of the Colorado 
River, Coconino Co., 15 Dec. 1939. Collected by Dr. 11. C. Bryant of the National 
Park Service in whose honor it is named. 

Type in Herb. Calif. Acad. Sci. No. 293940. Also collected in sand at the mouth of 
Hermit Creek in the Grand Canyon, 10 April 1917. Eastwood 5991 (G, F). 

The type of A. Bryantii is apparently part of the collection referred to by 
Kearney and Peebles (Fl. PI. Ariz. 486. 1942) as representing an undescribed 
species allied to A. palans M. E. Jones, but, although a real affinity may exist 
between the two, it has been found convenient for purposes of diagnosis to 
compare it rather with A. luokiacotsis A. Gray, with which it agrees more 
closely in the quality of the pubescence, in the violet, not reddish purple 
coloration of the corolhi, and in the less marked curvature of the legume. From 
A. mokiacensis, known only from a few localities on either side of the Colo- 



VoL.XXV] BARNEBT: PUGILLUS ASTBAGALOEUM ALTER 157 

rado River immediately below the Grand Canyon, A. Bryaniii differs in the 
strongly compressed, more slender and sparingly strigose pods which are of 
thinner texture and spread horizontally from the axis of the raceme. Indeed 
the texture and compression of the pod, with its acute ventral suture, led me 
at first to regard the species as more intimatelj^ connected with forms of A. 
arrectus A. Gray {Tiuni ser. Arrecta Rydb.) or with Hamosa ser. Tricarinatae 
Rydb., but the complete absence of stipe and the cylindric calyx-tube make 
such an alignment improbable, and the species is more naturally associated 
with A. mokiacensis. 

As far as can be seen from the limited material at hand, A. Bryaniii is very 
constant in vegetative characters, but the shape of the pod is evidently 
variable. Normally, as always in Eastwood 5991, the pod is narrowly lance- 
attenuate in outline, the cross-section at the middle being about 3 mm. high. 
Attached to Dr. Brj^ant's specimen, however, there are several loose pods, 
essentially similar in structure and pubescence but half as broad again, the 
cross-section being 4.5 mm. high and the outline lanceolate, more abruptly 
acute rather than attenuate at apex. Although the plant from which these 
pods have fallen has not been studied, their measurements are included in 
the description of the species and they have been treated as an integral part 
of the type. A similar latitude of variation has been found by the writer to 
occur in a number of species in the genus. 

Astragalus Pattersonii A. Gray, sens, strict. 

Jonesiella Pattersonii Eydb. 

Arizona: red clay hills 2 miles east of Fredonia, Coconino Co., alt. 4900 ft. Ripley and 
Barneiy 4361. 

The specimens are almost exactly typical, with narrowly elliptic, subsessile, 
strictly erect pods. This form of the extremely polymorphic species was known 
to Rydberg only from Colorado, and has not been reliably reported outside 
of that state, though a closely allied form with spreading pods is common in 
the Navajo Basin. The A. Pattersonii of Tidestrom and Kittell (Fl. Ariz, and 
N. Mex. 214. 1941), if one may judge from the synonymy, is an aggregate 
containing a number of diverse forms. 

Astragalus ensiformis M. E. Jones var. typicus Barneby nom. nov. 

A. ensiformis M. E. Jones, Rev. Astrag. 226. 1923. A. ursinus M. E. Jones, Proc. Calif. 
Acad. Sci. II, 5:658. 1895, non A. Gray. Hamosa ensiformis Eydb. 

Arizona: gentle stony slopes among junipers in Mokiak Pass, north of Wolf Hole, 
Mohave Co., alt. 5300-5500 ft., 4 May, fl. and fr. Ripley and Barneby 4311. 

The type-locality of A. ensiformis, "4 miles above Pagumpa," is very close 
to our station and the species has been collected elsewhere only by Peebles and 
Fulton in northern Navajo County (cf. Kearney and Peebles, Fl. PI. Ariz. 
488. 1942). The name is universally cited as having been published in 1895, 
but this is incorrect, Jones, it is true, described the species quite minutely 



158 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

in that year (I. c), but under the name A. ursinus A. Gray, merely suggesting 
A. ensiformis as an alternative should the plant prove as distinct as he sus- 
pected, and the combination was not validly applied until the Revision of 
Astragalus in 1923. Kydberg, who did not see the flowers, associated A. ensi- 
formis with A. Congdonii S. Wats, and A. drepanololus A. Gray, whereas, 
with its tubular calyx and enlarged, scarious stipules, it clearly belongs to 
his series Malacae of Hamosa and is closely related to A. Minthorniae (Rydb.) 
Jeps. In fact a collection from southern Utah, described below as a new 
variety, forms a passage between the two. 

Astragalus ensiformis M. E. Jones var. gracilior P.arneby var. nov. 

(Plate 17, fig. 28) 
Astragalus ensiformis M. E. Jones var. gracilior Barneby var. nov. a var. typico nob. caule 
internodiis extensis duplo elatiori, floribus post anthesin baud deflexis, racemo fruetifero 
minime secundo et praesertim legumine graciliori minus compresso patule erecto eviaen- 
tius hirsiitulo noc appresse striguloso diversa. Ab aflfini A. Minthorniae (Rydb.) Jcps., 
cujus liabitum in memoriam nonnihil ledigit, imprimis legumine magis arcuato baud 
dense sericeo-villoso facile separanda. 

Utah: in stiff soil among sagebrusb, 5 miles south of Veyo, Washington Co., iilt. 
4900 ft. Ripley and Barneby 4951. Type in Herb. Calif. Acad. Sci. No. 300413. 

So different was the superficial appearance of this variety from that of 
A. ensiformis var. typicus that it was thought in the field to represent an 
undescribed species, but subsequent study has shown that the differences, 
though striking, are not fundamental, and it cannot be specifically separated. 
In the typical form the decumbent stem, with congested internodes, does not 
exceed 15 cm. in length, the fruiting raceme is strongly secund, the fiexuous 
pedicels are arcuate-reflexed and the sparingly strigose pods, though arched 
upwards, are essentially pendulous. The stems of the var. gracilior, on the 
other hand, are upright and slender, reaching a height of 3 dm., the raceme 
is not at all secund, while the more finely and densely pubescent pods are 
held erect or somewhat spreading on straight and rigid pedicels. There is 
no appreciable difference in the form of the flowers, though in the var. iypicus 
they are reflexed after fertilization. 

Astragalus chamaemeniscus Barneby 

Nevada: in sand under sagebrush near the Geyser Ranch, N. Lincoln Co., alt. 0050 ft., 
11 May, flor. Ripley and Barneby 4416. 

This species was known previously only from extreme northeast Nye and 
adjacent White Pine counties in the valley of the White River. 

Astragalus Arthu-Schottii A. Gray, sensu restricto 

A. ArtJiu-Seliottii A. Gray, Proc. Amer. Acad. G:209. 1863, pro max. parte, noii op. cit. 7: 
337. 1867. A. Coultcri sensu S. Wats., Bot. King, 66, 435. 1871; A. Gray, Bot. Calif. 
1:146. 1880; Sheld., Minn. Bot. Stud. 1:140. 1894 et auct. mult, pro parte, quoad syn., 
non Bth. A. lenfigiriosus var. borreganus M. E. Jones, Contrib. W. Bot. 8:3. 1898; 
Rev. Astrag. 126, tab. 25. 1923, A. lentiginosiis var. Coulteri sensu Jones, Rev. Astrag. 



Vol. XXV] BARNEBY: PUGILLUS ASTEAGALOBUM ALTER 159 

127. quoad syn. ; Jeps., F1. Calif. 2:357. 1936, quoad syn. et pi. cit. Schott. A. agninus 
Jeps., Mau. Calif. 577. 1925; Fl. Calif. 2:355. 1936; Harrison and Kearney, Journ. 
Wash. Acad. Sci. 22:227. 1932; Munz, Man. S. Calif. Bot. 270. 1935; Kearney and 
Peebles, Fl. PI. Ariz. 487. 1942. Cysiium Arthu-Schottii (A. Gray) Rydb., N. Amer. 
Fl. 24:407. 1923. C. agninum (Jeps.) Rydb., loc. cit. 408. 

Study of the somewhat elaborate synonymy and literature cited will show 
that 4- Arthu-Schottii is believed to be the oldest valid name for the plant 
which has been described and treated in comparatively recent years as A. 
lentiginosus var. horreganus Jones and A. agninus Jeps., and, while reasons 
might be advanced for discarding Dr. Gray's name as a permanent source of 
error and confusion, a reexamination of the facts tends to support its revival 
into current use. A. Arthu-Schottii was first encountered by Schott during 
the exj^loration of the Mexican boundary in 1855, but his collections were 
not cited by Gray in the Botany of the Survey, doubtless because the speci- 
mens available to him were without fruit. Several years later, having received 
from Fremont additional material of what he took to be the same. Gray de- 
cided to publish the species, including in his concept a plant collected in 
northern Sonora by Capt. Smith. Thus the original A. Arthu-Schottii was 
founded on three elements, cited by Gray (I. c.) in the following words: "on 
the ]\Iohave River, Fremont, 23 April. Diluvial banks of the Colorado near 
its mouth, 12 March, Schott. Boca Grande, Capt. E. K. Smith.", and each of 
which belonged to a different species. The Fremont specimen (NY) is a very 
depauperate, first-year individual of that form of A. Coulteri Bth. with small 
corollas which is common on the Mohave Desert of California and which is 
variously treated in modern literature as A. lentiginosus var. Coulteri or 
var. Frctnontii: the Schott element, as will be established below, belongs to 
A. agninus Jeps. : while the material from Boca Grande, a fragmentary speci- 
men with leaflets glabrous on the upper surface, though not susceptible of 
certain determination, is probably a form of A. lentiginosus var. yuccanus 
Jones (certainly not Cystium eremicum as annotated by Rydb.). In settling 
on the type of A. Arthu-Schottii the latter may be quickly discarded : not 
only was it cited last by Gray, but the only reference to it in the description 
seems to be "foliolis . . . pagina superiore interdum gUibrata," and it must 
have formed a quite unimportant part of the species that the author had in 
mind. The choice between the Fremont and Schott elements, however, is more 
difficult, for the description fits both with almost equal exactness. Neverthe- 
less the phrases dealing with the calyx — "calycis dentihus suhulatis tuba 
ohlongo-campanulato vix dimidio longiorihus" — and with the keel — "carina 
majuscula suhrecta vexillo paullo breviore" — are matched more closely by the 
corresponding organs of Schott's plants than by those of Fremont's and the 
description seems to have been prepared primarily from the former, the first 
collected and more copious specimens. For this reason, as well as to avoid 
the anomaly of excluding from the species named in honor of Schott the very 
elements which he himself collected, it seems proper to regard the Boundary- 
Survey Xo. 256 in herb. Gray as the type. 



160 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

As stated above, the material of A. Arihu-Schottii available to Gray was 
without frnit, and this is true both of the sheet now in herb. Gray as well as 
of tlie two in the Torrey herbarium (NY). In the Field Museum, however, 
there are three additional isotypic sheets, apparently from Schott's private 
herbarium, one of which has nearly mature legumes and belongs, quite incon- 
testably, to A. agninus Jeps. Even without this further evidence the identity 
of Schott's plant would be almost certain from locality alone. The six sheets 
seen by the writer are variously annotated "Colorado Desert near Fort Yuma," 
"Colorado Desert, Sonora" and "Between San Diego and the Colorado," and 
whether or not these represent one or more separate gatherings it is clear that 
all came either from extreme southern Imperial County, California, or from 
closely adjacent Sonora, a territorj^ from M^hich A. Coultcri is totally un- 
known yet w'here A. agninus has been frequently collected. Nevertheless, start- 
ing with Watson in 1871, the Schott collection has been regailarly determined 
by all students of the genus as A. Coiilteri and is so annotated by Sheldon 
(NY) and cited by Jones and Jepson. The error is perhaps traceable back 
to Gray's publication, as early as 1867, (/. c), of a revised description of 
A. Arthu-ScJiottii taken from fruiting material communicated by Cooper 
from the Mohave Desert ("Camp Cad^v and elsewhere," G.) W'hicli is con- 
specific with the early Fremont collection but not with that of Schott, i. e. 
A. Coulteri var. It should be borne in mind that, at the time when Gray pro- 
posed A. Arthu-Schottii, A. Coulteri was known to him only from Bentham's 
description, and it is not surprising therefore that, when A. Coulteri was 
finally identified with the Cooper plant in fruit, the whole oiA. Arthu-Schottii 
was also thought to belong there. 

In the North American Flora (/. c.) Rydberg abandoned the accepted inter- 
pretation of A. Arthu-Schottii, recognizing it as a species in the genus Cyn- 
tium, but the effect is somewhat marred by the fact that he recognized Cystium 
agninum and C. Coulteri as well. If one may judge from specimens in the New 
York Botanical Garden so annotated by Rydberg himself, his C. Arthu- 
Schottii contained, in addition to the original Schott collection (which he also 
regarded as typical), a number of plants from the Mohave Desert such as 
Parish 4957, Elmer 3641, etc., which are inseparable from the earlj' Fremont 
collection, although that actual specimen he referred to C. Coulteri (Bth.) 
Rydb. ! It should also be remarked that in the same work {I.e., in syn.) Rydberg 
incorrectly cited A. lentiginosus var. horreganus as having been published by 
Jones in the Revision of Astragalus, whereas it actually had appeared as 
early as 1898 (cf. supra) and in addition failed to note Jones' excellent figure. 

A. Arthu-Schottii is a plant of rather restricted distribution in the south- 
w^estern deserts from interior San Diego County, California, at Borrego Wells, 
Orcutt, type of var. horreganus (PO), Jepson 8883, type of A. agninus (NY, 
fragments), to Yuma, Arizona, Beckett & Nobles 8695 (G, F), southward 
just into Sonora about San Luis, Harrison & Kearney 8434 (F), and north 
along the Colorado River to eastern San Bernardino County, California, at 



VOL.XXV] BAENEBY: PUGILLUS ASTEAGALORUM ALTEE 161 

Silver Lake, J. T. Howell 3586 (CAS) and Nipton on the borders of Nevada, 
Ripley d- Barnehy 3360. The species is very closely allied to A. Coulteri, which 
it replaces south of the Coachella Valley, but differs in the annual root, lax 
fruiting racemes and scarcely inflated pod. 

Astragalus lentiginosus Dougl. var. csesariatus Barneby var. nov. 

(Plate 17, figs. 29-30) 

Astragalu.s lc7)figinosus Dougl. var. ccesariaUts Barneby var. nov. inter Cysfia neogea Eyd- 
bergiana sive Astragali Diphysos A. Gray adnumeranda, caulibus prostratis, floribus 
congestis et legumine coriaceo A. tehatchapiensi (Eydb.) Tidestr. proxime affinis, sed 
ab eo floribus maximis ca. 19 mm. longis saturate violaceis, dentibus calycinis longio- 
ribus acutis, legumine aliruptius acuminato necnon racemorum axi calycibus bracteisque 
pube e pilis nigris laxius appressis constituta crebre indutis, satis superque diversa. A. 
lentiginosus var. nigricalycis M. E. Jones, qui etiam in iisdem finibus passim oceurrit, 
caulibus paucis elatis, raceme laxiori, floribus minoribus saepissime ochroleucis, pube 
crispa implexa et legumine omnino albo-villosulo nee partim atro-strigoso a nostra dis- 
tantius recedit. 

California : grassy slopes of the Temblor Range above McKittrick, Kern Co., alt. 2900 
ft., 13 April, fl. and fr. Eipley and Barneby 3243. Type in Herb. Calif. Acad. Sci. 
No. 289572. 

The proposed variety is here referred to A. lentigiiwsus — sensu latissimo — 
in order to fit into Dr. Jepson's treatment of this complex group in the Flora 
of California, but the disposition is probably not final. The writer has spent 
many months in study of A. lentiginosus and its allies and it is now believed 
that a taxonomic arrangement based on a specific concept intermediate be- 
tween the extremes of Rydberg, who recognized 33 species in Cystium, and 
of Jones, who regarded all the known races as varieties of A. lentiginosus, 
is not impossible. In spite of the fact that the specific limits would be some- 
what blurred, it would appear feasible to recognize at least three major 
species, A. lentiginosus, A. Coulteri Bth. and A. diphysus A. Gray (approxi- 
mately corresponding with Rydberg's three series Lentiginosa, Coulteriana 
and Diphysa of Cystium), under which the remainder, with several as yet 
undescribed, might reasonably be classified as varieties. In this case the var. 
ccesariatus would be referred to A. diphysus and associated closely with 
A. tehatchapiensis (Rydb.) Tidestr. (Proc. Biol. Soc. Wash. 50:21. 1937, as 
tehachapiensis) yet easily separable by the larger, brilliantly colored corolla, 
acute calyx-lobes, sinuous stems, and by the pubescence of loosely appressed 
black hairs which invest the w^hole axis of the raceme, the pedicels and calyces, 
and which recur, mixed with white villi of similar structure, on the valves of 
the more abruptly cuspidate pod. A. lentiginosus var. nigricalycisM. E. Jones, 
the common race of the San Joaquin Vallej^ differs from ours in its taller 
stems, more lax racemes of smaller ochroleucous flowers, and by the tangled, 
curly pubescence of the stem, herbage and legume, the valves of which are 
larger and of thinner texture. 



162 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th see. 

A. lentiginosus var. ccesariatus is not uncommon on the eastern slope of the 
Temblor Range, where it forms large clumps among the ephemeral spring 
grasses, man}' annuals characteristic of the flora of the Great Valley foothills, 
and such perennial herbs as Senecio Breweri Davy and Delphiniiim hesperium 
var. recurvatum Jeps. The stems, arising from a heavy root and multicipital 
caudex, are very numerous, reclinate below but with ascending tips, and the 
subcapitate or short and dense racemes of large purple flowers with their 
contrasting striate eye on the banner make the variety one of the showiest of 
California Astragali. The specific epithet is derived from the Latin 'ccesaries' 
and refers to the abundance of dark hairs in the inflorescence. 

Astragalus striatiflorus M. E. Jones, Proc. Calif. Acad. 
Sei.II, 5:643. 1895 

(Plate 17, figs. 19-24) 

Utah: locally abundant in white sand and on rock-ledges beneath sandstone cliff>s, 10 
miles north of Johnson Ranch, east of Kanab, Kane Co., alt. 5600 ft., 6 May, flor. 
Eipley and Barnehy 4358. Ibid., 5 June, fl. and fr. No. 4815. Also beneath the white 
cliffs 5 miles south of Zion Park Junction, Kane Co., alt. 6100 ft. No. 4365. 

This exceedingly obscure species lias been known up to the present only 
from an inadequate fragment, devoid of pods, which was gathered by Jones 
*'in red sand, above Springdale" (the western entrance to Zion Park, Wash- 
ington County, Utah) in September, 1894. The type collection, Jones 6080k 
(PO) , consists of a single very mature plant from which even the flowers have 
now fallen. The characteristic habit, however, the pubescence and curiously 
connate stipules still visible in the dried material, when taken in conjunction 
with the description of the flower and its attenuate keel, leave little doubt that 
our recent collections are determined correctly. Jones was apologetic about de- 
scribing a species of Astragalus wdthout knowledge of the fruit, but, con- 
vinced by the floral and other peculiarities of his material that he had before 
him a species "certainly belonging to the hiflati and probably near to A. ser- 
pens" (Z. c. ) , he was unable to resist giving it a name. That his proposition was 
valid is now obvious, but in referring A. striatiflorus to the Inflati (approxi- 
matel}^ the genus Phaca as defined by Rydberg) Jones was far from the mark : 
actually the pod, while inflated, is bilocular and the affinity of the plant is 
probably with A. platytropis A. Gray. In the text of the Revision of Astraga- 
lus the species is not mentioned, but in the Index (which is not paginated) it 
is reduced without comment to A. Silerauus M. E. Jones, a true member of 
the Inflati with discrete stipules, quite different flowers and elongated stems. 
Rydberg, to whom the species was entirely unknown, cited A. striatiflorus in 
his list of omitted synonyms in Astragalus (N. Amer. Fl. 24 :455. 1929), and 
could do little else but follow Jones' earlier reduction. As the species remains 
virtually undescribed to this day, it has been thought advisable to prepare 
a revised description with emphasis on the floral and carpological characters. 



VOL.XXV] BARNEBY: PUGILLUS ASTEAGALOEUM ALTER 163 

Astragalus striatiflorus M. E. Jones, char, fruct. 

A low cespitose perennial herb with a fleshy taproot and loosely branched subterranean 
caudex, the stems and herbage densely and softly villous-caneseent with white spreading 
and somewhat curly hairs. Stems very short, decumbent, clothed below with loosely imbri- 
cated stipules, the upper 2 or 3 internodes about 1 cm. long. Stipules very broadly obovate, 
truncate and often mucronate, 2-3 mm. long, scarious and many-nerved (the nerves more 
or less anastomosing), adnate neither to the stem nor to the petiole but connate around 
both to form a cup, strigose-villous without. Leaves 2-3 cm. long, the slender rachis and 
petiole subequal. Leaflets 11-13, petiolulate, obovate to oblanceolate, obtuse, truncate and 
mucronulate or rarely a little emarginate, often conduplicate, contiguous, the terminal 
smallest. Racemes short, loosely 3-5-flowered, on peduncles about equalling the leaves. 
Bracts ovate, acute or acuminate, scarious, longer or shorter than the spreading pedicels. 
Calyx purplish, villous-hirsute with white or some black hairs, the tube campanulate or 
obscurely obconic, 3-4 mm. long, oblique at the orifice, the acute teeth 2-3 mm. long, hetero- 
morphous, the ventral pair deltoid-acuminate, the lateral pair and the dorsal one linear- 
subulate. Corolla 9-12 mm. long, ochroleucous or suffused with lavender, the banner striate 
with purple lines and the keel deeply purple-tipped. Banner with suborbicular blade 8-9 mm. 
broad, narrowed into a very short claw, arcuate much below the middle nearly to a right- 
angle, the margins spreading but not reflexed. Wings 8-9 mm. long, the blades hmate 
obtuse, about 2 mm. broad, including the large basal auricle 6 mm. long. Keel-petals equal- 
ing the wings, the blades 6-7 mm. long, obliquely lanceolate and arcuate upwards, strongly 
gibbous above the claw and there attached to the wings, the upper margins concave at the 
middle, the lower margins straight below, toward the middle abruptly convex and arched 
upward to the slightly declined apex of the long-attenuate but obtuse beak, free for 1 
mm. below the summit. Style filiform, at full anthesis exserted about 1 mm. from the keel, 
geniculate below the capitate stigma. Pod bladdery-inflated, bilocular, 12-1.5 mm. long, 
spreading, sessile but seated on a minute boss within the calyx and readily deciduous there- 
from, in outline broadly ellipsoid, rounded and somewhat oblique at base, abruptly acute 
and apiculate at the apex which is crowned by the coiled persistent style, shallowly sulcate 
along either suture, the ventral suture nearly straight, filiform, the dorsal introflexed to 
form a thin double partition 3 mm. high produced across the entire width of the cavity, the 
valves membraneous, densely villous with white curled spreading hairs, mottled with purple, 
the cross-section suborbicular 6-7 mm. broad and a little less high, emarginate at the sulcate 
sutures. Seeds about 8 in either locule, obliquely reniform, compressed, alwut 2 mm. long, 
olive-green. 

A. striatiflorus is set off from all known species of Astragalus by a combina- 
tion of characters which excludes it from a place in any natural group hitherto 
defined. An attempt to determine the species from Kydberg's monograph 
leads the enquirer directly to Cystium platy trope (A. Gray) Rydb., or, more 
deviously, to A. anisus M. E. Jones, but the similarity between these plants 
is, I believe, rather superficial and based entirely on the structure of the pod. 
In the present paper it is not proposed to discuss the taxonomic position of 
A. anisus, which differs {inter alia) from A. platytropis and A. striatiflorus 
in the elongate flower and cylindric calyx-tube and which is probably cor- 
rectly placed by Jones and Rydberg as an aberrant relative of tlie Mollissimi 
(A. mollissimus Torr. et spp. affin.). But the differences between the two last 
and the relationship of each to A. lentiginosus sens, latiss., i. e. Cystium of 
Rydberg', will be examined in more detail. 

It has already been implied that a similarity between the pods of two given 



164 CALIFOBXIA ACADEMY OF SCIEXCES [Pkoc. 4th ser. 

species of Astragalus is not necessarily a criterion of affinity, but this assertion 
must be amplified, for, thus baldly stated, it stands in direct contradiction to 
Rydberg's most important principle of classification. All of Rydberg's later 
work on the genus, which culminated in his monographic revision for the 
North American Flora, is based on the primary assumption that carpological 
characters, even when of the most minute and quantitative nature, are of 
supreme value in the definition of groups or, in his terminology, of genera. 
Diversity in structure of the flower, stipules or pubescence he discarded as 
of quite secondary importance. By logical application of this principle, Ryd- 
berg was led to assume an affinity between species, scattered perhaps from the 
Levant to the Andes, which happened to have a similar width of septum in 
the pod, yet simultaneously to refer to widely separated groups two species 
intimately connected by ties of morphology and geography in which the sep- 
tum was found to be strongly or weakly developed. Of course this method was 
satisfactory in parts, for many allied species do, as one would expect, bear 
similar pods, but in some cases it produced polyphyletic groups of great arti- 
ficiality. Had Rydberg himself admitted this, it could be easily forgiven. But 
interior evidence from his published writings leaves little doubt that he pro- 
posed his genera as genuine phylae, as evolved expressions of a single heredity 
marked by a carpological structure common to all included members. And 
implicit in that belief was the conclusion that any two members of one genus 
are more closely related to each other than either can be to a member of another 
genus. With this principle in mind let us take an example from Rydberg's 
own work. 

Tiv))i Wilsonii (Greene) Rydb. is a species of Astragalus endemic to the 
high plateaux of northern Arizona, described by Greene as related to A. diphy- 
sus A. Gray (Cystium diphysum Rydb.) and reduced by Jones in synonymy 
to a variety of A. lentiginosus Dougl. The pod is lanceolate, leathery, slightly 
but not conspicuously inflated, with the valves introverted dorsally to form 
a partition and thus partially bilocular. The septum does not quite reach the 
ventral suture, but the gap is not above one millimeter in width. With this 
we may compare Cystium agninum (Jeps.) Rydb., discussed elsewhere in this 
paper. Here we have a pod of similar outline, little inflated, but the septum 
traverses the whole width of the cavity (sometimes less : cf . Jeps. ex char. : 
". . . extending half way or nearly to the ventral suture") and the gap between 
the sutures is as a rule closed. Rydberg rightly disregarded the lack of infla- 
tion in the fruit of C. agninum, otlierwise anomalous in his genus Cystium, 
yet attached sufficient importance to the equally quantitative character of 
the septum to refer A. Wilsonii to Tinm, and we are led to believe that despite 
their manifest similarity in all other respects to A. coulteri and A. diphysns, 
both Rydbergian Cystia, these two species of the southwestern states are more 
closely related to, respectively, the old-world A. Cicer L., genotype of Cystium 
and A. sulcatus L., genotype of Tium, than tliey are to one another. 

This relationship is not. of course, ipso facto, impossible. But once tlie rela- 



Vol. XXV] BAENEBY: PUGILLUS ASTRAGALOEUM ALTEE 



165 



tionship is admitted we must also accept that the evolution of the septum 
stopped abruptly at an incalculably distant date and has since been station- 
ary, while the differentiation of species has continued along other lines. 
However, when the marvelously diverse carpology of Astragalus is consid- 
ered as a whole, and when one recollects the manner in which the pods of 
species accepted by Rydberg himself as closely allied may differ in size and 
shape, in length of stipe, in the degree of compression and inflation, and even 
(cf. Tium platycarjnim and T. racemosum) in the very width of septum, it 
becomes clear that all parts of the fruit have been equally subject to evolu- 
tionary progress and recession in comparatively recent times. Indeed there 
seems to be no reason why the similarity of structure between the pods of Old 
and New World species, remarked by Rydberg and other taxonomists before 
him, cannot be explained as well by parallel development as by a common 
heredity. In this case no single carpological character, nor even general agree- 
ment in structure of the fruit, can be accepted as a reliable basis of a natural 
classification throughout the genus unless supported by other criteria of 
proved value : and the names of generic segregates such as Cysiium and Tium, 
founded on European or Asiatic species, have a very questionable place, 
whether applied to genera, subgenera or sections, in the taxonomy of Ameri- 
can Astragalus. 

From this statement of generalities we may well return to the particular 
subject of the present note, A. striatiflorus and its immediate allies. In the 
interests of clarity the major characters of the species are contrasted in the 
following table with A. platytropis and A. lentiginosus. 

A. lentiginosus 
strongly caulescent 
strictly appressed 
free or obscurely 

adnate to petiole 
lunate obtuse 

shorter than wings 
included 

with septum formed 
as in A. striatiflo- 
rus, the ventral 
suture sometimes 
(especially in allied 
species) very nar- 
rowly introflexed 
but never below in- 
sertion of funicles 

It will be seen at once that A! striatiflorus differs from its supposed allies 
(or, more precisely, from other species with bilocular, inflated pods) in the 
attenuate keel, exserted style, stipules and pubescence, and additionally from 
A. platytropis, of which it has nearly the habit, in the structure of the septum. 
A. platytropis (plate 26, figs. 25-28), on the other hand, is unique in the 
concretion of the stipules, in the septum and broadly truncate keel. In the 





A striatiflorus 


A. platytropis 


Stems 


shortly caulescent 


subacaulescent 


Pubescence 


villous 


loosely strigose 


Stipules 


connate around stem 


connate abaxially 




and petiole 


about the petiole 


Keel 


lance-attenuate 


oblong-truncate 




equalling wings 


equalling wings 


Style 


exserted 


included 


Pod, bilocular 


with septum formed 


with septum formed 


and inflated 


by intrusion of 


by a narrow inflex- 




dorsal suture, the 


ion of the dorsal 




ventral filiform 


suture and a deep- 
ly produced wing 
from the ventral 
which extends be- 
low the insertion 
of the funicles 



](i(i CALIFOBXIA ACADEMY OF SCIENCES [Pkoc. 4th ser. 

opinion of the writer the sum of characters peculiar to each isolates these 
species so fundamentally from the rest of Astragalus as to entitle them to 
rank as representative of sections in the genus. The following names are there- 
fore proposed. 

Astragalus sect. Orocystium Barneby sect. iiov. Cystium ser. Platytropia Rydb.. X. Ainer. 
Fl. 24:405. 1929. 

Legumen vesicario-inflatum, siitura dorsali anguste introplicata ventralique intus pio- 
ducta dimidiam loculi altitudinem superanti biloculare: carina oblongo-truncata : 
stipulae circa petiolum abaxialiter concretae, altrinsecus liberae. Species imica : 
A. platytropis A. Gray. 

Astragalus sect. CystieUa Barneby sect. nov. 

Legumen vesicario-inflatum, sutura dorsali introflexa biloculare: stylus ad antliesin 
exsertus: carina attenuata: stipulae multinerviae utrinque concretae cupuliformes. 
Species unica : A. striatiflorus M. E. Jones. 

Following the conclusion reached above that the New World species re- 
ferred to Cystium by Rydberg are probably not allied to the original elements 
of Steven, and as this group has no acceptable sectional name in the form of 
a substantive, I also propose : 

Astragalus sect. Diplocystium Barneby nom. et. stat. nov. Astragalus ser. Astragalus sect. 
Diphysi A. Gray, Proc. Amer. Acad. 6:192. 1863. Astragalus ser. Eu-asiragalus sect. 
Lentiginosus Slield., Minn. Bot. Stud. 1 :168. 1894, exclus. spp. plur. Cystium Rydb., Bull. 
Torr. CI. 32:659. 1905; N. Amer. Fl. 24:405, sequ. 1929, exclus. C. platytropide ; Amer. 
Journ. Bot. 17:232. 1930, non Cystium Stev., nee Astragalus subgen. Cercidothrix sect. 
Cystium Bge. Astragalus sect. Inflati M. E. Jones, Rev. Astrag. 123, sequ. 1923, quoad sp. 
ultimam p. max. p. Astragalus subgen. Uiphysus Tidestr., Contrib. Nat. Herb. 25:3]5. 
1925. Species typica: A. lentiginosus Dougl. ex Hook. 

While some pains have been taken to point out what are believed to be 
fundamental differences between these three sections, the converse and, I 
suppose, more valuable task of tracing their origin and real place in the genus 
cannot be attempted here. In the absence of exact cytological data such specu- 
lation is apt to be valueless and in any case is not susceptible of proof. I am in- 
clined to suspect, nevertheless, that Diplocystium is derived from a common 
source with the Mollissimi: Orocystium and CystieUa seem to be more closely 
connected with each other than with any known species, but, in associating 
A. platytropis with A. cohrensis A. Gray, Jones may have hit on a real line 
of affinity. 



EXPLANATION OF PLATE 



PLATE 17 

Figs. 1-9. Astragalus musimonum Bavuel^y, spec. nov. (1) Pod x 2. (2) 
Cross-section of pod x 3. (3) Flower x 2. (4) Calyx x 2. (5) Banner x 2. (6) 
Wing-petal X 2. (7) Keel x 2. (8) Leaf x 1. (9) Stipules x 2. 

Figs. 10-18. Astragalus Bryantii Barueby, spec. nov. (10) Podxl. (11) 
longitudinal cross-section of same x 1. (12) transverse section of same x 3. (13) 
Pod of broader type x 1. (14) transverse section of same x 3. (15) Flower x 2. 
(16) Wing-petal x 2. (17) Keel X 2. (18) Leaf X 1. 

Figs. 19-24. Astragalus striatiflorus M. E. Jones. (19) Pod x 2. (20) Cross- 
section of pod x 2. (21) Flower X 2. (22) Keel x 2. (23) Seed x 4. (24) Stip- 
ules X 3. 

Figs. 25-28. Astragalus platytropis A. Gray. (25) Cross-section of pod x 1. 
(26) Keel X 2. (27) Stipules x 3. 

Fig. 28. Astragalus en-siformis var. gracilior Barneby, var. nov. (28) pod x 
11/2. 

Figs. 29-30. Astragalus lentiginosus var. ccesariatus Barneby, var. nov. (29) 
Pod x 11/2. (30) Flower X 11/2. 

Fig. 31. Astragalus Whitedii fma. speirocarpoides Barneby fma. nov. (31) 
Pod X 2. 



[168] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 3 



[BARNEBY] PLATE 17 




[ 169 J 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXV, No. 4, pp. 171-176 Jiiiie 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 4 

SUBLITTORAL MARINE ALGAE OF THE 
MONTEREY PENINSULA 

BY 

GILBERT M. SMITH 

Professor of Biology (Botany), Stanford University 



TV TUMEROUS INVESTIGATIONS liBve been made on the marine algae of the Pacific 
1\| Coast of Nortli America but attention has been directed almost exclu- 
sively to those collected in the intertidal zone. Statements concerning sub- 
littoral algae (those growing below extreme low tide level) have been based 
upon the assumption that algae found cast ashore on the beach and not known 
to grow locally in the intertidal zone must have come from deep water. Here 
and there in the literature is mention of a few algae dredged along the western 
shores of North America. The only exception is the region in the vicinity of 
Friday Harbor, Washington, where there has been an intensive exploration 
by dredging. Of the 126 species of red algae that Kylin (1925) records from 
this area, 75 were found only in the intertidal zone, 33 were found only by 
dredging, and 18 were found both in the intertidal zone and in deep water 
(5-10 fathoms). 

For several years I have been engaged upon a survey of the marine algae 
of the Monterey Peninsula, California. In addition to collecting intensively 
in the intertidal zone, numerous collections have been made from a depth of 
15-60 feet below the extreme low tide level. At the time this investigation was 
commenced the only definite records of sublittoral algae from the Monterey 
Peninsula were the following: Dictyo7ieuropsis reticuhifa (Saund.) G. M. 
Smith by Saunders (1895) ; Myriogramme spectahilis (Eaton) Kylin by Nott 
(1900); Macrocystis pyrifera (L.) C. A. Ag. and Nereocysfis Luetkeana 
(Mert.) Post, and Rupr. by McFarland (1912) ; and Lithothamnion mon- 
feveyicnm Foslie by Foslie (1906). 

[171] 




172 CALIFORXIA ACADEMY OF SCIENCES [Proc. 4th sek. 

The first collections from tlie sublittoral by the writer were made with the 
assistance of three students (Vernon Brock, Charles Danforth, and Herbert 
Armstrong) in the summer of 1934. By means of a diving helmet, several algae 
were collected from a depth of 15-35 feet in the vicinity of Mussel Pt. Since 
1934 all collections from deep water have been by dredging from a small boat 
equipped with a 20-horsepower engine. The only really favorable place for 
dredging is the submerged shale reef lying at a depth of 30-35 feet a quarter 
mile east of the municipal wharf at Monterey. Here the rock is so soft that 
large pieces of rock with algae attached to them are broken loose by the dredge. 
Elsewhere on the Monterey Peninsula the rock on the ocean bottom is granitic 
and it is extremely difficult to break off pieces with the dredge. Here, also, 
the ocean bottom is so rough that the dredge is continually catching between 
the rocky crags, and on any dredging expedition one can expect to spend at 
least half the time trying to dislodge the dredge. In spite of these difficulties 
the sublittoral region between Mussel Pt. and Pt. Pinos has been rather thor- 
oughly explored. So, also, has the land-locked cove (Stillwater Cove) adjoin- 
ing Pebble Beach. No attempt has been made to dredge between Pt. Pinos 
and Cypress Pt. because of the danger in taking a small boat close to the shore 
in this wave-swept portion of the coast line. 

When blades of algae are found loose in the dredge there is always the prob- 
lem of distinguishing between those from plants growing in the sublittoral 
and those broken off from intertidal plants and lying unattached on the ocean 
bottom. For this reason every effort was made to obtain intact plants attached 
to pieces of rock and thus know with certainty that the alga is from the sub- 
littoral. In the following list of algae from the sublittoral one or more speci- 
mens of all the species have been found attached to rock dredged from the 
ocean bottom. Some of the algae brought up by the dredge are not found in 
the intertidal zone of the Monterey Peninsula, or are known only from isolated 
individuals collected there. These are considered true sublittoral algae. Other 
■of the algae are regularly found in the intertidal zone. These are interpreted 
as algae the upper limit of whose vertical distribution is approximately the 
mean low tide level. For the sake of comparison the occurrence or non- 
occurrence in the intertidal zone of the Monterey Peninsula is given for each 
of the species, as is also the intertidal range with respect to mean low tide level. 

PHAEOPHYTA 

ECTOCARPALES 

Ectocarpus granulosus (J. E. Smith) C. A. Ag. At 30-35 feet near munici- 
pal wharf, Monterey; at 15-20 feet, Stillwater Cove. Common between the 
0.5 and -1.5 foot tide levels. 

SPHACELARIALES 

Sphacelaria didichotoma Saund. At 15-20 feet, Stillwater Cove. Known 
from the intertidal zone. 



VOL.XXV] SMITH: SUBLITTOEAL ilAEINE ALGAE 173 

DESMARESTIALES 

Desmarestia linearis Gardner, mss. At 60 feet near Monterey. Not known 
from the intertidal zone. 

Desmarestia herhacea (Turn.) Lmx. At 30-35 feet near municipal wharf, 
Monterey. Common between the 0.5 and -1.5 foot tide levels. 

PUNCTARIALES 

Stictyosiphon tortilis (Rupr.) Reinke. At 30-35 feet near municipal wharf, 
Monterey. Not known from the intertidal zone. 

LAMINARIALES 

Dictyoneurum calif or nicum Rupr. At 25 feet near Mussel Pt. ; at 15-20 
feet, Stillwater Cove. Common between the 0.5 and -1.5 foot tide levels. 

Dictyoneuropsis reticulata (Saund.) G. M. Smith. At 30-35 feet near mu- 
nicipal wharf, Monterey; at 20-25 feet near Pt. Aulon. Not known from the 
intertidal zone. 

Nereocystis Luetkeana (Mert.) Post, and Rupr. Numerous beds growing 
at a depth of 30-50 feet. Isolated plants found between the 0.0 and -1.5 foot 
tide levels. 

Macrocystis pyrifera (L.) C. A. Ag. Numerous beds growing at a depth of 
30-50 feet. Not known from the intertidal zone. 

Pterygophora calif ornica Rupr. Growing in extensive stands at a depth of 
20-60 feet between Cypress Pt. and Pescadero Pt. Isolated plants found be- 
tween the 0.0 and -1.5 foot tide levels. 

FUCALES 

Cystoseira osmundacea (Menzies) C. A. Ag. At 20-35 feet near Mussel Pt. 
and near Pt. Aulon. Common between the 0.0 and -1.5 foot tide levels. 

RHODOPHYTA 

BANGIALES 

Go7iiotrichum elegans (Cliauv.) Zanard. At 35-40 feet near municipal 
wharf, Monterey ; at 25-30 feet near Mussel Pt. ; at 20-25 feet near Pescadero 
Rock. Isolated plants collected at mean low tide level. 

Goniotrichopsis siiblitt oralis G. M. Smith. At 30-35 feet near municipal 
wharf, Monterey. Not known from the intertidal zone. 

Erythrotrichia carnea (Dillw.) J. G. Ag. At 30-35 feet near municipal 
wharf, Monterey, and near Pt. Aulon. Common between the 2.0 and -1.5 foot 
tide levels. 

NEMALIONALES 

Rhodochorton concrescens Drew. At 25-30 feet near Pt. Aulon; at 35-40 
feet near Pt. Pinos. Known from the mean low tide level. 

Rhodochorton Amphiroae Drew. At 25-30 feet near Pt. Aulon. Known from 
the mean low tide level. 



174 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th ser. 

CRYPTONEMIALES 

Pikea pinnata Setch. At 30-35 feet near municipal wliai-f , ]\Ionterey ; at 
35-40 feet near Mussel Pt. Isolated plants found between the 0.5 and -1.5 
foot tide levels. 

Pikea californica Harv. At 20-25 feet near Pt. Anion. Widespread but 
scarce between the 0.0 and -1.5 foot tide levels. 

Peyssonnelia pacifica Kylin..At 30-35 feet near municipal wharf. ]\Ion- 
terey; at 20-25 feet, Stillwater Cove. Fairly common between the 0.0 and 
-1.5 foot tide levels. 

Lithothamnion montereyicum Foslie. At 70 feet near Monterey. Not known 
from the intertidal zone. 

Bossea Orhigniana (Dene.) Manza. At 30-35 feet near municipal wharf, 
Monterey ; at 20-25 feet near Pt. Anion. Not known from the intertidal zone. 

Calliarthron Setchelliae Manza. At 20-25 feet near Pt. Aulon; at 30-35 feet 
near Pt. Pinos. Common between the 0.0 and -1.5 foot tide levels. 

Halymenia californica Smith and Hollenb. At 30-35 feet near munici]ial 
wharf, Monterey. Not known from the intertidal zone. 

Prionitis lanceolata Harv. At 40-60 feet near Mussel Pt. ; at 20-25 feet near 
Pt. Aulon and at Stillwater Cove. Abundant between the 1.0 and -1.5 foot 
tide levels. 

Callophyllis marginifructa Setch. and Swezy. At 30-35 feet near municipal 
wharf, Monterey ; at 12-15 feet near Pt. Anion; at 15-20 feet, Stillwater Cove. 
Scarce between the 0.0 and -1.5 foot tide levels. 

CallophyUis megalocarpa Setch. and Swezy. At 15-20 feet near Mussel Pt. 
and at Stillwater Cove. Scarce between the 0.0 and -1.5 foot tide levels. 

Callocolax neglectus Schmitz. At 30-35 feet near municipal wharf. ]Mon- 
terey. Not known from the intertidal zone. 

GIGARTINALES 

Schizymenia pacifica Kylin. At 30-35 feet near municipal wharf, Monterey. 
Common between the 0.0 and -1.5 foot tide levels. 

Agardhiella Coulteri (Harv.) Setch. At 20 feet near Mussel Pt. and at 
Stillwater Cove. Abundant between the 1.0 and -1.5 foot tide levels. 

Plocamivm pacificum Kylin. At 20-25 feet near Pt. Aulon and near Pesea- 
dero Rock. Common between the 0.0 and -1.5 foot tide levels. 

Gracilaria Sjoestcdfii Kylin. At 30-35 feet near municipal wharf, Monterey 
at 40-60 feet near Mussel Pt. Abundant between the 1.0 and -1.5 foot tide 
levels. 

Gracilann rohusta Setch. At 15-25 feet near Pt. Aulon. Isolated plants 
found between the -0.5 and -1.5 foot tide levels. 

Ahnfeltia gigartinoides J. A. Ag. At 12-15 feet near Pt. Aulon. Not known 
from the intertidal zone. 

Stenogramma californica Harv. At 30-35 feet near municipal wharf. ]\Ion- 
terey ; at 20-25 feet near Pt. Anion. "Relatively scarce between the 0.5 and -1.5 
foot tide levels. 



Vol. XXV] SMITH: SVBLIITOBAL MABIXE ALGAE 175 

Gigartina Boryi Setch, and Gardii. At 12-15 feet near Pt. Anion. Common 
between the 1.0 and -1.5 foot tide levels. 

Gigartina corymhifera (Ktz.) J. G. Ag. At 20-25 feet near Mussel Pt. and 
at Stillwater Cove. Common between the 0.0 and -1.5 foot tide levels. 

Gigartina Harveyana (Ktz.) Seteh. and Gardn. At 25 feet near Pt. Anion. 
Common between the 1.0 and -1.5 foot tide levels. 

RHODYMENIALES 

Rhodymenia calif ornica Kylin. At 20-25 feet near Pt. Anion and at Still- 
water Cove. Common between the 0.0 and -1.5 foot tide levels. 

Rhodymenia yaciflca Kylin. At 60-75 feet near Mussel Pt. ; at 15-20 feet, 
Stillwater Cove. Common between the 0.0 and -1.5 foot tide levels. 

Gastroclo7iium Coulteri (Harv.) Kylin. At 20-25 feet near Mussel Pt.; at 
15-20 feet, Stillwater Cove. Abundant between the 2.0 and -1.5 foot tide levels. 

Coeloseira compressa Hollenb. At 20-30 feet near Pt. Anion. Not known 
from the intertidal zone. 

CERAMIALES 

Afitithamnion defectum Kylin. At 30-35 feet near Pt. Pinos. Not known 
from the intertidal zone. 

Antithamnion dendroideum G. M. Smith. At 30-35 feet near municipal 
wharf, Monterey. Not known from the intertidal zone. 

Antithamnion glandidiferum Kylin. At 30-35 feet near municipal wharf, 
Monterey. Rare between the 0.0 and -1.5 foot tide levels. 

Platythamnion heteromorphum J. G. Ag-. At 30-35 feet near municipal 
wharf, Monterey. Not known from the intertidal zone. 

Platythamnion villosum Kylin. At 30-35 feet near municipal wharf, Mon- 
terey ; at 25-30 feet near Pt. Anion. Rare between the 0.0 and -1.5 foot tide 
levels. 

Platythamnion pectinatum Kylin. At 30-35 feet near municipal wharf, 
Monterey. Rare between the 0.0 and -1.5 foot tide levels. 

Pleonosporium dasyoides (J. G. Ag.) DeToni. At 35-50 feet near Pt. Anion; 
at 15-20 feet, Stillwater Cove. Scarce between the 0.0 and -1.5 foot tide levels. 

Pleonosporium vancouverianum J. G. Ag. At 30-35 feet near municipal 
wharf, Monterey. Not known from the intertidal zone. 

Spermothamnion Snyderae Farl. At 30-35 feet near municipal wharf, Mon- 
terey ; at 35-40 feet near Mussel Pt. ; at 30-35 feet near Pt. Pinos ; at 20-25 
feet, Stillwater Cove. Common between the 0.5 and -1.5 foot tide levels. 

Griffithsia pacifica Kylin. At 25-30 feet near Pt. Anion ; at 30-35 feet near 
Pt. Pinos; at 20-25 feet near Pescadero Rock. Scarce between the 0.5 and -1.5 
foot tide levels. 

Microcladia Coidteri Harv. At 20-25 feet near Mussel Pt. and at Stillwater 
Cove. Abundant between the 1.0 and -1.5 foot tide levels. 

Branchioglossum Woodii (J. G. Ag.) Kylin. At 30-35 feet near municipal 
wharf, Monterey, and near Pt. Pinos. Very rare between the -0.5 and -1.5 
foot tide levels. 



17G CALIFORNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

Polyneura latissinia (Harv.) Kylin. At 30-35 feet near municipal wharf, 
Monterey, and near Mussel Pt. ; at 12-15 feet near Pt. Anion. Common be- 
tween the 0.0 and -1.5 foot tide levels. 

Phycodrys Setchellii Skottsb. At 30-35 feet near municipal wharf, Mon- 
terey; at 20-30 feet, Stillwater Cove. Very rare between the 0.0 and -1.5 foot 
tide levels. 

Niendurgia Andersoniana (J. G. Ag.) Kylin. At 20-25 feet near Pt. Anion 
and at Stillwater Cove. Very scarce between the 0.0 and -1.5 foot tide levels. 

Myriogramme spectdbUis (Eaton) Kylin. At 20 feet near Mussel Pt. ; at 
70-90 feet near Pt. Aulon. Not known from tlie intertidal zone. 

Cryptopleura violacea (J. G. Ag:) Kylin. At 30-35 feet near municipal 
wharf, Monterey ; at 20-25 feet near Pt. Aulon ; at 15-20 feet, Stillwater Cove. 
Abundant between the 0.5 and -1.5 foot tide levels. 

Botryoglossum Farlowianum (J. G. Ag.) DeToni. At 30-35 feet near mu- 
nicipal wharf, Monterey; at 20-25 feet near Pt. Aulon; at 15-20 feet, Still- 
water Cove. Abundant between the 0.0 and -1.5 foot tide levels. 

Dasyo'psis densa G. M. Smith. At 30-35 feet near municipal wharf, Mon- 
terey. Not known from the intertidal zone. 

PoJysiphonia calif ornica Harv. At 20-25 feet near Pescadero Rock. Common 
between the 1.0 and -1.5 foot tide levels. 

Polysiphonia flaccidissima var. Smithii Hollenb. At 35-40 feet between 
Mussel Pt. and Pt. Aulon. Not known from the intertidal zone. 

Pterosiphonia dendroidea (Mont.) Falkenb. At 20-25 feet near Pt. Aulon 
and near Pescadero Rock. Abundant between the 0.0 and -1.5 foot tide levels. 

Pterosiphonia Baileyi (Harv.) Falkenb. At 20-25 feet near Mussel Pt. and 
near Pt. Aulon. Common between the 0.5 and -1.5 foot tide levels. 

Hcrposiplionia rigida Gardn. At 30-35 feet near municipal wharf, Mon- 
terey. Rare between the 0.0 and -1.0 foot tide levels. 

HerposipJionia pygmnea Plollenb. niss. At 20-25 feet near Pt. Aulon ; at 
35-40 feet near Pt. Pinos. Not found in the intertidal zone. 

Laurencia spectahilis; Post, and Rupr. At 15-20 feet, Stillwater Cove. 
Abundant between the 1.0 and -1.5 foot tide levels. 

Laurencia Gardneri Hollenb. At 30 feet near Mussel Pt. ; at 35-40 feet near 
Pt. Aulon ; at 30-35 feet near Pt. Pinos ; at 20-25 feet near Pescadero Rock. 
Isolated plants found between the 0.0 and -1.5 foot tide levels. 

BIBLIOGRAPHY 

Foslio, M. 1906. Algologiske Notiser II. Kgl. Norske Vidcnsk. ScLsk. Skr. 1906, No. 2:1-28. 
Kylin, H. 1925. The marine red algae in the vicinity of the Ijiological station at Friday 

Harbor, Wash. Lunds Univ. Irsskr. N. F. 21, Nr, 9:1-87. 47 figp 
McFarland, F. M. 1912. The kelps of the central Californian coast. Appendix M (pp. 194- 

208), Senate Document 190, 2d. Session, 62d. Congress. 
Nott, C. P. 1900. Nitophylla of California. Proc, Calif. Acad. Sci. 3 Ser. Bot., 2:1-62. 9 pi, 
Saunders, DeA. 1895. A preliminary paper on Costaria with description of a new species. 

Rot. Gaz. 20:54-57. 1 pi. 



i 

I 



PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXW No. 5. j)j). 177-188, pis. 18-19 June 1. i()ji 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 5 

THE GANDER OAK, A NEW HYBRID OA 
FROM SAN DIEGO COUNTY, CALIFORNI 

BY 

CARL B. WOLF 

Botanist, Rancho Santa Ana Botanic Garden 



IX THE NEARLY a ceiitiirv and a half since Nee published the first two species 
of Quercus from California (Quercus agrifolia and lohata) the list has 
grrown so that now at least eig"hteen generally recopfnized species are believed 
to constitute the oak flora of the state. In addition to these species there are 
several rare forms which have evoked considerable interest. One of the first 
of tliese to be brought to the attention of botanists was the Oracle Oak ( X 
Quercus morehus Kellogg, Proe. Calif. Acad. Nat. Sci., 2 :36, 1863), long sus- 
pected of being a hybrid between Quercus Kelloggii Newb. and Quercus 
Wislizenii A. DC, but only recently so demonstrated by cultuijil experiments 
at Rancho Santa Ana Botanic Garden (Rancho Santa Ana Bot. Gard.. Occas. 
Papers, 1:47-52. 1938). 

In 1935 Mr. Frank F. Gander, botanist of the Museum of Natural History 
at San Diego, California, discovered another peculiar oak somewhat similar 
to the Oracle Oak. This tree grew along the road between Santa Ysabel and 
Lake Henshaw. Shortly thereafter ]\Ir. Gander informed us of his find and 
we visited the area, located the original tree and several others like it. In the 
fall of 1939 we obtained acorns of this new oak for propagation at Rancho 
Santa Ana Botanic Garden. The results of our studies in the field and in the 
Garden are embodied in tlie present paper. It is a pleasure to publish this 
oak as new in honor of ]\Ir. Gander in the Proceedings of the California 
Academy of Sciences, since one of the parents of this hybrid is Quercus Kel- 
loggii, a species honoring Dr. Albert Kellogg, one of tlie founders of the 
Academy, and also a most devoted student of our oaks. Ilis ])eautiful draw- 

[177] 




178 CALIFOBXIA ACADEMY OF SCIEXCES [Pkoc. 4th ser. 

ings of oaks were published in 1889 (Illustrations of West American Oaks, 
with the text by Dr. E. L. Greene). It has also seemed appropriate that 
this article should appear in the volume dedicated to Miss Alice Eastwood 
because she has long been interested in oaks and published one of the last of 
the generally recognized entities for California {Quercus Alvordmna Eastw., 
Handbook of the Trees of California, p. 48, pi. 27, fig. 4, 1905) . She is a charter 
member of the' Rancho Santa Ana Botanic Garden Advisory Board whei'e her 
enthusiasm for the Garden's experimental work has been a constant stimulus 
for us to go forward with the numerous, painstaking, slow and often dis- 
appointing cultural projects, many of which seem to yield so little in pro- 
portion to the time and energy required for their completion. 

X Quercus Ganderi C. B. Wolf, hybr. nov. 

{Quercus Kelloggi'Newh. X Quercus agrifolia var. oxyadenia [Torr.] J. T. 
Howell.) 

Tree 10-16 m. high; 6-10 m. spread; trunk up to 1 m. in diam., the bark 
dark brown or gray, comparatively smooth and essentially like that of Quercus 
Kelloggii. In aspect the tree is erect, usually is taller than broad and more 
nearly resembles Q. Kelloggii than it does Q. agrifolia var. oxyadenia. Leaves 
nearly evergreen, turning yellow in January and shedding shortly before the 
new leaves appear. Branchlets densely puberulent, but in the second year 
becoming nearly glabrous. Petioles 1-2.5 cm. long; blades coriaceous, slightly 
cupped, glossy green above, but microscopically stellate-pubescent, especially 
along the midribs and veins, gray-green beneath and at first heavily stellate- 
pubescent, but in age becoming much less so, 5-10 cm. long, 3-6 cm. wide, 
shalloAvly and irregularly lobed with 3 or 4 main lobes on each side, these in 
turn irregularly margined and bearing 1^ bristle-tipped teeth. Acorns soli- 
tar}' or 2-3 in a cluster, produced abundantly some years, and apparently 
maturing the first season ; styles 3, about 2-3 mm. long; cup 12-18 mm. high, 
15-25 mm. broad, the inner surface hairy at base, nearly glabrous at margins, 
scales thin, light tan with a darker border and mostly with ciliated margins, 
well imbricated in 6-8 rows; nut with lower one-third covered by the cup, 
oblong, 25-35 mm. long, lightly pubescent to nearly glabrous without, densely 
pubescent within, rich light brown at maturity. (The acorns are scarcely 
distinguishable from those of Q. Kelloggii, except that the nut is somewhat 
slenderer pointed.) Staminate aments slender, pubescent, borne in clusters of 
several ; each ament 4r-6 cm. long, bearing about 15-20 flowers, each with 6-8 
stamens. 

Arbor erecta, oirciter 10-16 m. altitndine; rajiiiilis junioribiis tonientellis ; foliis lialiitus 
Quercus Kelloggii, 5-10 cm. longLs, 3-6 cm. latis; pagiiia superiore sub-glabra, inforiore 
stellata-tomentosa; cupulis 12-18 mm. altitudine, squamis planus; glandibus oblongis, 
25-35 mm. longis. 

Habitat. Lower borders of the Arid Transition and the upper borders of 
the Upper Sonoran Life Zones at elevations of 3000 to 3200 feet, in association 



Vol. XXV] WOLF: THE GANDER OAK 179 

with Quercus Kelloggii, agrifolia var. oxyadenia and Engelmannii, growing 
on hillsides and near occasional streams in granitic type soils. 

Range. Known only from the mountains of San Diego County, California. 
All of the known trees are located in a rather restricted area extending from 
the Volcan Indian School, located north of Santa Ysabel, to a mile or two 
west of Lake Henshaw Dam and to the vicinity of Mesa Grande. 

Type Locality. San Diego Count}', California, about 300 yards north of the 
entrance to the Volcan Indian School on California State Highway No. 79 
between Santa Ysabel and Lake Henshaw, at 3100 feet elevation, in decom- 
posed granite soil, near an occasional stream just west of the highway. Type 
Specimen : Carl B. Wolf and Percij C. Everett, Herb. No. 9543 {Prop. No. 
3555), collected October 20, 1939, the tree being 16 m. high, 10 m. spread, its 
trunk branching into three main forks near the ground, two of these being 
about 46 cm. in diam., and the third about 20 cm. in diam. The Type Specimen 
is deposited in Rancho Santa Ana Botanic Garden Herbarium, sheet no. 23585. 
Isotypes widely distributed as indicated below. 

Localities and Specimens Examined. All of the localities mentioned below 
are in San Diego County, California. 

1). Near the entrance to the Volcan Indian School between Santa Ysabel 
and Lake Henshaw, at 3100 feet elevation: Frank F. Gander, Nov. 14, 1935, 
and May 6, 1937; C. B. Wolf and P. C. Everett, Herh. No. 9483, January 10, 
.1939; P. C. Everett, Herh. No. 9529, July 28, 1939; C. B. Wolf and P. C. 
Everett, Herh. No. 9543 {Prop. No. 3555), Oct. 20, 1939, Type Collection. 
(All of the above specimens were made from the same tree.) 
• 2) . Between Santa Ysabel and Lake Henshaw, on the Morelli Ranch, located 
about 2 miles south of the junction of the road to Warner's Hot Springs and 
Henshaw Dam, 3000 feet elevation : C. B. Wolf, Herh. No. 9487, January 24, 
1939. from a tree 10 m. high, 10 m. spread, and a trunk 60 cm. in diam.; and 
from tlie same tree, C. B. Wolf and P. C. Everett, Herb. No. 9542 {Prop. No. 
5554), October 20, 1939. 

3). Mesa Grande, M. V. Hood, Nov. 24, 1940. 

4) . One and eight-tenths miles from Mesa Grande on the road to Lake Hen- 
shaw, at 3200 feet elevation : C. B. Wolf, Herb. No. 9488, January 24, 1939, 
from a tree 11-12 m. high, 6-7 m. spread, and a trunk 35 cm. in diam. ; C. B. 
Wolf, Herh. No. 9489, January 24, 1939, from a tree 12-13 m. high, 11-12 ra. 
spread, and a trunk nearly 1 m. in diam. 

5) . Between Mesa Grande and Lake Henshaw f north of the above locality) . 
Several trees were noted on January 24, 1939, on the slopes away from the 
road, but for lack of time no collections were made. 

6) . On the hills to the south and west of Lake Henshaw Dam. The trees noted 
in this area on January 24, 1939, appeared to be the northwest limit of dis- 
tribution for the Gander Oak in this region, but for lack of time no specimens 
Avere collected. 

In its native habitat the individual trees of the Gander Oak are difficult to 



180 CALIFOEKIA ACADEMY OF SCIENCES [Proc. 4th ser. 

detect (luring most of the j^ear except upon close examination, for they are 
mixed with the Kellogg Oak, which the.y closely resemble. However, in the late 
fall or early winter they can readily be spotted on the hillsides. At those times 
the leaves of the Kellogg Oak have been shed, while those of the Gander Oak 
are beginning to turn yellow. In January of 1939 we were able to locate be- 
tween 20 and 30 Gander Oaks in the vicinity of Lake Henshaw and Mesa 
Grande merely by driving slowly along the roads that traverse the areas. 
Exploration of the adjacent hillsides not visible from these roads would likely 
result in increasing this figure considerably. It is also probable that the Gan- 
der Oak occurs elsewhere in San Diego County, but thus far no additional 
localities have been brought to our attention. All of the trees of the Gander 
Oak which we have seen in the wild appear to be tirst-generation crosses be- 
tween the two parents, despite the fact that fertile acorns appear to be pro- 
duced somewhat regularly. However, this lack of second-generation offspring 
in the wild is in keeping Avith the behavior of many other wild hybrids. 

Wild trees of X Querciis Ganderi can be distinguished from Quercus Kel- 
loggii by the less deeply lobed and slightly cupped leaves with considerable 
pubescence beneath, and by the longer persistence of the foliage. Separa- 
tion of X Quercus Ganderi from X Quercus morehus is also relatively easy, 
although the leaves of the two are very similar in size and outline. The latter 
species has nearly plane leaves whose under surfaces are practically glabrous. 
Furthermore, X Quercus morehus requires two seasons to mature its acorns, 
whereas, X Quercus Ganderi appears to mature its acorns in a single season 
(as does Querciis agrifolia var. oxyadenia). 

In October, 1941, Mr. John M. Tucker of the Department of Botany of the 
University of California, Berkeley, California, forwarded us an oak specimen 
{Tucker No. 418) collected on the University of California Hastings Wild 
Life Reserve in the Santa Lucia Mountains of Monterey County. This speci- 
men appears to be a hybrid between Quercus Kelloggii and Quercus agrifolia, 
but we have not been able to visit the area and see the tree growing in the wild, 
nor has it been possible to obtain acorns for propagation. Tucker's specimen 
closely resembles X Quercus Gauderi, but is essentially glabrous. The status of 
this oak should be determined by additional field studies and by experimental 
methods. If it should be demonstrated that it is a hybrid, as suggested above, 
it could hardly be included in the present concept of XQnercus Ganderi 
unless that concept be enlarged and Quercus agrifolia var. oxyadenia be re- 
duced to synonomy under Quercus agrifolia. 

Cultural Studies 

Because of the practical difficulties involved in making experimental crosses 
ill Ihe field we have not attempted to produce X Quercus Ganderi by hybridiz- 
ing Quercus Kelloggii and Quercus agrifolia var. oxyadenia. We have, how- 
ever, been able to observe the behavior of seedlings of X Quercus Ganderi 
and feel that they have yielded sufficient data to justify our conclusion that 



Vol. XXV] WOLF: THE GANDEE OAK 181 

the tree is a hybrid between Querciis Kelloggii and Quercus agrifolia var. 
oxyadenia. 

In October, 1939, Mr. Everett and I hand-picked over 400 acorns from the 
Type Tree of X Quercus Ganderi (Herb. No. 9543, Prop. No. 3555), and 11 
acorns from a second tree in the same region (Herh. No. 9542, Prop. No. 3554). 
These acorns were subsequently planted in our lathhouse, where by April, 
1940, there were 311 seedlings of the former and 8 of the latter. In contrast to 
the seedlings of X Qiiercus morehus, grown previously, these were remarkably 
vigorous and attained average heights of over 30 cm. the first season. 

Plantings of 189 seedlings (random selections) of the Gander Oak {Prop. 
No. 3555) w^ere made in the Garden on March 10, 1941. Only 61 of these sur- 
vived to October, 1942. This unusually high loss can be accounted for in part 
because at that time the Botanic Garden Nursery was moved and it was neces- 
sary to dig the seedlings and hold them for considerable time before planting. 
The remainder of the seedlings of this same lot were lined out in nursery rows 
where over 50 are still alive. In the same nursery rows are 5 seedlings of Prop. 
No. 3554. These seedlings in the Garden and in the nursery rows now show 
the usual intergradation of characters displayed when seeds of a hybrid are 
propagated. Those seedlings closely resembling Quercus Kelloggii are now 
few in number, for, under our conditions, that species has always been diffi- 
cult to grow. The rest of the seedlings now alive represent several typical 
Quercus agrifolia var. oxyadenia, a large number of forms very much like the 
original X Quercus Ganderi, and many of varying characteristics. Some have 
the leaf shapes nearly like Quercus Kelloggii, but are exceptionally pubescent. 
Others are like Quercus agrifolia var. oxyadenia, but are glabrous. Still others 
have numerous intermediate leaf characters and are even more unlike either 
of the parent species of the original cross. It will be many years before acorns 
are produced by these seedlings, but they will likely show an almost continu- 
ous series of variations between the slender form produced by the one parent 
and the thick form by the other. It is also probable that some of these seedlings 
will require two seasons to mature acorns while others should do so in a single 
season. 

X Quercus Ganderi may eventually prove of some merit for horticulture, 
for it has many of the fine features of Quercus Kelloggii, plus semi-evergreen 
foliage, a vigorous growth rate and hardiness at low elevations. Acorns gath- 
ered from some of the wild trees could be gi'own in the nursery for two or three 
years, so that selections of outstanding horticultural types could be made 
before making permanent plantings. 

The accompanying drawings on Plates 18 and 19 of Quercus Kelloggii, 
agrifolia var. oxyadenia, Ganderi and the nursery seedlings grown from the 
latter were prepared from tracings (except the drawings of the acorns) . It is 
hoped that they will convey to the reader not only the characteristics of the 
parental species and the original hybrid, but also the great variation exhibited 
by the seedlings. 



182 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th SER. 

DISTRIBUTION OF HEKBARimi SPECBIEXS 

Specimens referred to in the text and in the figures are all on deposit in Eancho Santa 
Ana Botanic Garden Herbarium with the exception of the two specimens by F. F. Gander 
and the one by M. V. Hood, which are in the San Diego Museum of Natural History Her- 
barium. Duplicates of our collections of X Querent Ganderi have been distributed as follows: 

California Academy of Sciences, San Francisco, California : Nos. 9483, 9487, 9488, 9489, 
9529,9543; 

Dudley Herbarium, Stanford University, California : Nos. 9483, 9487, 9488, 9489, 95S9, 
9542,9543; 

Field Museum of Natural History, Chicago, Illinois: No. 9543 ; 

Los Angeles Museum, Los Angeles, California: No. 9543 ; 

Missouri Botanical Garden, St. Louis, Missouri: No. 9543 ; 

New York Botanical Garden, New York City, N. Y. : No. 9543; 

Philadelphia Academy of Natural Sciences, Philadelphia, Penna. : No. 9543 ; 

Pomona College, Claremont, California: No. 9543 ; 

San Diego Museum of Natural History, San Diego, California : Nos. 9483, 9487, 9488, 
9489, 9529, 9543; 

United States National Herbarium, Washington, D. C: Nos. 9483, 9488, 9543 ; 

University of Arizona, Tucson, Arizona: No. 9543 ; 

University of California, Berkeley, California : Nos. 9483, 9487, 9488, 9489, 9529, 9543 ; 

University of California, Los Angeles, California: Nos. 9483, 9543 ; 

University of Colorado, Boulder, Colorado: No. 9543 ; 

University of Montana, Missoula, Montana : No. 9543 ; 

University of Washington, Seattle, Washington : No. 9543. 



EXPLANATION OF PLATES 



PLATE 18 

Figs. 1-24. Leaves of seedlings of X Quercus Ganderi C. B. Wolf ; collected 
from Botanic Garden plantings and nursery rows, January 13, 1943, Prop. 
No. 3555 (grown from acorns taken from Type Tree). 

Figs. 1-7. From seedlings most nearly like Quercus Kelloggii. 

Figs. 8-15. From seedlings most nearly like Quercus agrifolia var. oxya- 
denia. 

Figs. 16-24. From seedlings most nearly like X Quercus Ganderi and inter- 
mediate forms. 



[184] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES, VOL. XXV. NO. 5 [WOLF] PLATE 18 




[185] 



PLATE 19 

Figs. 1-10. Quercus Kelloggii Newb. 

Figs. 11-18. Quercus agrifoliavav. oxyadenia (Torr.) J. T. Howell. 

Figs. 19-30. X Quercus Ganderi C. B. Wolf. 

Fig. 1. C. B. Wolf, Herb. No. 2488 (wild). 

Fig. 2. ,7. T. Howell, Herb. No. 683 (wild). 

Figs. 3 and 4. C. B. Wolf, Herb. No. 3992 (wild). 

Fig. 5. C. B. Wolf and B. D. StarTc, Herb. No. 5482 (wild). 

Fig. 6. P. C. Everett, Herb No. 608 i (nursery seedlings of Prop. No. 1951). 

Fig. 7. C. B. Wolf, Herb. No. 5572 (nursery seedlings of Prop. No. 1951). 

Fig. 8. P. C. Everett, Herb. No. 6078 (nursery seedlings of Prop. No. 1952). 

Fig. 9. B. D. Starlc, Herb. No. 798 (acorn, wild). 

Fig. 10. C. B. Wolf, Herb. No. 2488 (acorn, wild). 

Fig. 11. J. T. Howell, Herb. No. 600 (wild). 

Fig. 12. B. D. StarTc, Herb. No. 5552 (wild). 

Fig. 13. 1. L. Wiggins and J. W. Gillespie, No. 4053 (wild). 

Fig. 14. B. D. StarTc, Herb. No. 5552 (wild). 

Fig. 15. 1. L. Wiggins and J. W. Gillespie, No. 4053 (acorn, wild). 

Fig. 16. B. B. StarTc, Herb. No. 5552 (acorn, Avild). 

Figs. 17 and 18. P. C. Everett, Herb. No. 6074 (nursery seedlings of Prop. 
No. 2026). 

Fig. 19. C. B. Wolf, Herb. No. 9489 (wild, as are also nos. 20-30). 

Fig. 20. C. B. Wolf and P. C. Everett, Herb. No. 9483. 

Fig. 21. C. B. Wolf and P. C. Everett, Herb. No. 9543 (Type). 

Figs. 22 and 23. C. B. Wolf and P. C. Everett, Herb. No. 9543 (acorns from 
the Type). 

Fig. 24. P. C. Everett, Herb. No. 9529 (young acorn). 

Fig. 2.5. C. B. Wolf, Herb. No. 9488. 

Fig. 20. C. B. Wolf, Herb. No. 9489. 

Fig. 27. C. B. Wolf and P. C. Everett, Herb. No. 9542. 

Fig. 28. C. B. Wolf and P. C. Everett, Herb. No. 9543 (Type). 

Fig. 29. C. B. Wolf and P. C.-Everett, Herb. No. 9487. 

Fig. 30. C. B. Wolf and P. C. Everett, Herb. No. 9542. 



[180] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES, VOL. XXV. NO. 5 [WOLF] PLATE 19 




[187] 



PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XX^^ No. 6, pp. 189-214, pis. 20-22 J""c 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 6 

THE GENUS DRYMARIA IN, AND ADJACENT T 

THE SONORAN DESERT 

BY 

IRA L. WIGGINS 

Professor of Biology (Botany), Stanford University 

THE TAXONOMiST frequently finds that it is necessary to study not only 
those represenstatives of a genus that occur within a circumscribed area in 
which he is interested, but numerous other species from contiguous regions as 
well. This necessity became apparent at once when I attempted to construct a 
key to the species of Drymaria known to occur within the limits of the Sonoran 
Desert, for the relationships of several of the desert species were obviously 
with forms growing in the mountains nearby, or in the drier parts of New 
Mexico, Texas, and northern Mexico. As these close relatives were studied, still 
others had to be taken into account until eighteen entities entered into the 
problem. Of this number, only four or five actually grow within the desert, 
but it seemed desirable to present the total results of the study rather than 
to restrict the paper to a treatment of those species which occur regularly in 
the desert. 

Willdenow recognized four species, three of them new, when he set up the 
genus Drymaria in 1819 (in R. and S. Syst. Veg. 5 : 406, 1819), and based the 
new genus on Holosteiim cordatiim Linn. Five years later DeCandolle (Prodr. 
1 : 395, 1924) listed five species, the fifth having been described by Humboldt, 
Bonpland, and Kunth in 1823 (Nov. Gen. et Sp. 6 : 17-21, pL 515-516, 1823). 
Since that time there has been a steady increase in the number of species recog- 
nized in the genus. At present, one hundred nine specific and varietal names 
have been applied in Drymaria. It is probable that a thorough investigation 
of the genus would require reduction of certain combinations and the de- 
scription of other forms not as yet recognized. 

Most of the species and varieties of Drymaria occur in Mexico and the west- 

[ 189 ] 




190 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th SER. 

ern part of South America. Sixty-four, exclusive of the new entities proposed 
liere, have been described from North America, about thirty from South 
America, several from the West Indies and Central America, one species is 
nearly cosmopolitan in the tropics, and one occurs in Australia. Over fifty 
species have been recorded from Mexico. 

The genus includes both annual and perennial herbs, the latter sometimes 
slightly suffrutescent at the base. Most species are clearly annual or perennial, 
with little or no tendency for the annuals to persist beyond a single growing 
season. But occasionally a plant of an annual species ma}', under particularly 
favorable conditions, become a biennial or short-lived perennial. 

The species vary in habit from prostrate, through procumbent and ascend- 
ing, to strictly erect plants. The stems may be simple, unbranched, or pro- 
fusely branched from either the base or in the upper part of the plant. The 
stems and branches are slender, often nearly filiform in such species as D. 
tenella, D. debilis, and D. carinata. The internodes may be rather short, as in 
D. crassifolia, with the leaves exceeding the internodes and forming dense 
mats, or they may be greatly elongated and much surpassing the leaves, as 
in the closely related D. holostcoides. The herbage is pubescent in some species, 
glabrous or glabrate in others. The pubescence may be glandular or non- 
glandular in different species. 

The leaves are useful in separating the genus into two main groups, one 
of which possesses leaves that are ovate, cordate, or broadly elliptic and from 
half as wide as long to distinctly broader than long. The other group has leaves 
that are linear, filiform, or narrowly oblong or subulate. These leaves show 
little tendency toward an elliptic outline and are always several times as long 
as broad. A few species have fleshy or leathery leaves, but most of them have 
thin, herbaceous leaves of delicate texture. 

The arrangement of the flowers in the inflorescence is racemose or cymose. 
The former arrangement is found in D. holosteoides, C. crassifolia, and D. 
pacliyphijlla. In these species the flowers appear to be in dense clusters in 
tiie axils of most leaves and at or near the apices of the branches. Examination 
of this arrangement reveals that the flowers are borne in a short raceme, each 
flower subtended by a small, scarious bract. The minute bracts nearly conceal 
the rachis. Such racemes are axillary to the larger leaves of the fascicles 
formed at each node. In other species studied the inflorescence is cymose. A 
schematic representation of this type of inflorescence as exhibited by most of 
the annual species of Drymaria is shown in PI. 20, fig. 1, and a diagram of 
the inflorescence of D. sperguloides is shown in fig. 2. In the latter figure, the 
small, solidly inked circles represent abortive flowers. 

Floral characters useful in differentiating species of Drymaria include the 
shape, length, comparative length and width, venation, and texture of the 
sepals; the shape, depth of the lobing, length as compared with the length of 
other floral structures, and the presence or absence of appendages between 
the main lobes of the petals ; and the comparative length of claw and blade 



Vol. XXV] If'IGGIXS: DEYMAEIA IX SONURAN DESERT 191 

of the petals. Flowers of comparable age must be selected in making these 
comparisons, however, for the petals are persistent and the mature capsules 
are usually much longer than the ovary at anthesis. 

The aestivation of the sepals is, in all cases examined, cochlear, a modifica- 
tion of the imbricate pattern. In this type of sepal arrangement one sepal 
is wholly outside the others, one has both margins overlapped by adjacent 
sepals, and of the other three each has one margin overlapped by an adjacent 
sepal and the other margin overlapping that of its neighbor. The outermost, 
exposed sepal is on the abaxial side of the calyx, while the one with both mar- 
gins covered is on or near the adaxial side. The margins that are covered by 
the edges of adjacent sepals are almost always more broadly scarious-winged 
than the exposed margins (PL 20, fig. 3). This does not hold in the case of 
D. carinaia and its variety, for in these plants the thicker, herbaceous part 
of the sepals is reduced to a narrow midrib and the sepals are nearly or quite 
symmetrical. 

All species studied have stamens that are slightly dilated at the base to form 
a shallow, connate cup. 

It is doubtful that there is any value in trying to differentiate between 
species with "stipitate" and "sessile" ovaries, for all species examined had 
the ovaries elevated to some degree on a slender, short stipe. Though short, 
this stipe was distinctly demonstrable in every case. 

The seeds are so small that definitive measurements are difficult to make. 
Differences in the depth of the notch adjacent to the hilum, the sculpturing 
of the surface, the over-all pattern of the surface markings, and sometimes 
the color of the seeds can be used to advantage. The seeds of D. viscosa are 
striking in that they are smooth and pale cream or pale buff instead of tes- 
sellated, muriculate, or tuberculate and dark colored as the seeds of most other 
species are. 

During the course of this study the material of Drymaria at the California 
Academy of Sciences, in the herbarium of the University of California, and 
in the Dudley Herbarium at Stanford University has been available con- 
stantly. The types of a number of species were studied, notes taken, and photo- 
graphs of them made during a visit to the herbarium of the Royal Botanic 
Garden at Kew, Surrey, and to the Gray Herbarium of Harvard University 
in 1937. These notes and photographs were valuable aids in determining 
specific limits. The type specimens of Brandegee's species, deposited in the 
herbarium of the University of California, and an isotype of D. depressa 
Greene, in the herbarium of the California Academy of Sciences, clarified the 
status of several poorly known species. My sincere thanks are due, and gladly 
given, to Miss Alice Eastwood for permitting me to borrow material from the 
herbarium of the California Academy of Sciences. Appreciation is hereby 
expressed, also, to Dr. Herbert L. Mason, of the University of California ; 
to Mr. A. D. Cotton, Keeper of the Herbarium of the Royal Botanic Garden, 
Kew, England; and to Dr. M. L. Fernald, of the Gi-av Hei-barinm at Harvard 



192 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th ser. 

University, for the privilege of studying- material and types deposited in the 
herbaria under their supervision. 

In citing specimens, the herbaria in which the specimens are deposited are 
indicated by the following abbreviations : 

California Academy of Sciences CA 

Dudley Herbarium, Stanford University D 

Gray Herbarium, Harvard University G 

Royal Botanic Garden, Kew, England K 

University of California at Berkeley UC 

Taxonomic Treatment 
Drymaria Willd. in R. and S. 

S.yst.Veg. 5:406, 1819. 

Annual or perennial plants wdth slender, prostrate, decumbent, ascending, 
erect or scrambling stems. Branches simple or profusely branched, sparsely 
to densely foliate. Leaves linear, subulate, elliptic, ovate, or cordate, thin and 
herbaceous or thick and fleshy or leathery, glabrous or pubescent, glandular 
or egiandular ; stipules usually present, subulate or linear, persistent or fuga- 
cious. Flowers small, in cymes or racemes, sessile or pedicellate; sepals 5, 
1-3-nerved, similar or slightly dissimilar, the middle portion herbaceous and 
often puberulent or grandular, one or both of the margins scarious-winged ; 
petals (in ours) deeply cleft, in some species bearing additional narrow ap- 
pendages near base of the cleft; stamens 5, the filaments slightly dilated and 
connate at the base ; anthers ovate, much shorter than the filaments. Style 
lender, from much shorter than, to about equal to, the ovary; stigma 3-lobed 
or 3-cleft, the lobes shorter or longer than the style. Capsule ovoid to sub- 
globose, longitudinally dehiscent, mostly 3-valved, 1-celled. Seeds obovoid 
to obovoid-reniform, variously sculptured, most dark brown, dull or slightly 
shining. 

Type species. — Holosteum cordafian Linn. 

Key to the Species 
Leaves about as broad as, or broader than long. 

Flowers in short, crowded, nmbel-liko rncenies; leaves leathery or fleshy, often glau- 
cous; plants mostly prostrate or decumbent. 
Stamens equaling or surpassing the ovary at anthesis; style one-fourth as long 
as, or equaling, the ovary ; petals with 4-R appendages near base of inner 
margins of main lobes. 
Herbage somewhat puberulent, scarcely glaucous; petals 3-3.5 mm. long, 
about equaling the sepals; style 1.4—1.6 mm. long; internodes ex- 
ceeding the leaves 1.7). holosteoides. 

Herbage glabrous, conspicuously glaucous; petals 2-2.2 mm. long, shorter 
than the sepals; style 0.6-0.8 mm. long; internodes mostly shorter 

than the leaves 2. D. eras.ii folia. 

Stamens about half as long as the ovary at anthesis; style less than one-fifth as 
long as the ovary; petals with only one nppendage at base of each mnin 
lobe 3. Z>. pachiiplnjlla. 



Vol. XXV] WIGGINS: DRTMARIA IN HONORAN DESERT 193 

Fln^vers in cymes; leaves thin, not fleshy nor leathery, rarely glaucous; plants mainly 
erect or ascending. 
Sepals strongly carinate, th£ herbaceous mid-portion narrowly linear. 

Claws of petals half as long as, and gradually widening to, the blade; petals 

slightly exceeding the sepals; plant annual 4. D. carinata. 

Claws of petals very short, less than one-sixth as long as, and flaring abruptly 
to, the limb; petals often twice as long as the sepals; plants peren- 
nial 4a. D. carinata var. perennis. 

Sepals not carinate, subequal, the herbaceous mid-portion elliptic. 
Petals surpassing the sepals; sepals not strongly ribbed. 

Sepals oblong-elliptic or narrowly oblanceolate, obtuse at the apex; petals 
5-6.5 mm. long; pubescence of short hairs 0.4 mm. long or less. 
Plants perennial ; leaves shallowly cordate ; sepals glabrous or spar- 
ingly hairy near base 5. D. polystachya. 

Plants annual ; leaves truncate or broadly cuneate at the base ; sepals 
copiously glandular-puberulent on lower two-thirds. 

5a. D. polystachya var. diffusa. 
Sepals broadly ovate, acute at the apex; petals 3.8-4.5 mm. long; pubes- 
cence of crinkled, spreading hairs to 1 mm. long . . .6. D. dehilis. 

Petals shorter than the sepals; sepals strongly 3-ribbed l.B. Fendleri. 

Leaves linear or narrowly oblong, several times as long as broad. 

Herbage heavily viscid-puberulent throughout; seeds smooth, not tessellate nor muricu- 

late 8.D. viscosa. 

Herbage viscid-puberulent only on pedicels and sepals, or glabrous; seeds tessellate, 
muriculate, or otherwise roughened. 
Petals fimbriate-appendaged. 

Herbaceous part of sepals obtuse, glabrous; stigmas subsessile; petals with 
1 lobe at base of each main lobe ; leaves pseudo-verticillate. 

9. D. sperguloides. 
Herbaceous part of sepals acute, glandular-puberulent; stigmas not subses- 
sile; petals with 2-4 appendages at base of each main lobe; leaves 
not pseudo-verticillate. 
Petals 6 mm. long, much exceeding the sepals ; appendages long, slender, 
nearly equaling the main lobes ; leaves narrowly elliptic. 

10. D. arenarioides. 
Petals 4 mm. long, barely exceeding the sepals ; appendages oblong, half 

to two-thirds as long as the main lobes; leaves narrowly linear. 

11. D. peninsidaris. 
Petals merely bifid, without appendages along inner margins of main lobes. 

Petals slightly exceeding the sepals ; plants puberulent. 

Herbaceous part of sepals ol:)tuse ; leaves mostly 5 mm. long or less ; claws 

of petals narrow, naked 12. D. Johnstonii 

Herbaceous part of sepals acute; leaves mostly over 10 mm. long; claws 

of petals broadly winged, as wide as the limb 13. D. effusa. 

Petals shorter than the sepals; plants glabrous, or base of sepals and upper 
parts of internodes sparsely puberulent. 
Stems erect, branching above; leaves narrowly linear, 1 mm. wide or less; 
seeds minutely but sharply muriculate. 

Sepals acute ; herbage glabrous 14. D. tenella. 

Sej)als acuminate ; upper parts of internodes and bases of sepals 

puberulent 14a. D. tenella var. nodosa. 

Stems decumbent or spreading, branched from the base, leaves oblong, to 
3 mm. wide; seeds tessellate with low, rounded bosses. 

15. D. depressa. 



19-4 CALIFORNIA ACADEMY OF SCIENCES [Proc. -Ith ser. 

1. Drymaria holosteoides Benth. Bot. Voy. Sulph. 16, 1844 

(Plate20, figs. 4-12) 

Drymaria Featchii Curran, Proc. Calif. Acad. Sci. (II), 1:227, 1888. 
Mollugophytum holosteoides. M. E. Jones, Extract from Contr. West. Bot. 18 :35, 1933. 

Annual, with slender, spreading, decumbent or prostrate stems 4-40 em. 
long, some of them often stolon-like and forming secondary rosettes at rooted 
nodes, internodes 2-10 cm. long, mostly conspicuously exceeding the leaves, 
spreadingl}^ puberulent and somewhat glandular, older stems glabrate ; stip- 
ules lacking or represented by narrow scarious wing between bases of a leaf- 
pair ; leaves mostly in fascicles of 4-6 at each node, blades spatulate-elliptic, 
3-7 mm. wide, 4—12 mm. long, somewhat fleshy or leathery, faintly 3-nerved 
beneath, glabrous or sparsely puberulent near the base, lightly glaucous, on 
slender petioles 2-10 mm. long; flowers in dense, short racemes in axils of 
most leaves, each flower subtended by a narrow, ovate, hyaline bract about 
1 mm. long; pedicels slender, 3-9 mm. long, glandular-puberulent; sepals obo- 
vate, obtuse or rounded at the apex, 3-3.5 mm. long, the middle portion 
herbaceous, glandular-puberulent at the base, glabrous and faintly glaucous 
above, the scarious margins conspicuous; petals persistent, about equaling the 
sepals, 2.8-3.2 mm. long, white, bifid about half way to the base and bearing 
2—4 narrow segments near the base of inner margin of each main lobe, the claw 
short, broad, somewhat erosulate ; stamens about equaling the petals, nearly 
twice as long as the ovary ; filaments slightly dilated at the base ; ovary about 
1.5 mm. high; style equaling the ovary; capsule broadly ovoid, 1.5-2 mm. 
high ; seeds obovoid-reniform, 0.6-0.8 mm. long, dark brown to black, dull, 
minutely rounded-tuberculate in longitudinal lines. 

Type locality :—"CsLpe San Lucas," Baja California, Mexico. 

Distribution: — Sandy areas from the vicinity of Calmalli to La Paz, and 

possibly to Cape San Lucas; Cedros Island; Magdalena Island; Tiburon 

Island, Mexico. 

Specimens examined: — Magdalena Bay, Hinds, 1841, (K, type. This is the only specimen 
of D. holosteoides in the Kew Herbarium collected on the Voyage of the "Sulphur," and in 
spite of the statement in Bentham's publication to the effect that the plant was obtained at 
"Cape San Lucas" the type locality is probably Magdalena Bay, on Magdalena Island). 
Madgalena Bay, Brandegee, March 1889, (UC) ; Bryant, 1888, (UC, type of D. Featchii) ; 
hills east of Calmalli, Purpus 151, (D, UC) ; rocky hillside 4 miles east of San Ignacio, 
Wiggins 7898, (D, UC) ; 2 miles west of San Ignacio, Eeed 6156, (D) ; Coyote Bay, Con- 
cepcion Bay, Johnston 4178, (CA) ; Gentry 4059, (D) ; Concepcion Bay, Shreve 7095. (J)) ; 
5 miles south of head of Concepcion Bay, Wiggins 5456, (CA, D, UC) ; Mulege, Johnston 
3690, (CA) ; San Luis Gonzales Bay, Johnston 3329, (CA) ; San Francisco Island, John- 
ston 2949, (CA) ; dry washes between Medano and Venancio, Wiggins 5531, (D, UC) ; La 
Paz, Johnston 3048, (CA, UC) ; Brandegee, Feb. 2, 1890, (UC) ; Cedros Island, Stewart 
48-49, (CA) ; Willard's Point, Tiburon Island, Sonora, Johmton 42G3, (CA, D, UC). 

A long-standing confusion concerning the identity of this species and the 
closely related 1). crassifolia was cleared up by Brandegee (Zoe 2: 66-70. 
1891). 



Vol. XXV] WIGGINS: DRYMABIA IN SONOBAN DESEBT 195 

The petals in D. holosteoides are more pronouncedly fimbriate at the base 
of the sinus than they are in D. crassifolia, and the style is nearly or quite 
as long as the ovary in holosteoides, but only half as long in D. crassifolia. 
The densely matted habit of crassifolia is the most readily observed character 
for separating the two species. 

2. Drymaria crassifolia Benth. Bot. Voy. Sulph. 16, 184-1 

(Plate20, figs. 20-26) 
Mollugophyium crassifolium M. E. Jones, Extract from Contr. West. Bot. 18 :35. 1933. 

Prostrate, matted, glabrous and glaucous perennial ; stems profusely 
branched, 3-20 cm. long, internodes 0.5-2.5 cm. long, mostly obscured by the 
numerous, fasciculate leaves; stipules as in D. holosteoides; petioles slender, 
3-9 mm. long; leaf-blades broadly ovate, broadly rhombic-spatulate, or ellip- 
tic, mostly rounded at the apex and broadly cuneate at the base, 2.5-8 mm. 
broad, 4-12 mm. long, fleshy, glaucous, indistinctly 3-nerved beneath; flowers 
numerous in short cluster-like racemes, pedicels 5-12 mm. long ; sepals slightly 
dissimilar, broadly elliptic, rounded, acutish, or faintly emarginate at the 
tip, 1.4—1.8 mm. wide, 3-3.4 mm. long, fleshy, glaucous, faintly veined ; petals 
2-3 mm. long, bifid about to middle, usually with 2 short, rounded or truncate 
appendages at base of inner margin of each main lobe; stamens 1.8-2 mm. 
long, about equaling the ovary; style 0.4—0.6 mm. long, about one-fourth as 
long as the ovary, the stigma-lobes minute ; capsule ovoid, about equaling the 
sepals ; seeds about 0.8 mm. long, obovoid-renif orm, minutely reticulate in 
longitudinal lines, black, dull. 

Tyi)e locality: — Cape San Lucas, Baja California, Mexico. 

Distribution : — Sandy soil and along washes. Cape Region, Baja California. 

Specimens examined: — Cape San Lucas, Hinds, 1841, (K, type) ; Cabo San Lucas, 
Shreve 7258, (D) ; on beach sands, between *the town of Cabo San Lucas and the end of 
the peninsula. Whitehead 906, (D) ; seashore, Todos Santos, M. E. Jones 27046, (D, UC) ; 
San Jose del Cabo, Purpus 265, (UC) ; Anthony, Mar.-June, 1897, (TJC) ; Brandegee, Oct. 
7, 1890, and Nov., 1902, (UC). 

In commenting on D. holosteoides, Johnston wrote (Proc. Calif. Acad. Sci. 
IV. 12 : 1024. 1924), "Drymaria crassifolia . . . is a very closely related form 
known only from San Jose del Cabo, and with little more than its perennial 
habit to distinguish it." It seems to me that the heavily glaucous herbage, the 
shorter stamens, the shorter petals with less strongly fimbriate appendages, 
the shorter internodes and matted habit, are characters of sufficient magni- 
tude to warrant retaining D. crassifolia as a distinct species. Further field 
work may bring to light additional localities in which it grows. 

3. Drymaria pachyphylla Wooton and Standley, Contr. U. S. 
Nat. Herb. 16: 121, 1913 
(Plate 20, figs. 13-19) 
Annual, with slender, spreading, prostrate stems 3-15 cm. long, the inter- 
nodes 2—12 cm. long, much surpassing the leaves, which often are confined 
to terminal tufts ; leaves mostly pseudoverticillate in 4's, the bases of each 



196 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. 4th sek. 

pair connate in a narrow, scarious membrane ; petioles slender, 5-7 mm. long, 
blades broadly elliptic, rhombic-ovate, or broadly ovate, 3-11 mm. wide, 
5-14 mm. lony, somewhat fleshy, green or faintly glaucous, indistinctly 3- 
nerved ; flowers in small, racemose clusters, about 3-15 flowers on each short 
rachis, each subtended by a narrowly ovate, hyaline bract 1.5-2 mm. long; 
pedicels 1-4 mm. long; sepals broadly ovate, 1.5-2 mm. wide, 2.5-3 mm. long, 
rounded at the apex, green and herbaceous, with membranous, white margins 
about 0.2 mm. wide ; petals about equaling or slightly shorter than the sepals, 
bifid one-third of way to base, each lobe bearing a broad, blunt appendage 
half as long as itself near the base of the inner margin ; stamens 1-1.2 mm. 
long, half as long as the ovary; style 0.2 mm. long, the stigmas scarcely 
longer; capsule broadly ovoid to subglobose, about equaling the sepals, yel- 
lowish to pale brown; seeds 0.6-0.8 mm. hmg, minutely granular-tessellate. 
dark brown, dull. 

Type locality: — On the plains south of the White Sands, Dona Ana County, 
New Mexico. 

Distribution: — Sandy soil, southeastern Arizona, through New Mexico to 
western Texas and in adjacent Mexico. 

Specimens examined: — Arizona: Chiricaliua Mts., M. E. Jones, Sept. 22, 1931, (D, UC). 
New Mexico: plains south of the White Sands, Dona Ana Co., Wooion 405, (D, UC, iso- 
types). Mexico: Torreon, Coahuila, Furpus 472, 1903, (UC) ; Filipinas, Coahuila, Furpus 
4942, (UC). 

This is the species that Sereno Watson (Proc. Am. Acad. 17: 329. 1882) 

listed from Coahuila as Z>. crassifolia. 

4. Drymaria carinata Brandegee, Zoe 2:70, 1891 
(PIate20, figs. 27-31) 

Annual, with erect or ascending, weak, slender stems and glabrous herbage 
(sparsely puberulent in the variety) ; stipules narrowly subulate, 2-4 mm. 
long, white, subpersistent ; leaves broadly ovate, the blades 14^16 mm. long, 
5-18 mm. wide, apiculate, from broadly cuneate to shallowly cordate at the 
base, thin, finely but distinctly 3-veined, bright green above, slightly paler 
beneath; petioles half as long as the blades to equaling them; inflorescence 
loosely cymose ; peduncles and pedicels filiform ; pedicels 2-8 mm. long, each 
subtended by an ovate, acute, scarious bract 1-1.5 mm. long; sepals broadly 
ovate-lanceolate, strongly incurved, bluntly carinate, 2-2.5 mm. long, the 
green midvein narrow; petals 1.5-2.5 mm. long, bifid two-thirds of way to 
base, the lobes laiu-eolate to narrowly spatulate, entire or faintly emar- 
ginate, devoid o! marginal appendages; the claw narrow; stamens about 
equaling or slightly exceeding the sepals, filaments linear; ovary 1-1.2 mm. 
high, short-stipitate; capsule subglobose, about equaling the sepals; seeds 
obovate-reniform, 0.4-0.6 mm. long, deep amber to light brown, minutely 
tessellated. 

Type locality: — Sieri-a de la Laguna, Baja California, Mexico. 

Disfrihufioii : — "Mountnins of southoi-n l^>a.ia California. 



Vol. XXV] WIGGINS: DBYMAEIA IN SONOEAN DESERT 197 

Specimens examined: — Sierra de la Laguna, Brandegee 34, Jan. 21, 1890, (UC, type); 
Laguiia Mts., M. E. Jones 24193, (CA) ; Sierra de la Trinidad, Brandegee, Nov. 1902, 
(UC) ; Miraflores, Brandegee, Oct. 14, 1890, (UC) ; El Taste, Brandegee, Sept. 14, 1893, 
(UC) ; San Felipe, Purpus 488, 1901, (UC) ; under overhanging rocks, sides and bottoms 
(if wash 22 miles south of La Paz, Wiggins 5610, (D). 

4a. Drymaria carinata Brand, var. perennis, Wiggins, var. nov. 

(Plate 20, figs. 32-34) 

Ab D. carinata typiea radicibus perennibus, caulibus basim lignosis, nodis 
sparse minutissimeque scaberulosis, lobis petalorum latis ciirtisque differt. 

Type specimen, deposited in the Dudley Herbarium, Stanford University, 
no. 263280, was collected in humus on cliff of rocky canyon slope, in oak-pine 
forest. La Laguna, Sierra de la Laguna, alt. 6000 feet, Baja California, 
Mexico, H. S. Geniry 4415, March 25, 1939. 

Additional specimen examined. — Sierra de San Francisquito, Brandegee, 
March 30, 1892, (UC). 

There is a elearcut and constant difference between the shape and size of 
the petals of the variety perennis and of those of the typical annual form 
(PL 20, figs. 30, 32-34). The lobes of the former are distinctly broader in 
comparison with the length, are practically the same width from base to tip, 
and are more nearly parallel than are those of D. carinata. In perennis, the 
claw is shorter and broader, and flares more abruptly to the blade than in 
the typical material. 

5. Drymaria polystachya Brandegee, Zoe 2:70, 1891 

(Plate21, figs. 2-5) 

Perennial from a gnarled or tortuous, woody root ; stems slender, spread- 
ing to ascending, 1-3 dm. long, freely branched, sparingly to moderately 
pilosulous or glabrate; herbage pubescent and slightly glandular; stipules 
linear-subulate, 1-2 mm. long; leaf -blades broadly cordate, 6-20 mm. wide, 
6-15 mm. long, the apex abruptly short-acuminate or acute, subsessile or on 
petioles 2-7 mm. long; inflorescence of terminal and subterminal cymules; 
peduncles slender, 1-8 cm. long, a pair of narrowly ovate, scarious bracts 
1.6-2.2 mm. long at each node of the inflorescence; pedicels subfiliform, 
0.6-2 cm. long ; sepals subequal, oblong-elliptic, 1.5-1.7 mm. wide, 4.5-5.5 mm. 
long, obtuse or rounded and erosulate at the apex, the broad greenish central 
portion glabrous or sparingly puberulent, faintly 3-nerved, the margins 
scarious; petals bifid about to the middle, 5.5-6.5 mm. long, devoid of append- 
ages, the claws narrow, about 2 mm. long, naked ; stamens about 3 mm. long ; 
ovary ovoid, nearly equaling the stamens; capsule 3.5-^.5 mm. long; seeds 
obovoid-reniform, slightly compressed laterally, brown, finely tessellated in 
numerous longitudinal rows. 

Type locality: — Cliffs near San Jose del Cabo, Baja California, Mexico. 

Distribution: — Among rocks and on cliffs, southern Baja California. 

Specimens examined: — San Jose del Cabo, Brandegee 35, Mar. 14, 1890, (UC, type, D, 



I 

198 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. 4th ser. ; 

isotype) ; Brandegee, March lU, 1892, (UC) ; Grabendorfer, March, 1899, (UC) ; Sieira 
de la Trinidad, Brandegee, Nov., 1902, (UC). 

Brandegee described the leaves as "... 1 em. or less broad, ^2 cm. long, on i 
petioles of nearly the same length." However, most of the leaves are sub- 
sessile or on petioles only 2-3 mm. long. 

The date on the type in the herbarium of the University of California reads 
"March 14, 1890," and the label bears the number "35." A specimen in the 
Dudley Herbarium also bears the number "35," but is dated, "March 24, 
1890." It is not clear whether the discrepanc}' in the dates on the two speci- 
mens is a clerical error made in copying the labels, or two separate collections 
are involved. I am inclined toward the former explanation. No mention of 
the habitat is made on either label, so one must rely solely upon Brandegee's 
statement in the original publication for the information that the plant grew 
on "cliffs" near San Jose del Cabo. 

5a. Drymaria polystachya Brand, var. diffusa (Rose) Wiggins, comb. nov. 

(Plate 21, figs. 6-8) 
Drymaria diffusa Eose, Contr. U. S. Nat. Herb. 1 :130. pi. 12, 1892. 

Annual, with weak, spreadingiy-ascending stems, giandular-puberulent 
throughout; leaves with very slender petioles 4-10 mm. long, the blades 
3-10 mm. wide, 3-8 mm. long, truncate or broadly cuneate at the base ; sepals 
narrowly oblanceolate, about 3.5 mm. long; petals 4.5-5 mm. long; otherwise 
as in the typical form of the species. 

Type locality: — Carmen Island, Baja California, Mexico. 

Distrihiitio7i: — Known only from Carmen Island and the mainland of Baja 
California immediately adjacent. 

Specimens examined: — Carmen Island, Palmer 819, Nov. 1-7, 1890, (UC, 2 isotypes) ; 
shady talus slope under basaltic cliffs, above Punta Escondido, Sierra Giganta, alt. 2000 
to 3000 feet, Gentry 3751, (UC), 

6. Drymaria debilis Brandegee, Proc. Calif. Acad. Sci. II. 2 :131, 1889 

(Plate21, figs. 9, 10) 

Annual ; stems subfiliform, weak, spreading-ascending or scandent, 1-2.5 
dm. long, sparingly pilosulous, the internodes 3—4.5 cm. long; stipules linear- 
subulate, somewhat tortuous, 1-1.5 mm. long ; petioles slender, 2-10 mm. long, 
pilosulous; leaf -blades broadly ovate, 3-10 mm. long and wide, acute or 
abruptly short-acuminate at the apex, abruptly and broadly cuneate or trun- 
cate at the base, sparingly pilosulous on both sides or glabrate above ; in- 
florescence loosely cymose ; pedicels filiform, 4—14 mm. long, sparsely pubes- 
cent; sepals 2.6-3.2 mm. long, pilosulous, ovate-acuminate, or the scarious 
wings obtusish at the apex, faintly 3-nerved; petals 3.8-4.2 mm. long, bifid 
about to the middle, the lobes broadly spatulate, the claws narrow and naked; 
stamens about equaling the sepals, the filamentas linear, flattened; capsule 
ovoid, nearly equaling the sepals ; seeds reddish brown, obovoid-reniform. fi 
0.6-0.8 mm. long, minutely tessellate in fine, longitudinal lines. 



Vol. XXV] WIGGINS: DEYMAEIA IN SONOBAN DESERT 199 

Type locality: Piirisima, Baja California, Mexico. 
Disirihution: — Known only from the type locality. 
Specimens examined: — Purisima, Brandegee, Feb. 13, 1889, (UC, type). 

In general appearance D. debilis closely resembles D. carinata, but differs 
from that species in having acute, broader sepals that are not at all carinate ; 
more pronouncedly pilosulous herbage, and somewhat shorter petals. The 
crinkled hairs of the pubescence are often 1-1.2 mm. long, but there are very 
few glandular hairs mixed with the non-glandular ones. The paucity of glands 
helps to separate D. dehilis from D. polysfachya and its variety, diffusa. 

7. Drymaria Fendleri S. Wats. Proc. Am. Acad. 17:328, 1882 

(Plate21, figs. 11-19) 
Drymaria Fendleri var. perennis M. E. Jones, Extract from Contr. West. Bot. 18:65. 1933. 

Erect or ascending annual (rarely biennial or perennial under favorable 
conditions) with glandular-puberulent herbage; stems slender, 1-3.5 dm. 
long ; internodes to 12 cm. long ; stipules bifid into linear-subulate lobes 1.5-2 
j mm. long, persistent; leaves thin, broadly ovate or ovate-cordate, 4r-15 mm. 
long. 6-22 mm. wide, acute or rounded but abruptly apiculate at the apex, 
truncate or shallowly cordate at the base, on slender petioles 2-7 mm. long, 
the margins often crinkling on drying ; inflorescence cymose ; peduncles 
slender, 1-6 cm. long, usually strongly glandular-puberulent, bearing a pair 
of ovate, scarious bracts 2-2.5 mm. long at the node ; pedicels slender, 1-2 mm. 
long, or some flowers subsessile in 2-5-flowered clusters ; sepals lanceolate, 
1-1.4 mm. wide, 4-5 mm. long, strongly 3-ribbed, acute to acuminate, the cen- 
tral portion green, glandular-puberulent, marginal wings scarious ; petals 
3.5-4 mm. long, bifid about to the middle, the lobes linear-spatulate, claws 
narrow and naked; stamens half as long as the petals; capsule ovoid, 2-2.5 mm. 
high, 8-15-seeded; seeds obovoid-reniform, 0.6-0.8 mm. long, reddish brown, 
minutely tessellated in longitudinal rows. 

Ty2:)e locality: — "New INIexico." 

Bistrihntion: — Foothills and mountains, New Mexico and Arizona to cen- 
tral Mexico, and the Cape Region of Baja California. 

Specimens examined: — Ne-\v Mexico: without further locality, Fendler 60, 1847, (G, 
type) ; Thurber 1073, (G) ; C. Wright 866, 1851, (G) ; La Cuesta, Bigelow, 1853, (G) ; (all 
of the foregoing specimens are mounted on a single herbarium sheet, and all were cited by 
Watson in his original description) ; Silver City, E. L. Greene, Sept. 14, 1880, (UC) ; granite 
hills, Kingston, Sierra County, Metcalfe 1378, (CA) ; under junipers, mountain slope near 
San Jose, San Miguel County, Barnehy 2627, (CA). Arizona: amid big boulders, Prescott, 
Eastwood 16825, (CA) ; near Prescott, Eusche, Sept. 1-5, 1929, (CA). Baja California, 
Mexico : Cota Eanch, Laguna Mts., 14 miles E. of Todos Santos, M. E. Jones 27050, (D, UC, 
isotypes of var. perennis Jones) ; Saucito, near Sierra San Francisquito, Brandegee, Oct. 14, 
1893, (UC) ; Sierra de la Laguna, Brandegee, Jan. 21, 1890, (UC) ; Mirafiores, Brandegee, 
Oct. 14, 1890, (UC) ; Arroyo Hondo, Sierra de la Laguna, Brandegee, Oct. 22, 1893, (UC) ; 
San Felipe, Purpus 469, Jan.-March, 1901, (LTC). Sonora, Mexico: San Bernardo, Rio 
Mayo, Gentry 1338, (UC). 



200 CALIFOSXIA ACADEMY OF SCIENCES [Proc. 4th ser. ; 

The specimens from Baja California are slightly more robust than tliose 
from Arizona and New Mexico, but in qualitative characters the southern 
and northern specimens are identical. A careful search revealed no morpho- 
logical characters accompanying the perennial habit of Jones' material, so his 
variety, D. Fenclleri var. perennis is reduced to synonomy under this species. 

8. Drymaria viscosa S. Wats, ex Orcutt. AVest Am. Sci. 2:57, 

1886, nonien nudum. Proc. Am. Acad. 22:469, 1887 

(Plate21, figs. 20-27) 

Annual with prostrate or spreading branches, dichotomously branched 
stems 5-20 cm. long, and glandular-puberulent leaves, pedicels, and sepals; 
internodes to 6 cm. long, glabrous or sparsely glandular-puberulent ; stipules 
filiform-.subulate, 1-1.5 mm. long, glandular; leaves often pseudo-verticillate, 
linear to linear-oblanceolate, 1.5 mm. wide or less, to 15 mm. long, acute or 
obtuse at the apex, gradually narrowing to a slender petiole 1-3 mm. long; 
inflorescence cymose; peduncles subfiliform, to 3.5 cm. long; pedicels 1-3 mm. 
long or some flowers subsessile ; sepals narrowly to broadly elliptic, 0.6-1.2 
mm. wide, about 2.5 mm. long, the green midrib ending in an excurrent apicu- 
lation slightly below the apex of the scarious, glabrous part ; petals bifid 
about two-thirds of the way to the base, 2 mm. long, the linear-spatulate lobes 
and narrow claw naked ; stamens about 1 mm. long, equaling the ovary ; style 
half as long as the ovary; cap.sule broadly ovoid, nearly equaling the sepals; 
seeds obovoid-eircinate, 0.6 mm. high, pale cream, with light amber angles 
and dorsal band, smooth or slightly wrinkled, but not tessellate nor muricu- 
late, dull. 

Tyije locality: — Socorro, northern Baja California, Mexico. (Misspelled 
"Socono" in the original publication). 

Distribution: — In sandy soil from about 30° 20' N. Lat. (vicinity of San 

Quintm) southward to the Cape Region, Baja California. 

Specimens examined: — Socorro, Orcutl 1330, (G, type, D, UC, isotypes) ; Lagoon Head, 
Palmer 767, March, 1889, (CA) ; Magdalena Island, Orciitf 51, March, 1917, (CA) ; Bran- 
degee, March 13, 1889, (UC) ; mainland near Ascension Island, Anthony, April 17, 1897, 
(UC) ; Cahnalli, Purpun, Jan.-March, 1898, (UC) ; along beach on sand dunes, 4 miles south 
of Guadalupe, Whitehead 830, (D) ; sandy wash 27.4 miles south of Pozo Aleman, Wiggins 
7862, (D) ; San Gregorio, Brandegee, Feb. 1, 1889, (D). 

The light colored, nearly or quite smooth seeds and the heavily viscid herb- 
age of this species are distinctive. In general appearance the plant resembles 
Mollugo cerviana (L.) Seringe. 

9. Drymaria sperguloides A. ( Jray, Am. Acad. Arts and Sci. 

Mem. 11.4:11, 1849 
(Plate 21, figs. 28-38) 
Mollugophytum sperguloides M. E. Jones, Extract from Contr. West. Bot. 18:3o. 1933. 

Slender, erect, simple or dichotomously branched, glabrous annual 0.5-2 
dm. high ; leaves in fascicles of 4-6, pseudo-verticillate, linear, 0.5-3 cm. long, j 



Vol. XXV] WIGGIXS: DRYMABIA IN SONOBAN DESEBT 201 

the stipules trianoiilar-siibiilate, 1-1.5 mm. long, scarioiis ; flowers in terminal 
and subterminal cymnles, on peduncles 0.5-2.5 cm. long; pedicels snbfiliform, 
1-5 mm. long ; sepals dissimilar, narrowly to broadly elliptic, 1-1.5 mm. wide, 
about 2.5 mm. long, cuculate, 1-nerved, the apex rounded or obtuse, central 
green portion terminating slightly below the apex of the scarious margins, 
glabrous; petals 2-2.5 mm. long, bifid about to the middle, 2 linear appendages 
two-thirds as long as the main lobes arising at the bottom of the sinus, claw 
wanged, nearly as broad as the blade ; stamens about 1 mm. long, three-fourths 
as long as the ovoid ovary ; stigma-lobes subsessile ; capsule broadly ovoid to 
subglobose, 3-5 mm. long, exceeding the sepals ; seeds obovoid-circinate, not 
compressed, 1-1.2 mm. long, dark brown, dull, minutely rounded-tuberculate 
in longitudinal rows. 

Ti/pe locality: — "Valley of Santa Fe Creek in the mountains," New Mexico. 

Distrihution: — Mountain slopes and meadows, Arizona to western Texas 
and northern Chihuahua. 

Specimens examined: — New Mexico: without furtlier locality, Fendler 55, 1847, (G, 
type) ; C. Wright 867, (G, UC) ; Mogollon Mountains, on or near west fork of Gila Eiver, 
Socorro County, Meicalfe 349, (D). Texas: near Presidio del Norte, Parry, (G). Arizona: 
Patagonia Mts., Kearney and Peebles 10042, (UC) ; near San Francisco Mts., Fidion 7368, 
(CA) ; EastvieAv, Eincon Mts., Blumer 3583, (D) ; Barfoot Park, Chiricahua Mts., Blumer 
1433, (D). Chihuahua, Mexico: hills and mesas near Guerrero, Pringle 1192, (D). 

The label on the type sheet bears no locality other than "Novo-Mexicanae," 
but the inscription "Drymaria sperguloides n. sp." is in the handwriting of 
Asa Gray, and it is assumed that the more detailed locality data were obtained 
by him from Fendler's notes or through correspondence. 

The large capsules and seeds are distinguishing characters. The pseudo- 
verticillate arrangement of the leaves is sufficient, even in young plants, to 
separate it from D. tenella and D. effusa, both of wdiich have linear leaves 
and a similar habit. 

10. Drymaria arenarioides Willd. in Roem. and Schult. Syst. 

Veg. 5:406, 1819 

(Plate 21, fig. 39; Plate 22, figs. 1-8) 
Drymaria franlcenioides H. B. K. Nov. Gen. et Sp. 6:21. pi. 515, 1823. 

Prostrate, glandular-puberulent, canescent, short-lived perennial with pro- 
fusely branched stems 3-20 cm. long; stipules subulate, 1-1.5 mm. long, 
persistent, slightly glandular-puberulent ; leaves more or less fasciculate, 
linear-lanceolate to narrowly elliptic, 1-2.5 mm. wide, 5-15 mm. long, granu- 
lar-puberulent and slightly canescent, acute, tapering gradually to a short, 
slender petiole 1-2 mm. long; flowers axillary and in terminal, 1-3-flowered 
cymules; pedicels slender, 3-10 mm. long, often deflexed in fruit; sepals dis- 
similar, narrowly lanceolate to ovate-lanceolate, 1-2.2 mm. wide, 5-6 mm. 
long, acute to short-acuminate, weakly 1-nerved, the margins scarious ; petals 
5-6.5 mm. long, divided into 6 lobes about to the middle, the main, outer ones 



202 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th sek. 

1-1.2 mm. wide, the inner 4 in 2 pairs of filiform appendages slightly shorter 
than the main lobes; stamens about 1.5 mm. long, equaling the ovary; style 
half as long as the ovary; capsule ovoid, 4—5 mm. long; seeds orbieular-eir- 
cinate, slightly compressed laterally, with a pronounced, broad, dorsal groove 
and deep lateral depressions, uniformly and minutely granular-tessellate, 
dull, brown. 

Tyjye locality: — Pachuca, Mexico. 

. Distrihution : — Northern Sonora and Chihuahua and southward into cen- 
tral Mexico. 

Specimens examined: — Chihuahua: hills and plains near Chihuahua (City), Pringle 715, 
(UC) ; Chihuahua, Purpus 1123, (UC). Sonora: gravelly slopes 20 miles south of Llano, 
Wiggins 7235, (D) ; Shreve 7322, (D) ; southern edge of oak belt 35 km. south of Nogales, 
Ferris 8810, (D) ; rocky benches, Nogales road, about 1 mile north of Cumetal, Ahrams 
13178, (D). Hidalgo: Ixmiquilpan, Purpus 483, Feb., 1905, (UC). Zacatecas: Purpus 133, 
1903, (UC). 

The filiform central lobes of the petals are surprisingly uniform in length, 
width, and point of insertion. There is a visible vascular strand, similar to 
those running to the main lobes, leading to each pair of the appendages. These 
bundles depart from the main petal bundle just below the sinus. 

The seeds are distinctive in that the tessellations are not arranged in rows, 
but form a low, close mosaic over the whole surface. The broad, dorsally 
grooved back, and the tight coil of the embryo are noteworthy also. 

ll.Drymaria peninsularis Blake, Journ. "Wash. Acad. Sci. 14:285, 192-4 

(Plate22, figs. 9-16) 

Prostrate, decumbent, or ascending annual, or sometimes persisting as a 
perennial, with many freely branched stems 0.5-2 dm. long; herbage pale 
green or slightly glaucescent, glandular-hirtellous throughout ; internodes 
2-25 mm. long; stipules scarious, undulate, entire, 0.6-0.8 mm. long; leaves 
linear, somewhat fleshy, sessile, obtuse or rarely acute, lower ones 1-2.5 cm. 
long, the upper rarely over 4 mm. long, shorter than the internodes ; flowers 
solitary in the axils from about the middle of the stems to the apex ; pedicels 
2-12 mm. long (in fruit) , erect to strongly deflexed ; sepals slightly dissimilar, 
narrowly to broadly ovate-oblong, 1-2 mm. wide, 3.5-4 mm. long at anthesis, 
4r^.5 mm. long in fruit, obtuse or rounded, obscurely 1- or 3-nerved, the cen- 
tral part green, herbaceous and glandular-puberulent, the margins scarious, 
glabrous ; petals white, 3.5-4 mm. long, curved, so just equaling the sepals, 
4— 5-fid nearly to the middle, the outer lobes broader and longer than the cen- 
tral ones, claws slightly dilated and erosulate near the base, nearly as broad 
as the limb ; stamens about two-thirds as long as the sepals, slightly surpassing 
the ovary; style and stigmatic lobes about equal ; capsule ovoid, 3-4 mm. long ; 
seeds round-reniform, plump, 0.7-0.9 mm. long, fuscous, finely and bluntly 
tessellate, the bosses near the base of the dorsal surface in longitudinal rows, 
not aligned elsewhere. 



Vol. XXV] WIGGINS: DRYMARIA IN SONOBAN DESERT 203 

Type locality: — Cape Region, Baja California, Mexico. 
Distribution: — Known only from the Cape Region of Baja California, 
Mexico. 

Specimens ei-amined : Cape Region, Pur pun 423, (UC, isotype) ; Cabo San Lucas, Bran- 
degee 30, March 17, 1892, (D, UC) ; Brandegee, Nov., 1902, (UC) ; Canyon Salado, Purpus 
337, (UC) ; coast south of Pescadero, Brandegee, Nov., 1902, (UC) ; Arroyo Salado (San 
Jose del Cabo), Purpus 496, (UC) ; El Taste, Brandegee, Sept. 11, 1893, (UC) ; Cabo San 
Lucas, Johansen 541, (D). 

The leaves are narrower and the upper ones more reduced in Z>. peninsnlaris 
than they are in D. arenarioides, with which the former was long identified. 
The sepals of D. arenarioides are more acute than they are in D. peninsnlaris, 
and the appendages in the sinuses of the petals are longer and narrower in 
the former species than they are in the latter. The seeds of the two species 
are quite similar. 

12. Drymaria Johnstonii Wiggins, sp. nov. 

(Plate22, figs. 17-21, 36) 

Planta annua gracilis, ramulis decumbentibus vel ascendentibus glaucis 
5-10 em. longis gianduloso-puberulentibus ; stipulae subulatae 0.5-0.8 mm. 
longae ; folia opposita vel f asciculata linearia subcarnosa 2-5 mm. longa ses- 
silia apicem obtusa vel rotundata ; tlores axillarii solitarii, sepalis 2.2-2.5 mm. 
longis ellipticis obtusis, margine membraneis, petalis semibifidis 2.5-3 mm. 
longis albis nudis sepala vix superantibus; staminibus ovarium vix superanti- 
bus ; ovarium 1 mm. altum ; capsula anguste ovata 2.5-3 mm. longa sepala 
vix superantia; semina 2-5 nigra obovata minute granulo-muriculata 0.6 mm. 
longa. 

Annual, with freely branching, slender, decumbent or spreadingly ascend- 
ing stems 1-1.5 dm. long and slightly fleshy, glaucescent leaves, puberulent 
throughout with stout, short-stipitate, whitish glandular and simple hairs; 
internodes 5-14 mm. long; stipules subulate, scarious, 0.5-0.8 mm. long, 
persistent; leaves opposite or the lower ones sometimes fasciculate, linear, 
slightly fleshy, 2—5 mm. long, or basal ones up to 1 cm. long, faintly 1 -nerved, 
sessile, rounded or obtuse at the apex; flowers axillary, solitary; pedicels 
slender, 1-2 mm. long at anthesis, 4-8 mm. long in fruit ; sepals herbaceous- 
fleshy. 2.2-2.5 mm. long, elliptic, obtuse at the apex, the scarious, hyaline 
margins evident on the inner sepals and the covered edges of the outer sepals; 
petals white, bifid nearly to the middle, devoid of appendages, 2.5-3 mm. 
long, slightly surpassing the sepals ; stamens equaling or slightly surpassing 
the ovary ; style 0.2 mm. long or less, much exceeded by the stigma-lobes ; 
capsule narrowly ovoid, acute, 2.5-3 mm. long, equaling the sepals or sligh'tly 
exceeding them, 2-5-seeded ; seeds black, 0.6 mm. long, obovoid-reniform, 
minutely and uniformly granular-muriculate, dull. 

Tyjye in the herbarium of the California Academy of Sciences no. 82814, 
collected in crevices of rock on mesa near crest of island, the isthmus, Espiritu 
Santo Island, Baja California, Mexico, Ivan M. Johnston 3972, May 31, 1921. 



204 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Only one plant was seen and the species has not been re-eollected. Johnston 
(Proc. Calif. Acad. Sci. [IV], 12:1023, 1924) determined it, tentatively, as 
D. arenarioides and stated that the plant seemed undescribed. He also said 
that the plant on the mainland of Baja California which had been called 
D. arenarioides was different, both from his Espiritn Island plant and the 
material called I), arenarioides from eastern Mexico. Subsequently S. F. Blake 
described the peninsula plant as D. peninsularis, but he did not have access 
to this material of Johnston's. 

D. Johnsfonii closely resembles a depauperate plant of D. peninsidaris, but 
differs from that species in having few leaves over 4—5 mm. long, petals en- 
tirely devoid of appendages, and in having seeds that are less tightly cirei- 
nate, ungrooved dorsally, and in being much more finely muriculate over the 
Avliole surface. 

13. Drymaria effusa A. Gray, PI. Wright. 2:19, 1853 
Slender annual 0.8-2 dm. high, dichotomously branched above the second 
or third nodes, minutely puberulent, or the internodes becoming glabrate; 
internodes 2-4 cm. long, or the lowest slightly longer; stipules filiform- 
subulate, 1.5-2.2 mm. long, entire; basal leaves obovate-spatulate, 1.5-3 mm. 
wide, 3-10 mm. long, rounded, broadly cuneate at the base, green, sparsely 
puberulent, on petioles 1-2 mm. long, or subsessile ; cauline leaves filiform 
or narrowly linear, sessile, 1-2.5 cm. long, or the uppermost reduced, faintly 
1-nerved, minutely puberulent ; flowers solitary in the axils of the upper 
leaves and of scarious, ovate bracts 0.5-1.5 mm. long; pedicels filiform. 2-4 
mm. long ; sepals ovate, slightly dissimilar, 1.4-1.8 mm. long, obtuse or rounded 
at the apex, inconspicuously 1-nerved, minutely but closely puberulent. the 
winged margins scarious; petals bifid almost to the center, 2-2.8 mm. long, 
exceeding the sepals, devoid of appendages; claws narrow and naked; sta- 
mens about equaling the sepals ; ovary 1 mm. high ; style 0.4—0.6 mm. long, 
about equaled by the filiform stigma-lobes; seeds (immature) reddish brown, 
muriculate. 

Tyj^e locality: — "Mountains east of Santa Cruz, Sonora." 
Distribution: — Mountains in Santa Cruz, Cochise, and Pima counties, Ari- 
zona, and southward into adjacent northern Sonora. 

Specimens examined: — Sonora : mountains east of Santa Cruz, C. Wright 869, Sept., 1851, 
(G, type). Arizona: Babocomari, Santa Cruz County, Thiirber 995, (G) ; Sonoita Valley, 
BoihrocTc 169, (G) ; "S. Arizona," Lemmon 509, 1881, (G) ; Huachuca Mountains, Price, 
Aug. 14,1895, (D). 

14. Drymaria tenella A. Gray, Mem. Am. Acad. IT. 4 : 12, 1849 

(Plate 22, figs. 22-27, 34) 

Erect, slender annual 0.5-2 dm. high, dichotomously branched above the 
second or third node, glabrous, or glandular puberulent in the variety; inter- 
nodes to 3.5 cm. long; basal leaves spatulate-obovate, 1-2.5 mm. long on pedi- 
cels of equal length ; cauline leaves linear, sessile, 3-15 mm. long, acute, in- 



Vol. XXV] WIGGIXS: DBYMARIA IN SOXORAN DESERT 205 

conspicuously 1-nerved, glabrous; stipules white, scarious, linear-subulate, 

1 mm. long- or less, erect ; flowers sliort-peclicellate or subsessile in the axils 
or terminal on slender peduncles 5-10 mm. long ; sepals ovate-lanceolate, 
0.8-1.2 mm. wide, 2.8-3.2 mm. long, acute, strongly 3-ribbed, the margins 
.scarious; petals white, 2.3-2.8 mm. long, bifid to middle or slightly below, 
the lobes oblong, entire or minutely emarginate, devoid of appendages, claws 
narrow and naked; stamens slender, about equaling the 1-1.3 mm. high ovary; 
style 0.2-0.4 mm. long, stigmatic lobes about as long; capsule ovoid, about 

2 mm. high, 8-12-seeded ; seeds obovoid-reniform, 0.6-0.8 mm. long, minutely 
but sharply muriculate, reddish brown. 

Tyjie locality: — Eight miles west of Las Vegas, New Mexico. 

Distribution: — Mountains of Arizona, New Mexico, and adjacent Chihua- 
hua and Sonora, and in northern Baja California, Mexico. 

Specimens examined: — New Mexico: eight miles west of Las Vegas, Fendler 56, Aug., 
1847, (G, type) ; Pinos Altos Mts., E. L. Greene 332, Sept. 8, 1880, (G) ; Sawyer's Peak, 
Grant County, Metcalfe 1428, (CA) ; \vithout definite locality, C. Wright 868, 1851, (G). 
Arizona: Manning Camp, Eincon Mts., Pima County, Bhnner 3368, (D) ; rliyolite spur just 
N. of Wilgus Ranch, Chiricahua Mts., Cochise County, Blumer 1639, (D) ; Barfoot Park, 
Chiricahua Mts., Blumer 1411, (D). Chihuahua: pine plains, base of the Sierra Madre, 
Pringle 1194, (D). Baja California, Mexico: margins of meadow at La Encantada, Sierra 
San Pedro Martir, alt. 2200 m., Wiggins and Demaree 4915, (D). 

This species is very similar to B. effiisa in general appearance, but it is 
separated from that species by the acuminate sepals and the lack of puberu- 
lence on the herbage. 

14a. Drymaria tenella A. Gray, var. nodosa (Engelm.), Wiggins, n. comb, 

(Plate 22, figs. 28-33, 37-38) 
Drymaria nodosa Engelm. apud A. Gray, Mem. Am. Acad. II. 4:12, 1849. 

Upper parts of internodes, peduncles, pedicels and herbaceous portions of 
the sepals minutely glandular-puberulent ; sepals short-acuminate ; otherwise 
as in typical I), tenella. 

Type locality: — Cosiquiriachi, Chihuahua, Mexico. 

Distribution : — Northern Chihuahua and northeastern Sonora. 

Specimens examined: — Chihuahua: pine plains, base of the Sierra Madre, Pringle 1195, 
(D) ; thin soil, rocky hills near Chihuahua, Pringle 716, (D). Sonora: Los Pinitos, alt. 6100 
ft.. Hart man 138, (UC). 

There are only two minor characters by which the variety can be separated 
from D. tenella, and they are of insufficient value to warrant retaining this 
entity as a distinct species. 

15. Drymaria depressa Greene, Leafl. Bot. Obs. and Crit. 1 : 153, 1905 

(Plate22, figs. 39-43) 

Annual, freely branched plant with slender branches 2-4 cm. long, de- 
pressed or spreading, glabrous, the internodes 0.5-2 cm. long ; stipules 0.5-1.2 
mm. long, whitish, slightly curled; leaves oblanceolate, 1.5-3 mm. wide. 



206 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

5-13 mm. long, obtuse at the apex, graduall}' narrowing to the sessile base, 
glabrous, faintly 3-nerved ; inflorescence of compact cymules; pedicels 1.5 
mm. long or less, minutely granular-puberulent ; sepals oblong, 2.2-2.6 mm. 
long, rounded at the apex, narrowly scarious-margined, glabrous, distinctly 
3-nerved; petals narrow, bifid to the middle, 2-2.2 mm. long, devoid of append- 
ages, the claws narrow, naked ; stamens slightly over half as long as the petals, 
and about equaling the ovary at anthesis; ovary short-stipitate ; style nearly 
as long as the ovary, the stig-ma-lobes half as long ; capsule narrowly ovoid, 
nearly equaling the sepals at maturity, 8-12-seeded ; seeds obovoid-renif orm, 
0.6-0.8 mm. long, light buff, uniformly and finely tessellate with low, rounded 
welts, dull. 

Tyjje local a y: — Sawyer's Peak, Grant County, New Mexico. 

Distribution: — Mountains in eastern Arizona and New Mexico. 

Specimens examined: — Open glade on Sawyer's Peak, Grant County, New Mexico, alt. 
9500 feet, Metcalfe 1430, (CA, isotype). 

Kearney and Peebles suggest that this species may not be distinct from D. 
effusa, but it seems to me that the broader leaves, compact habit, and lack or 
paucity of puberulence separates it from that species. I have not seen the Ari- 
zona material. 



EXPLANATION OF PLATES 



PLATE 20 
Fig. 1. Diagram of cyniose inflorescence common in Drymaria. 

Fig. 2. Diagram of inflorescence of D. sprrguloides; solidly inked circles 
indicate abortive flowers. 

Fig. 3. Diagram of aestivation of sepals in Drymaria. 

Figs. 4-12. I), holosteoides ; figs. 4-8, sepals, x 6.6; fig. 9, petal, x 13; figs. 
1(1-12, X 6.6; fig. 10, ovary and stamens; fig. 11, ovary; fig. 12, petal. 

Figs. 13-19. D. pachyphylla, x 6.6; fig. 13, petal; fig. 14, ovary and stamens; 
figs. 15-19, sepals. 

Figs. 20-26. D. crassifolia, x 6.6; fig. 20, ovary and stamens; fig. 21, petal; 
figs. 22-26, sepals. 

Figs. 27-31. B. carinata, figs. 27-29, x 6.6; figs. 30-31, x 13; fig. 27, petal; 
figs. 28-29, sepals; figs. 30-31, petal and sepal from different plant. 

Figs. 32-34. D. carinata var. pereniiis, x 13; fig. 32, petal before anthesis; 
figs. 33-34, petals from different plants at anthesis. 



[ 208 ] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 6 [WIGGINS] PLATE 20 




[ 209 ] 



PLATE 21 

Fig. 1. D. crassifolia, x 6.6 ; arrangement of leaves at a node, the dotted lines 
indicate a hyaline "sheath" of epidermal and subepidermal cells overlying the 
photosynthetic tissue. 

Figs. 2-5. D. polystachya, petal and sepals, x 6.6. 

Figs. 6-8. D. polystachya var. diffusa, sepals and petal, X 6.6. 

Figs. 9-10. D. debilis, petal and sepal, x 6.6. 

Figs. 11-19. Z>. Fendleri, x6.6; figs. 11-12, petal and ovary and stamens 
from a flower from New Mexico ; fig. 13, mature capsule from same plant; figs. 
14-19, sepals and petal from plant from Baja California, Mexico. 

Figs. 20-27. D. viscosa, x 13; fig. 20, petal; fig. 27, ovary and stamens; figs. 
21-26, sepals (fig. 22, lateral view of a sepal). 

Figs. 28-38. B. sperguloides, x 13 ; figs. 28-31, 35-36, sepals ; figs. 32-33, 
petals ; fig. 34, ovary and stamens ; fig. 37, mature capsule and persistent sepals; 
fig. 38, arrangement of leaves and branches at a node. 

Fig. 39. D. arevarioides, petal, x 6.6. 



[210] 



PROC. CALIF, ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 6 [WIGGINS] PLATE 21 





[211] 



PLATE 22 

rigs. 1-8. D. arenarioides, x 6.6; figs. 1-5, sepals; fig. 6, ovary and stamens; 
figs. 7-8, petals. 

Figs. 9-16. D. peninsularis, xS.6; figs. 9-13, sepals; figs. 14-10, petals. 

Figs. 17-21, 36. D. Johnstonii, x 13; fig. 17, flower at antliesis; figs. 18-19, 
sepals; fig. 20, ovary and stamens; figs. 21, 36, petals and adjacent stamens. 

Figs. 22-27, 34. B. tenella, x 6.6; figs. 22-26, sepals; fig. 27, ovary and sta- 
mens ; fig. 34, petal. 

Figs. 28-33, 37-38. D. tenella var. nodosa, x 6.6; figs. 28-30, 37-38, sepals; 
fig. 31, ovary and stamens; figs. 32-33, petals. 

Figs. 39-43. P. depressa, x 13 ; fig. 39, petal ; figs. 40-43, scpaLs. 



[212] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV, NO. 6 [WIGGINS] PLATE 22 




[213] 



PROCEEDINGS 



OF THE 

CALIFORNIA ACADEMY OF SCIENCES 



Fourth Series 
Vol. XXV, No. 7, pp. 215-220 



June 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 7 
WATER AND PLANT ANATOMY 

BY 

GEORGE J. PEIRCE 

Professor of Botany, Emeritus 
Stanford University, California 



FORTUNATELY the processcs of Nature go on whether man understands them 
or not ; but curiosity, which has led man to enquire about everything inside 
and outside himself, has brought not only the satisfaction sought, but has 
stimulated continued enquiry, has led to the development of powers formerly 
unsuspected, and to the recognition of reasons why the powers are not un- 
limited. It has been discovered that living things require air to breathe, water 
to absorb, food to take in ; what it is in air, water, food which is indispensable 
and not merely stimulating; and some of the relations between the worlds 
outside and inside the living thing. One of these relations concerns itself with 
the intake and outgo of one or other of these things — air, water, food. In this 
paper in honor of an associate of many years I wish to consider one of these 
relationships, thereby contributing also to an understanding, or at least a 
conjecture, as to why grass, chaparral, or forest should be the natural cover 
of respective areas. 

The body of the higher land plants consists of tissues among which the vari- 
ous functions are divided and by which the various needs are met ; but, while 
there is division of labor, each tissue and each organ of the plant may perform 
secondary functions, or be the seat of subsidiary phenomena. Thus the green 
leaf, primarily for the production of food, is the seat of maximum gas ex- 
change and of water loss. The green leaf represents in its shape, size, dis- 
tribution, anatomy, duration, and even number, the adjustment which each 
individual leaf, as well as the whole plant, makes among these, and many 
other, less evident, phenomena. Similarly the root, primarily for the meehani- 

[ 215 ] 




216 CALIFORNIA ACADEMY OF SCIENCES [Proc. -Ith ser. 

cal support of the plant and the intake of water, must provide also for passing 
the water on to other parts of the body. But obviously these tasks vary accord- 
ing to diverse or changing external conditions, the intake of water from moist 
soil in the rainy season appearing to be easier and more understandable than 
by the same root from the same soil in the dry season. The facile statements 
that water intake by roots is an osmotic phenomenon, or that "work is done,'' 
ignore the fact that, under conditions very different from those of well- 
watered Europe, the relative densities of the solutions in root and in soil are 
also very different. 

In order to provide the path of water movement from soil to air through 
the body of the plant there must be a vascular system, again so flexible that it 
will meet the need of the plant in all of its circumstances, ranging from satu- 
rated soil and air to soil and air almost dry. Combining mechanical support 
and water lift, one finds the vascular tissues variously strengthened and the 
vascular tissues themselves variously composed of elements as different as 
the small-one-celled tracheids of the wood of the conifers and the large many- 
celled vessels of the annual herbaceous plants of the rainy season. Further- 
more, the same individual plant may regularly produce different conducting 
tissue in the same year according to the external environment. 

I should like now to cite some of the evidence on which these statements rest, 
specifically in regard to the conducting tissues of the vascular land plants. 

As shown in previous papers,' the quantity and the physical state of water 
in the conducting tissues of plants, and their parts, varies from a liquid exert- 
ing outward pressure to a vapor under internal tension or suction : from 
columns of liquid water more or less completely filling the lumina of ducts 
and tracheids to sheets of liquid water on and in the material of the walls of 
ducts and tracheids, and enclosing masses of water vapor filling the lumina 
of the vascular elements. We are familiar with the phenomena of bleeding, 
which are the evidence of internal pressure, and only less familiar with the 
phenomena of transpiration or evaporation, which result in tension (trac- 
tion) within the water mass in the plant bod5\ At seasonal or other intervals 
the water in the vascular system of the land plant undergoes these complete 
changes, from liquid to vapor, from pressure to tension ("negative pressure") . 
Perennial land plants must survive these changes or succumb, either locally 
or extensively. Many annual plants cannot maintain themselves when the 
procession of the seasons from wet to dr}^ entails these changes in the quantity 
and the state of the water within their conducting systems. Examination 
reveals that their vascular anatomy does not meet the requirements of ade- 
quate water holding and water lifting. 

Between the so-called "water reservoirs,'" which are cells and tissues evi- 
dently enlarged and otherwise fitted for containing water in their vacuoles, 

^ Peirce, G. J. The state of water in ducts and tracheids. Plant Physiology, 11, 1936, and 
literature there cited. 

' See Haberlandt, G. Physiologische Pflanzenanatomie, p. 371 et seq. Ctli ed. 1924 or later, 
on Avater storage and water storing tissues. 



Vol. XXV] PEIRCE: WATER AND PLANT AN ATOMY 217 

and the -vvet walls which not merely contain but actually hold water with great 
tenacity, there are many intermediates, differing from one another as do the 
environments in which the plants live. These wet walls vary in thickness as 
they do in hardness and stiffness. They may look as if they were mechanically 
strong where little or no mechanical strain can develop ; but they may show 
by their chemical composition that they fix — adsorb and incorporate — water, 
perhaps in the manner in which the water of crystallization is incorporated 
in blue copper sulphate. It is recognized that the crystalline form and the 
crystalline state of copper sulphate depend upon the two constituents of the 
crystals, water and copper sulphate. Similarly, but to a certain point only, 
and also only partially, the shape, and the thickness, of cell walls depend both 
on the chemical composition (cellulose derivatives and their accompaniments) 
of the cell wall and water. Collenchyma tissues, which are mechanically weak 
though looking as if they might have tensile or compressive resistance, do store 
amounts of water which are large in proportion to the mass. The amount of 
heat required to drive off all of this water indicates the force with which this 
water is bound : and the shrinkage shows no less plainly how much the form 
depends upon the constituent water. 

Turning now to specific instances as illustrating these general statements, 
. one may consider Montia perfoliata (Donn) Howell, "Miner's lettuce," a suc- 
culent, low, annual herb which begins soon after the first rains have consid- 
erably wet the soil of natural and cultivated areas, and which expires while 
the moisture content of the air and of the soil is still high. The appearance of 
the root system suggests that it is adequate as an absorbing organ ; the area 
and composition of the leaves and their surface are such that transpiration 
(evaporation) into warm, dry air, while considerable, need not be excessive, 
provided the means of moving water, even through the limited distance of the 
height of these small plants, be adequate. But examination of cross and longi- 
tudinal sections of the stem discloses conducting tissues in vascular bundles 
too few and too small to meet the need. Furthermore, the wood (xylem) of 
the vascular bundles consists of ducts (vessels) of relatively large size and 
thin Avails, and comparatively few and thin-walled tracheids. These plants 
are mechanically weak, but they successfully carry their own weight and the 
increasing load of seeds and fruit, breaking only when unusual force is ap- 
plied. The supporting tissues of Miner's lettuce are sufficient to keep it erect : 
but when the strain of drying air and soil is put upon the conducting tissues, 
they are unequal to the need of moving water rapidly from the roots which 
take it in to the leaves which lose it. The composition of the vascular bundles 
is diversified and interesting, but they are inadequate and the plants die. 

If one turn to other native herbaceous plants, the "tar weeds" (Hemizonia 
spf), one finds much longer lived annuals which start from seed at about the 
same time as Montia but which persist throughout the dry season till the rains 
come again. The plants form more or less perfect rosettes of largish rather 
fleshy leaves quite close to the ground. These are the food-manufacturing 



218 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

org-ans wliieli are active for a long time and which lose water comparatively 
slowly because of their position close to the gronnd, their hairy covering-, their 
water-proofing by cell-walls and secretions. The stalks which bear the flowers 
from early summer till late fall are slender, dry, woody, and their leaves ai-e 
small and narrow. The surface of stalks and leaves is waterproofed. The struc- 
ture is compact, as one would expect in a plant of such xerophytic habit, with 
adequate mechanical support. The vascular bundles are composed of a strik- 
ingly large proportion of thick-walled tracheids, with few and small ducts. 
The limited area of the surface of the flowering stalks and the composition of 
their vascular bundles insures a sufficient water transport to offset the water 
loss into the dry air. 

If one compare these two annuals with a plant which has a perennial under- 
ground root, namely Echinocystis fahacea Naud., common manroot, one sees 
long, slender stems which climb by means of branched tendrils very sensitive 
to contact. The leaves are large and thin, smooth or only sparsely spiny, not 
thoroughly waterproofed nor structurally provided with water-holding ma- 
terials. The stems are mechanical^ weak, except that their tensile strength is 
considerable, especially' when dry : but their vascular bundles have long been 
the admiration of the plant anatomist because of the arrangement and the 
strikingly large size of the duets and sieve-tubes. Tracheids make up only a 
small part of the wood. In the underground roots which, in older plants, may 
attain the size of a half barrel, or even larger, there is a great store of water ; 
but in spite of the spectacular vascular anatomy, the leaves and stem dry and 
die very soon after the onset" of the dry season. Furthermore, the leng-th to 
which the leafy stems will grow seems to be directly related to the humidity, 
as well as the temperature, of the air — the higher the humidit}' during the 
period in which air temperature makes growth possible, the longer the stems 
become. In "wet springs" when, because of occasional rains on thoroughly 
watered ground, air as well as surface soil remain damp, the stems may attain 
surprising lengths (30 feet) as they climb over fences, bushes, and grass. On 
the other hand, the same underground roots give rise to decidedly shorter 
stems in the "dry springs," when rains are infrequent or do not occur at all. 

One sees the same thing in the common garden sweet pea which, under 
various conditions of soil and air temperature, moisture, etc., produces slender 
stems of very different lengths from seeds of the same variety. Assuming the 
same treatment otherwise of soil and water, one's sweet pea vines will grow 
tall or remain shorter, they will continue to grow in length or they will stop, 
according to the need of carrying water in quantity and at speed through the 
stems from roots to leaves. Where air dryness makes transpiration from the 
leaves too rapid for the vascular bundles of the stems to offset, no matter how 
scrupulously the bed may be ground-watered, the stems promptl}^ cease to 
elongate. The water-carrying capacity of the stems, the size and composition 
of the vascular bundles, seem to determine the height of the plants. 

In all of tlie plants tluis cited, as well as in all the rest, tlie anatomical struc- 



Vol. XXV] PEIRCE: WATER AND PLANT ANATOMY 219 

ture of the stems may be considered to be hereditary ; but while the plants 
may vary according to circumstances, the limits of variation, as well as their 
behavior otherwise, seem to be set by hereditary characters, and we find the 
sweet pea failing to grow indefinitely in height in spite of the mechanical 
support of frame or trellis. The reason seems to be that the water-carrying 
capacity of its vascular system is limited. 

Aesculus calif ornica (Spach) Nutt., the buckeye, or horse chestnut, is a 
small, spreading tree which puts forth its light green leaves, and loses them, the 
earliest of our native perennials. Its leaves are large and thin ; one might call 
them delicate, but their skeleton is strong enough to carry a very considerable 
expanse. They fall apparently in accordance with the amount of water avail- 
able in the soil, those buckeye trees becoming bare earliest which are rooted in 
dryest or thinnest soil, and those retaining their leaves longest are rooted 
where more water is available. However, this is not necessarily the case ; young 
buckeyes from one to five or more years old, grown in well-watered gardens, 
and even grown in water cultures,^ in which there is a constantly adequate 
supph" of water as w^ell as mineral matters and air, lose their leaves before 
the cold of winter would make them fall. The reason for this behavior is to be 
found, I believe, in the anatomy of the wood. The xylem consists of strikingly 
large, relatively thin-walled ducts (vessels) with fewer tracheids or other thick- 
v.alled elements than might be expected. While the capacity to carry large 
volumes of water, if the water is abundantly supplied from the roots, is large, 
the size of the ducts and the thinness of their walls permit holding only small 
amounts of water. Hence the large leaves are insufficiently supplied wdth 
water as the dry season continues, and they gradually dry and fall. 

If one considers now the wood anatomy, the behavior of the foliage, and 
some of the other traits of two native oaks, Quercus loiata Nee, the Valley, or 
White Oak, and Q. agrifolia Nee, the Coast Live Oak, we shall be led further. 
Q. lohata puts out its summer canopy of leaves well after the coolness of mid- 
winter is past but with the soil water undiminished ; and it retains its foliage 
throughout the dry season, losing the leaves apparently when soil dryness 
may add to the difficulty of maintaining its foliage through cooling nights, 
and days in which photosynthesis may lag. The Live Oak, Q. agrifolia, on the 
other hand, while also losing its leaves once a year, puts out its new crop when 
the days have already become warm, and at least the surface soil has begun 
to dry. Then the leaves of the preceding year drop off. The leaves of Live Oak 
are ordinarily an inch or two long, compact in form, rather thick and leathery, 
smooth and shiny above, hairy underneath, and weU waterproofed. The leaves 

^ For six years I have maintained cultures of buckeye seedlings in one-quart milk bottles 
half filled with Knopp's Solution. These seedlings, one year old and older, up to six years, 
are bare from late August or early September tUl February or March in spite of the nutrient 
solution : and they put out new leaves in the spring at about the same time as similar seed- 
lings outside, rooted normally in the soil. The swelling of winter buds and the putting forth 
of leaves were not instigated in my cultures by fresh solution : nor do I see that there is any 
other general cause operating on the seedlings growing wild out of doors, as well as my cul- 
tures in bottles, than the rise in temperature which is one factor in the phenomena of spring. 



220 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

of White Oak are much larger, lobed, thin in proportion to their area, and 
not especially waterproofed. If one compare the woods of these two species 
of oaks one finds tlie xylem of AVhite Oak containinia: ducts of fairly large size, 
whereas that of Live Oak is made up predominantly of tracheids. The wood 
of the latter carries enough water from the roots, even in dry soil at the end . 
of the dry season, to keep the leaves from excessive drying, and from falling 
before the new crop of leaves emerges in the spring. 

The wood of the conifers, as is well known, is composed mainly of tracheids 
which, in proportion to their diameters, have thick walls. In these thick walls 
much water is tenaciously held. The majority of conifers are also evergreen, 
retaining their leaves for two or more years. Those which are not evergreen 
live where there is ample moisture in soil and air. The evergreen conifers are 
distributed from well-watered to desert areas. Their monotonous wood anatomy 
enables them to meet this great range of circumstances. The vascular systems 
of the conifers, all built upon the same general plan, are equal to the great 
variety of conditions under which different species and individuals live. But 
the vascular systems are associated with leaves of limited area, limited water 
loss, and even limited number. The vascular systems can carry enough water 
to supply the leaves at all seasons. Only in the severest of dry seasons, in years 
of light winter snowfall and of rainless summers, does one see signs of suffer- 
ing from lack of water. In such seasons the plants absorb what water they can 
from dry soil ; they lose what they must from leaves restricted in area and in 
number — many needles fall during such summers — ; they make limited 
growth in length and may even die back extensively. In spite of the superi- 
ority of their vascular systems, one finds the desert pines as scattered indi- 
viduals, not as close stands of forest or even chaparral. 

The amount of precipitation and the anatomy of the plants living in a given 
area are two of the factors which determine the character of the vegetation, 
limiting a natural area to grassland, chaparral, or forest, and correspond- 
ingly limiting what man can do in the area. Attempts to establish forest, with- 
out irrigation and other care, in regions of limited rainfall and naturally 
supporting only grass or even chaparral, seem unlikely to succeed because 
the water-carrying capacity of the wood is unequal to the need of moving 
water fast enough from the roots in dry soil to leaves in dry air at any but 
limited heights above ground. The height to which a plant can grow is the 
height to which its vascular system can carrj^ water. This is determined by its 
wood anatomy — which is inherited — and by its environment. The relation of 
the anatomy to water is the dominant factor, in our environment with its long 
dry seasons, in the complex which is the living thing. 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
\'ol. XXV, No. 8, pp. 221-234, pis. 23-24 Jiii^e 1, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 8 

A PLEISTOCENE FLORA FROM THE McKITTRlCl^ 
ASPHALT DEPOSITS OF CALIFORNIA 

BY 

HERBERT L. MASON 

Associate Professor of Botany and Curator of the Herbarium 
University of California 

THE KNOWN Pleistocene floras of California are largely coastal in their posi- 
tion and contain a group of species as fossils that are essentially maritime 
in their occurrence today. Of these by far the richest as to species content are 
the Tomales flora (1) in Marin County and the Carpinteria flora (2) in Santa 
Barbara County. Both of these contain a Monterey type forest dominated by 
pines and containing cypresses and live oaks as well as an understory of shrubs 
that is highly diversified as to genera and species. The asphalt deposit at 
Rancho La Brea (3) in Los Angeles County, although near the coast, contains 
a flora that is essentially interior in its nature and is best represented today 
by the open woodland vegetation on the slopes of the Tehachapi Range. The 
occurrence in this deposit of a few specimens of maritime species, however, 
suggests that at least a part of the drainage basin contributing to this deposit 
Avas populated by a maritime flora. It seems most likely that a mountain range 
such as the Santa Monica Moitntains lay between it and the sea and that the 
coastal slope of these mountains was covered by a maritime flora which ex- 
tended over the top and a short distance down the interior side much as does 
the coastal pine forest today on Cedrus Island off the Lower California coast. 
Only a very few strictly interior floras of Pleistocene age in California are as 
yet known and these contain very few species of plants. We therefore know 
very little as to the nature of the interior California floras during Pleistocene 
time and hence welcome any information, however meager, that will help us 
to piece together a picture of the vegetation of the interior for this period of 
the earth's history. 

[221] 




222 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

In the asphalt deposits at McKittriek in western Kern County, California, 
such a fossil flora occurs. The jMcKittrick deposit is located at the southwest- 
ern edg-e of a small oil town of the same name in the southwestern San Joaquin 
Valle}^ just east of the base of the Temblor Range. The elevation of McKittriek 
is 300 feet while the Temblor Range rises to about 4000 feet. The geology of 
the area has been reported by Arnold and Johnson (4) and will not be further 
treated here. The rich fossil faunas have been described and discussed by 
Merriam and Stock (5, 6) for the mammals and by Miller (7, 8) for the birds. 
Summaries of these latter papers are presented by Hay (9). Arguments as 
to the precise position of these beds in the Pleistocene have been presented 
and discussed at length by these authors. The plant species that are at all 
critical in this respect all go back to the Pliocene both north and south of this 
area and are still living both north and south of this area and hence can throw 
very little light, if any, upon this problem. The Pleistocene status of the flora 
is inferred from the associated fauna which is largely made up of now extinct 
species. 

The flora, however, does have a contribution to make as to the general eco- 
logical setting and can serve to a large extent in forming the basis for an 
interpretation of the habitat and the local climate for the portion of the Pleis- 
tocene represented. It will fill out the ecological picture presented in part by 
the mammalogist and in part by the ornithologist. 

Merriam and Stock (5, 6) report the occurrence of horses, bison, camels, 
antelope, bear, lion, tiger, wolf, ground sloth and mastodon, all of which are 
now extinct. These are largely plains or savanna types of animals. Miller 
reports a list of thirty-five birds of which water birds predominate by two to 
one. Only 10 per cent of the birds are now extinct. 

To date six identifiable plant species have been found. These consist of two 
gymnosperms and four angiosperms. Although the flora is meager these plants 
are sufficiently critical as to their climatic tolerance to serve as indicators of 
the nature of the flora and of the climate that prevailed in the McKittriek 
region during Pleistocene time. It may be that some future excavations will 
yield a flora comparable in quantity as well as in quality to other asphalt 
deposits in California but for the present we must be content with Avhat 
there is. 

The plants are represented in the deposit by fragments of poorly preserved 
fruiting structures and seeds as well as scraps of partially disintegrated 
twigs. These are imbedded in sand that has been impregnated with asphalt. 
The advanced state of deterioration of so much of the plant material suggests 
that the impregnation of the asphalt occurred after the plants were deposited 
and after preliminary decomposition had set in. There is not the evidence 
of a pool of asphalt such as is suggested by the conditions at Ranclio La Brea. 
As compared with equal volume of matrix from other asphalt deposits the 
plant material in the McKittriek is rare. Most of the species are represented 
by single specimens. In onlj^ one case, the Atriplex, is there sufficient material 



Vol. XXV] MASOX: PLEISTOCENE McKITTEICK FLOSA 223 

to give any notion as to the range of morphological variation of the structure. 
Fortunately, however, most of the material is of such a nature as to be readily 
identifiable and the following species may be recorded : 

Pinus monophylla Torr. 
Juniperus calif ornica Carr. 
Atriplex lentiformis (Torr.) Wats. 
Arctostaphylos glauca Lindl. 
Arctostaphylos ptmgens H.B.K. 
Echinocystis fahacea Naud. 

All of the above species found in the McKittrick flora are still living in the 
California flora today. The occurrence of all of them in a single Pleistocene 
flora presents a harmonious ecological picture that is not unlike an assemblage 
in the modern flora of the eastern end of the San Rafael Mountains in north- 
eastern Santa Barbara County, on the slopes of Santa Barbara Canyon a little 
above its entrance upon the Cuyama Valley. (United States Geological Sur- 
vey, Santa Ynez Quadrangle, R 25 W, T 9 N, in the Santa Barbara National 
Forest.) 

The San Rafael Mountains in this vicinity range from altitude^s slightly in 
excess of 600 feet to altitudes near 7000 feet. The great differences in altitude 
and the roughness of the terrain are reflected in great local diversity of the 
flora. A profile section of the San Rafael Range in this region in the direction 
of the storm paths will help to explain why a region so close to the coast 
supports such an arid type of vegetation. There are three prominent ridges 
separated by the canyons of Manzana Creek and the Sisquoc River. The west- 
ernmost of these ridges reaches an altitude of 4000 feet at Zaca Peak directly 
above the 1500-foot Santa Ynez Valley. The canyon of Manzana Creek drops 
precipitously to 1700 feet. The ridge between this canyon and the canyon of 
the Sisquoc River rises to a height of 3500 feet and is clothed with a very 
arid type of vegetation. The canyon of the Sisquoc drops to 2300 feet and the 
third and highest ridge culminates in Pine Mountain at 6800 feet with a 
northward spur of about 5800 feet forming the west wall of Santa Barbara 
Canyon. These ridges serve to force the moisture-laden winds from the ocean 
upward and causes them to deposit their load on the westernmost ridge and 
the upper part of the easternmost high ridge. This condition is reflected in 
the vegetation of these ridges. The western slope of the first ridge is clothed 
with a savanna type of cover except where the topography causes rapid run- 
off. Here the vegetation is chaparral. The top of this ridge is clothed with a 
forest of big cone spruce (Pseudotsuga macrocarpa), Coulter pine {Pinus 
Coulteri) and western yellow pine {Pinus ponderosa), which runs down the 
canyons and gullies on the east side. The second ridge, as was pointed out 
above, is clothed with a very sparse, arid type of chaparral. The third ridge 
is arid below the 4000-foot level but above is clothed with a very rich conif- 
erous forest of big cone spruce. Coulter pine, western yellow pine and sugar 
pine {Pinus Lamheriiana). It is the lower slopes of the east side of this ridge 



224 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

that contain the flora represented in the ^k-Kittrick asphalt deposit. Here is 
an arid type of flora of scattered stands of pinyon pine (Pinns monophylla) 
and California juniper {Juniperus calif ornica) and patches of manzanitas 
{Arctostaplnjlos glauca, A. pungens and A. Parryana). The other species 
represented in the fossil flora are abundant on the lower slopes and in the 
Pleistocene were probably much closer to the site of deposition than were the 
larger shrubs and trees. 

Precisely where the particular plants that found their way to preservation 
in the McKittrick asphalt grew during Pleistocene time, is difficult to deter- 
mine. It seems probable, however, that they were a part of the flora that occu- 
pied the Pleistocene Temblor Range and were borne down by the streams to 
the ponds and lakes that ultimately became silted up and infiltrated with 
asphalt. The pine, the juniper and the manzanitas, as well as the Echinocystis, 
probably occupied the higher slopes, while the Atriplexocc\i]}ied alkaline flats 
near the ponds and marshes. This occurrence would explain the abundance 
of Atriplex and the relative scarcity of the other species in the deposit, since 
these latter would have to be transported by a none too rich stream flow. 

Climatically the flora suggests an area of low rainfall but perhaps more 
than is received by the McKittrick region at the present time. This might lead 
some to argue that the flora indicates at least glacial time ; however, in view 
of tlie close proximity of a similar modern assemblage of species it is more 
logical to assume that some minor changes in topography to the west would 
have accomplished the same result during the Pleistocene at McKittrick. If 
the Temblor Range were higher by 1000 feet, it might support such a flora 
around its summit. Evidence as to glacial or interglacial time is negative so 
far as the McKittrick flora is concerned. 

Ecologically the flora is entirely consistent with the assemblage of mammals 
and of the birds. It was sufficiently open to permit grazing and browsing and, 
although the presence of Atriplex does not necessarily mean ponds and lakes, 
it frequently occurs near such sites today. 

Since all of the plants represented in the McKittrick flora are modern 
species the nomenclature here used is that of the modern plants. In one of 
the plants the nomenclature of the modern group is not as yet stabilized, 
owing to incomplete knowledge and hence uncertain taxonomic concepts. In 
this instance it is felt that it is not within the province of this paper to solve 
these problems, nor could they be solved on the basis of the material here 
presented. The nomenclature for this species therefore follows current usage. 
The problem will be raised in connection with the species in question. 

The specimens here cited are all deposited in the Palaeobotanical Collection 
of the University of California at Berkeley. 



VOL.XXV] MASON: PLEISTOCENE McKITTEICK FLOBA 223 

Pinus monophylla Torr. 

(Plate23, fig. 3) 
Pinus monophylla Torr. in rremont, Second Eeport, 319, 1845. 

Pinus monophylla is represented by a cone, of relatively large size for this 
species, that is poorly preserved. The structure is practically reduced to vas- 
cular tissue but enough of the surface features of a few of the scales is pre- 
served to show the essential nature of the apophysis and of the umbo as 
well as to clearly indicate the size and shape of the seed cavity. The cone is 
about 8 em. long and about 7 cm. wide. The scales are terminated by a some- 
what recurved, quadrate apophysis bearing a dorsal umbo. The seed cavity is 
ovoid, about 12 mm. long and 8 mm. wide, indicating a relatively large seed. 

Pi7ius monophylla ranges primarily to the east of the Sierra Nevada. A 
few stations have been recorded for its occurrence on the west slopes of the 
Sierra. It occurs also in the Tehachapi and the Mount Pinos region, and 
extends northward in the coast ranges to the Cuyama Valley in the San Rafael 
Mountains, and on the watershed of the headwaters of the Santa Ynez River. 

University of California Palaeobotanical Collections, Plesiotype no. 2810. 

Juniperus californica Carr. 

(Plate23, fig. 1) 

Juniperus californica Carr. Eev. Hort. 352, 1854. 

Juniperus californica Carr. var. breaensis Frost, Univ. Calif. Publ. Bot. 14:77, 1927. 

Juniperus californica Carr. is represented in the fossil flora by a seed 8 mm. 
long by 5 mm. wide that in no way differs from seed of the modern species as 
it occurs in the Tehachapi Mountains today. The seed coat is of medium thick- 
ness, and agrees with other Pleistocene material collected from the asphalt 
deposits at Rancho La Brea and described by Frost as distinct from the 
modern plants. 

The modern J. californica Carr. is an aggregate species that includes with 
varietal rank the desert and Great Basin form J. utahensis (Engelm.) Lem- 
mon. These two forms intergrade in the region of the Tehachapi Mountains 
so as to make their separation as distinct species impossible. They have been 
separated on the basis of the glandular nature of the foliage of typical /. 
californica. Another important difference is the relative thickness of the seed 
coat in the extremes of the two forms. Typical /. calif orriica has a thin seed 
coat while typical /. californica var. utahensis has a very thick seed coat. Both 
the characters of the seed coat and of the glandulosity intergrade in material 
collected by the writer from the Tehachapi Mountains and from the Sierra 
Liebre in California (see Mason 3737, Herbarium of the University of Cali- 
fornia). The fossil material from the McKittrick asphalt is of the intermediate 
type represented by this material. Because of its intermediate character it 
is impossible to designate whether it belongs to the species or to the variety. 

In the fossil record J. californica has been reported from tlie asphalt depos- 



0Qiq^ 



.'^xrJ^^*^ 




226 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th ser. 

its at Carpinteria and at Rancho La Brea. In the latter flora, however, the 
material was described as new under the name J. calif ornica var. hreaensis 
Frost. J. californica ranges from Lower California through the western 
]\Iojaye Desert to the Tehachapi Mountains and northward in discontinuous 
patches in the inner coast ranges to Shasta County and sparingly on serpen- 
tine soil in the Sierra Nevada footliills to Butte County. It is of interest to 
the present paper to record it in abundance in the inner south coast ranges 
that border the Cuyama Valley. 

L'niversity of California Palaeobotanical Collections, Plesiotype no. 2811, 

Atriplex lentif onnis (Torr.) Wats. 

(Plate 24, figs. 1, 2, 3) 
Atriplex lentif ormis (Torr.) Wats. Proc. Am. Acad. 9:118, 1874. 

The species is represented by a large number of fruiting bracts often enclos- 
ing fragments of the seed coats. They are flattened and wedge-shaped. Some | 
of them possess a stipe while others appear to have been sessile on the branch. 
They average 4 mm. in length and 5 mm. in M'idth. The bracts are united below] 
over the seed and are apparently free above. The upper margins are somewhat ] 
broken and it is difficult to reconstruct the character of these organs. There l 
is evidence to indicate a fringed, denticulate margin. The surface shows aj 
coarsely reticulate venation and is entirely devoid of any facial processes. 

It seems evident from a careful comparison of the specimens that but a 
single species of the genus is represented. The great range of variation exhib- 
ited by many of the modern species shows forms that are not unlike the fossils. 
How^ever, each species has a characteristic kind of fruiting structure and 
its variation centers around this type. Of the modern species now living in j 
California, A. lentif ormis seems closest both on morphological and ecological 
grounds. The area Avherein this flora is best represented today is rich in 
species of Airiplex and any or all of them might well have been living in the 
McKittrick locality during the Pleistocene. 

The genus Atriplex has been recorded from the Pleistocene Tomales flora] 
of central California where fossil material of the coastal species of A. hastata^ 
has been collected. 

University of California Palaeobotanical Collections, Plesiotype no. 2812. 

Arctostaphylos glauca Lindl. 

(Plate24, figs. 4, 5, 6) 
Arctostaphylos glauca Lindl. Bot. Reg. sub t. 1791, 1835. 

This large-fruited species of manzanita is represented in the fossil flora by 
frag-ments of fruits one of which is 12 mm. in diameter, with its carpels closely 
coalesced so as to present a very smooth surface. The material is identical Avith 
similar structures on the modern A. glauca and is clearly referable to that 
species. 



VOL.XXV] MASON: PLEISTOCENE McKITTRICE FLORA 227 

The modern plant is a characteristic species of the Upper Sonoran life zone 
in the blue-oak-dig:ger pine belt, and ranges from central California south- 
ward to southern California. It is frequent in the Tehachapi Mountains and 
the inner south coast ranges in association with juniper. Its occurrence in the 
fossil flora suggests conditions similar to those of these mountains at an 
altitude of about 3000 feet. 

Arctostaphylos glauca is knowai in the Pleistocene from both the Carpin- 
teria and the Rancho La Brea asphalt deposits. It is an indicator of arid 
interior conditions (Chancy and Mason, 2) . 

University of California Palaeobotanical Collections, Plesiotype no. 2813. 

Arctostaphylos pungens H.B.K. 

Arctostaphylos piingens H.B.K. Nov. Gen. et Sp. 3 :278, 259, 1818. 

This species is represented in the fossil flora bj' a fruit 4 mm. long and 6 
mm. wide with coalesced carpels. The individual carpels show unequal devel- 
opment and present a strongly ribbed appearance to one side of the fruit. 
The specimen is not unlike many that are of frequent occurrence on modern 
plants of Arctostaphylos pungens of the inner south coast ranges. 

Arctostaphylos pungens is a species of the arid hills bordering on the upper 
part of the Sonoran and the lower part of the Transition zones from central 
California southward. Its precise range cannot at present be stated because 
it is obvious that what has been going under the name of A. pungens in current 
floras of the region is not the same as the plant originally described under 
that name from the vicinity of Mexico City. Furthermore, there is consider- 
able confusion about the taxonomy of the group locally and until this problem 
can be straightened out by a monographer of the genus as a whole the precise 
name must stand in abeyance. It is possible that A. montana Eastwood and 
A. Hookeri Don may be involved in relationship here to such an extent as to 
invoh'e the nomenclature of the fossils. At any rate, the specimen under con- 
sideration is of a type like the modern plants of this group now growing in 
the inner south coast ranges of California and referred to in modern floras 
as A. pungens. 

University of' California Palaeobotanical Collections, Plesiotype no. 2814. 

Echinocystis fabacea Naud. 

(Plate 24, figs. 8, 9) 
Echinocystis fahacea Naud. Ann. Sci. Nat. ser. IV, 12 : 154, 1859. 

Fragments of the seed coat of this species preserved in this deposit differ 
in no way from the modern species that is so widespread in the inner and 
outer coast ranges of California. Identification was based upon the histologi- 
cal nature of the seed coat, where there are tw^o very different layers of cells 
evident, an inner layer composed of narrow cylindrical cells and an outer 
layer about three times as thick composed of manv irregularly rounded cells,-— r-3r—-s-. 




228 CALIFORNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

The inner face of tlie seed coat is lined with a very soft spongy tissue that 
readily sloughs off. Echitiocystis fahacea ranges throughout the inner and 
outer coast ranges of central California, crossing the Tehachapi Mountains to 
the Mojave Desert, where it has been reported in tlie vicinity of Muroc. It is 
not common in the Sierra Nevada foothills, but is abundant in the inner south 
coast ranges. 

University of California Palaeobotanical Collections, Plesiotype no. 2815. 

LITERATURE CITED 

1. Mason, H. L. Carnegie Institution of Washington Publication no. 415 : 83-179. 1934. 

2. Chanet, R. W., and H. L. Mason. Carnegie Institution of Washington Publication no. 

415:47-79.1933. 

3. Frost, F. H. University of California Publications in Botany 14 : 73-98. 1927. 

4. Arnold, R., and H. R. Johnston. Bull. U. S. Geological Survey no. 406:1-225. 1910. 

5. Merriam, J. C, and C. Stock. Science, n. s. 54:566-567. 1921. 

0. . Carnegie Institution of Washington Publication no. 347:11, 38—44. 1925. 

7. Miller, L. II. Condor 24 : 122-125. 1922. 

8. . University of California Department of Geology, Bull. 15 : 307-326. 1925. 

9. Hay, O. P. Carnegie Institution of Washington Publication no. 322B: 196-199. 1927. 



EXPLANATION OF PLATES 



PLATE 23 

Gymnosperms in the McKittriek Flora 

Fig. 1. Juniperus calif ornica Carr., drawing of the fossil, x 6. Univ. Calif. 
Palaeobot. Coll., Plesiotype no. 2811. 

Fig. 2. Seed of a modern plant of Juniperus calif ornica Carr., x 1. 

Fig. 3. Cone of Pinus monophylla Torr. x 1. Univ. Calif. Palaeobot. Coll., 
Plesiotype no. 2810. 



[230] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 8 



[MASON] PLATE 23 






[231] 



PLATE 24 

Angiosperms in the McKittrick Flora 

Figs. 1, 2, 3. Atriplex lentiformis (Torr.) Wats. Fruiting bracts, x 10. Univ. 
Calif. Palaeobot. Coll., Plesiotype no. 2812. 

Figs. 4, 5, 6. Arciostaphylos glauca Lindl. Fragments of fruit. Figs. 4 and 
5, X 4 ; fig. 6, X 5. Univ. Calif. Palaeobot. Coll., Plesiotype no. 2813. 

Fig. 7. Arciostaphylos pungens H.B.K. Endocarp of fruit, x 8. Univ. Calif. 
Palaeobot. Coll., Plesiotype no. 2814. 

Fig. 8. Echinocystis fabacea Naud. Cross section of the seed coat of a mod- 
ern specimen showing the two layers of the seed coat, x 15. 

Fig. 9. Echinocystis fabacea Naud. Cross section of the seed coat of the fos- 
sil, X 15. Univ. Calif. Palaeobot. Coll., Plesiotype no. 2815. 



[ 232 ] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES, VOL XXV. NO. 8 



[MASON] PLATE 24 










[ 233 ] 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
\'ol. XX\^ No. 9, pp. 235-244, 6 text figures Jiine 1 . 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 9 

THE PERENNIAL SOUTHWESTERN DATUR 
AND THE VALIDITY OF MATTHEW'S 
HYPOTHESIS IN PLANT GEOGRAPHY 

BY 

JOSEPH EWAN 

Instructor in Biology, Curator of Botany, University of Colorado 

EVIDENCES OF plivlogeiiy among major plant groups rest upon two lines of 
observational research, viz., comparative morphology and paleobotany, 
and upon two methods of experimental research, viz., serum diagnosis and 
cytogenetics. Among species within a single genus evidences of phylogeny 
rest again upon comparative morphology in the broadest sense, along with 
breeding experiments and cytogenetics, with either supporting evidence or 
the lack of it from the fields of ecology and plant geography. Interpretation 
of morphology is thrown into sharper perspective by subjecting the data to 
Haeckel's Biogenetic Law, Dollo's Law of Irreversibility, Darwin's evolu- 
tionary theory, Liebig's Law of the Minimum as applied to distribution prob- 
lems. Good's Theory of specific tolerance, Jordan's Law of Geminate Species, 
Eames' concept of "phyletic slides" within plant groups, and what I have 
chosen to call Matthew's Hypothesis of Peripheral Populations. The subject 
of the present paper is a consideration of Matthew's Hypothesis as it may 
prove valid in interpreting distribution phenomena in Datura metdoides DC. 
In his now classic paper entitled "Climate and Evolution" (1915) the Amer- 
ican vertebrate paleontologist, W. D. Matthew, presented a working hypothe- 
sis for the evolution of the higher vertebrates. He devoted three paragraphs 
in the beginning to his "Principles of Dispersal," with which we are chiefly 
concerned. His evolutionary theory embraced the concept that the evolution 
of land life has responded in adaptation to recurrent periods of aridity 

[ 23.5 ] 




236 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

through geologic time. In short, Matthew demonstrated that the major ad- 
vances of vertebrate evolution have coincided with shifts in megaclimates, 
especially those shifts toward uicreasing aridity. It is a significant corollary 
of Matthew's Hypothesis, in my opinion, that the angiosperms of arid and 
semi-arid regions exhibit conspicuous evolutionary advance in their mor- 
phology over their relatives of mesic regions. 

Matthew's Hypothesis 

Matthew's Hj'pothesis holds that, 

at any one time in the evolution of a taxonomic gro^ip the most advanced stages should 
be nearest the center of dispersal, the most conservative stages farthest from it. 

Or, to use the words of another vertebrate paleontologist, Prof. R. S. Lull, it 
may be stated that, 

"the most ancient members of a group are not to be found at the old center of evolution, 

but rather at the periphery of their migratory area." (Lull in George A. Baitsell, ed., 

Evolution of Earth and Man, pp. 118-119. 1929.) 

In his original paper Matthew points out that the evolution of a race "should 
'be at first most progressive at its point of original dispersal, and it will con- 
tinue this progress at that point in response to whatever stimulus originally 
caused it and spread out in successive waves of migration, each wave a stage 
higher than the previous one" (1915, 180). Assuming that a species is the 
product, in part, of its environment, and that environments are subject to 
change, "it is the environment itself, biotie as well as physical, that migrates, 
and primitive species are those which have followed it, while those which 
remained have had to adapt themselves to a new environment and become 
altered thereby." However, it should be remembered that probably "it is 
never the case that the environment of the marginal species is an absolute 
replica of the older environment of the race" (1915, 180-181). 

The American plant ecologist Stanley Cain expressed the concept in 1940 
as follows : "in wide-ranging species populations mere distance constitutes 
a type of isolation and marginal or other portions of a population are fre- 
quently recognizable . . . without the development of any conspicuous dis- 
continuity" (19-40, 214). Cain points out, furthermore, that these marginal 
populations are genetically more homozygous. In the instance of Daiuva 
meieloides, the native solanaceous perennial known by the Indian name 
"tolguacha," this homozygosity may be demonstrated by the occurrence of 
distinctive leaf types of representative individuals from geographically mar- 
ginal populations, as recorded in some of the larger herbaria. 

Materials and Procedure 

This study of Datura meieloides, it should be pointed out at the outset, is 
based Avholly upon herbarium specimens and is subject to confirmation from 
an even more statistical approach, especially in the field over the extensive 
geographic range of the species, in the manner of Fassett's "mass collection" 



Vol. XXV] 



EWAN: DATUEA 



237 



studies (1941). It may be of interest to record how this problem arose. De- 
Candolle's original description of Datura meteloides rests, not upon the usual 
preserved herbarium "type" specimen but upon a Sesse and Mocirio drawing, 
rather unfortunate for having been hastily copied from the original. The 
plant from which the drawing was made was collected at an unrecorded local- 




Fig. 1. Maximum natural range (generalized) of Datura meteloides DC. 



ity in, probably, central Mexico by Mocifio. By comparing this drawing with 
modern herbarium collections as to close similarity of leaf blade outline an 
attempt was made to fix, ixi a general way at least, the possible geographic 
source, that is, a putative "type locality," of the material which served as the 
basis of what must be considered, in lieu of an actual preserved plant, the type 
of the species. The taxonomic history of this plant, and certain problems in 
the systematics of it, are treated elsewhere by the present author (1944) . 



238 



CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 



This paper, then, arose from an examination of nearly 300 collections, 
primarily from two large herbaria, the U. S. National Herbarium, and the 
Rocky Mountain Herbarium of the University of Wyoming, with subsequent 
examinations of materials in the herbaria of the Academy of Natural Sci- 
ences at Philadelphia and the Field Museum. To the curators of these collec- 
tions I am grateful for the privilege of studying the materials in their charge. 
The collections were sorted according to closeness of fit in leaf blade config- 
uration. Four chief repetitions in leaf-form were encountered. These char- 
acteristic infraspecific variations in leaf -form — including equilateral and 
inequilateral, serrate and subentire forms — were then plotted on outline maps 
according to the place of collection as recorded on the herbarium labels of 
each. ^lature leaves were chosen from each sheet and tracings were made of 
representative collections. Almost invariably the collections were of flowering 
specimens, and the leaves, therefore, associated with flowering shoots. It was 
found that those collections showing closely comparable leaf -forms came, for 
the most part, from geographically limital localities. These data are given 
below. The total range of the species is shown in a generalized way by the 
accompanying map (fig. 1). 



Data from Datura meteloides 
Four distinctive groups showing close repetition of leaf-form are listed 
below. From each group representative collections are cited, chosen from 
widely separated and geographically peripheral localities. Pen line tracings 
and map positions are given for each collection. 



Leaf-form 
inequilateral 
ovate and 
subentire 



Leaf-form 
equilateral 
ovate and 
serrate 



Leaf -form 
equilateral 
ovate and 
subentire 

Leaf -form 
equilateral 
deltoid and 
serrate 



'' Locality 
1. San Bernardino, Calif 



Group One (fig. 2) 

Collector 



2. Santa Barbara, Calif. 

3. St. George, Utah 

4. Valencia Beach, L. Calif. 



G. E. Vasey 443 
W. H. Brewer 310 
M. E. Jones 6093 
Wiggins and 
Gillespie 3948 

Group Two (fig. 3) 

Locality Collector 

5. Holbrook, Ariz. Osterhout 6767 

6. Tarrant Co., Texas A. Ruth 583 

7. Fuerte, Sinaloa, Mex. Eose, Standley 

and Eussell 13452 

Group Three (fig. 4) 
Collector 



< 



f Locality 

8. Santa Ysabel, Calif. 

9. Colonia Diaz, Chih., Mex. 
in. Buckskin Mts., Ariz. 



Henshaw 68 

E. W. Nelson 6430 

Jones 6063n 



Group Four (fig. 5) 
Locality Collector 

11. Montezuma Co., Colo. Crandall in 1895 

12. Pasadena, Calif. G. B. Grant 270 

13. Mazatlan, Sinaloa, Mex. Ortega 5654 



Herb, and sheet no. 
USNH 156906 
USNH 322527 
EM 14182 
USNH 1491044 



Herb, and sheet no. 
EM 169488 
USNH 1523537 
USNH 636275 



Herb, and sheet no. 
USNH 238095 
USNH 360146 
USNH 260638 

Herb, and sheet no. 
USNH 963190 
EM 54824 
USNH 1208683 



Vol. XXV] 



EWAN: DATURA 



239 




Fig. 2. Group One of Datura meteloides leaf -forms. 




Fig. 3. Group Two of Datura meteloides leaf-forms. 



240 



CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

L 




Fig. 4. Group Three of Datura meteloides leaf -forms. 




Fig. 5. Group Four of DaUira meteloides leaf -forms. 



Vol. XXV] 



EWAN: DATURA 



241 



Collections from the more central portion of the range of Datura meteloides 
show a range of leaf-forms well represented by the five tracings made from 
Charles Wright 526 from Turkey Creek, Texas, which is the lectotype of 
Datura Wrightii Kegel (USNH 60043, cf. fig. 6, tracings 14-18 inc. all from 
same coll.). Collections closely comparable to Wright 526 in the range of 
variation of leaf -forms have been made at the stations indicated by the circles 




Fig. 6. Center of distribution of Daiura vieteloides as determined by conformity in 
leaf -form, with five leaves (nos. 14-18) from single coll. {WrigM 526) from locality 
at "14." 

in fig. 6. This, then, represents the biologic center of the species. Doubtless 
the species will be found to be present, in suitable river bottom and alluvial 
habitats, in northern Mexico as well, where the map suggests its absence. 

Discussion 
The center of distribution for Datura meteloides may be recognized as the 
Rio Grande catchment basin of New Mexico westward to the upper Gila River 
basin of Arizona. This conclusion is based upon a noticeable homogeneity in 
the populations as demonstrated by the limited range of variation of their 
leaf -forms. The peripheral populations are considered, therefore, to represent 
fragmentary remnants of the original species colony persistent today as a 
fringe about the now altered population mass comprising the present "center 
of distribution." 



242 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Gleason noted the agreement of marginal members of a group about a center 
of distribution in his studj^ of the genus Vernonia. He wrote, "that those spe- 
cies most nearly alike morphologically . . . ai-e always distributed about the 
same center" (1906, 152). C. H. MuUer found .Matthew's Hypothesis to obtain 
in certain species of Mexican Quercus (oral communication ; cf. also Am. Midi. 
Nat. 18: 844, 1937). In my studies of the Calif ornian flora instances which 
apparently corroborate Matthew's Hypothesis have been noticed in Monar- 
della lanceolata Gray, where small capitulae of closely comparable form and 
size are to be noticed in plants of Yosemite Valley on the north and San Diego 
County, California, on the south. In the mariposa called "lantern-of-the- 
fairies," Calochortus alhus DougL, closely parallel small-flowered forms may 
be recognized at almost the same geographically distant localities as in Monar- 
della,^ each an extreme limital population of the species range. Among the 
North American Delphiniums, infraspecific phases have been detected in the 
Calif ornian Delphinium Parry) Gray where small-flowered populations occur, 
again, at limital positions over the total range of the species. In Delphinium 
patens Greenei (Eastw.) Ewan, a very restricted subspecies of California, 
plants with identical glandular pedicels occur on Marysville Buttes in the 
northern Great Valley on the north and from Fresno County upper foothills 
on the south, with non-glandular plants of Delphinium paiens patens occupy- 
ing the interveniug area. 

An apparent antithesis of Matthew's Hypothesis as valid for Datura mete- 
loides occurs in Tradescantia yvhere the peripheral populations have been 
demonstrated by Anderson and Woodson (1935) to be derivitive tetraploid 
infraspecific races, with the ancestral diploid races occupying the center of 
distribution. These infraspecific races are indistinguishable in their morphol- 
ogy. The oldest members of the species populations are central, then, and 
not, as in Datura meteloides, marginal to the mass of the species. In Trad- 
escantia the center of distribution significantly is geologically an old land 
mass within the total range of the species. In Datura meteloides the peripheral 
populations occupy extremely diverse regions which are not often cognate 
geologically. Furthermore, these peripheral populations are not morpho- 
logically homogeneous, at least as to their leaf -forms, but differ within them- 
selves from the much more uniform plants of the center of distribution. 
There are, in fact, four distinctive leaf -forms existing more or less side by 
side within this marginal population; at least in coastal Southern California 
three of the four leaf -forms occur together. 

In the European crucifer Biscutella laevigata L. three polyploid infra- 
specific races — a diploid, tetraploid and hexaploid race — have been shown 
to exist by Irene Manton (1937) . Here the archaic diploids are "xerothermal" 
interglacial relicts along the river valleys of central Europe from Germany 
to the Balkans, nol'th and east of the Alps. The tetraploid plants occupy the 

^ "As is frequently the case with Lnhiatae, the extreme forms geoprraphically are often 
most similar, at least superficially." (Epling, A Revision of Salvia: Subgenus Calosphace, 
191,1939). 



Vol. XXV] EWAN: DATVEA 243 

Swiss and Austrian Alps, swept over by glaciers, as a subsequent invading 
population. Obviously there is no direct relation to glaciation in the distribu- 
tion of Datura meteloides. 

There is, however, with this Datura a clear floristic affinitj^ to the Sierra 
Madrean element of Southwestern United States and Mexico. Axelrod's map 
(1939, 59, fig. 2B) should be studied in this regard. This map shows the 
"supposed distribution of the Sierra Madrean element as a dominant" at 
Lower Pliocene time. This extent agrees very closely with the extreme geo- 
graphic range enjoyed by Datura meteloides at the present time. Indeed, the 
peripheral populations whose leaves exhibit these four distinctive leaf -forms 
characterized in the present paper fall very nearly upon the margin of this 
mapping of the Sierra Madrean floristic element by Axelrod. The only notable 
exception is the northeastern arm in its range lying to the east of the Llano 
Estacado of Texas (cf. fig. 1). This arm falls, perhaps significantly, to the 
east or coastward of the Fall Line, passing northward on the Coastal Plain 
sediments. The absence of Datura meteloides in the region to the west, that 
is, on the Llano Estacado, is believed by Prof. E. L. Reed to be due essentially 
to the lack of proper alluvial or river-bottom habitats in that region (per- 
sonal communication, 22 Feb. 1939).' The occupation of the Coastal Plain, 
and hence, the significance of the Fall Line as marking the upper limit of 
the Coastal Plain or the "boundary between Triassic and older formations on 
the inland and Lower Cretaceous (Comanchean) and later formations coast- 
ward," has been discussed by Pennell very fully (1935, 572-579). 

It will be noted that the marginal phases of Datura meteloides today are 
usually topographically isolated by mountain ranges of varying height and 
extent or by arid desert expanses with their playas and bolsons, serving as 
topographic barriers. Topographic isolation, to a degree then, has maintained 
these distinctive leaf -forms at these limital stations. 

Summary 

The central populations of Datura meteloides, the "tolguacha," though vari- 
able, are uniform to a degree and show in general an absence of such leaf-forms 
as are exhibited by the geographically peripheral populations. There exist on 
the margin of its range today four leaf-forms ; these leaf -form types are not 
geographically segregated, however, but sometimes occur together in the same 
marginal locality. 

An explanation may be : the population of the original center of distribu- 
tion was highly heterozygous and consisted of several infraspecific races (this 
needs cytogenetic confirmation). Applying the concept of Matthew, these 
several races moved outward in irregular waves of varying intensity and 
direction and for varying time durations. This movement may have been part 
of the larger general spread of the Sierra Madrean floristic element over the 
Southwest. The end result is a series of partial-populations now scattered on 

- Prof. B. C. Tharp finds no coll. from the Llano Estacado in Univ. Texas Herb. 



244 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

the periphery of the species range. Each partial-population is reminiscent in 
its leaf architecture of its descent from a parent race which, as a race, has 
been lost in the center of the species mass by the subsequent swamping and 
intermingling of races and the genetic stabilization of the species. 

LITERATURE CITED 

Anderson, Edgar and Kobert E. "Woodson. Species of Tradescantia indigenous to the United 
States. Contr. Arnold Arboretum 9:1-132. 12 pis. 1935. 

Axelrod, Daniel I. Miocene flora from the western border of the Mohave Desert. Carn. Inst. 
Wash. Publ. 516:1-129. 12 pis. 1939. 

Cain, Stanley. Some observations on the concept of species senescence. Ecol. 21:213-215. 
1940. 

Ewan, Joseph. Taxonomic history of Tolguacha, Datura meteloides DC. : a problem in the 
application of the Type Basis Concept. Khodora vol. 46. 1944. (In press.) 

Fassett, Norman. Mass collections : Bubus odoratus and B. parviflorus. Ann. Mo. Bot. Gard. 
28:299-374. 1941. 

Gleason, Henry .\llan. Eevision of the North American Vernonieae. Bull. N. Y. Bot. Gard. 
4:144-243. 1906. 

Manton, Irene. Problem of Biscutella laevigata L. Annals Bot. n. s. 1:439-462. 1937. 

Matthew, W. D. Climate and evolution. Ann. N. Y. Acad. Sci. 24:171-318. 1915. 

Pennell, Francis W. Scrophulariaceae of Eastern temperate North America. Monog. Acad. 
Nat. Sci. Phila. l:i-xiv. 1-650. 1935. 



PROCEEDINGS J^/*^ ^*^ ^^'^^ 

:^ I L I -^ R A R y 

OF THE '•*r.\ ;?>^f «*• ) 

CALIFORNIA ACADEMY OF SCIENCE^^jK^^^^^^^S^ 

Fourth Series 
Vol. XXV, No. lo, pp. 245-268, pi. 25 November 10, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 10 
A REVISION OF SOME ARIZONA CACTACEAE 

BY 

LYMAN BENSON 

Associate Professor of Botany 
Pomona College 

THE CACTACEAE, like otlier succulent groups, are frequently neglected be- 
cause of the common impression that specimens are impossible to prepare 
or are not worth much after they are made. It is hoped that recently published 
articles at least have helped to dispel this notion (Benson, 1939 ; Peebles, 
1942). Mr. Peebles has given a particularly thorough account of preparation 
of cactus specimens, and it is illustrated with excellent photographs. Lack of 
adequate series of herbarium specimens tends to place the burden of study 
of the Cactaceae upon examination of living individuals in the field. Despite 
the many advantages of this procedure and its absolute necessity, it is im- 
possible to make adequate comparisons from individual to individual without 
intervening lapse of time, and it is difficult for one person to visit all of the 
localities in which a particular species is to be found. 

When "The Cacti of Arizona" was published (Benson et al, 1940), the 
writer was impressed by the close relationship of a number of described 
species, but evidence of intergradation was not available in the plants exam- 
ined in the field or in the inadequate herbarium material. Continued study 
in the field and preparation and acquisition of additional herbarium speci- 
mens have resulted in the following rearrangement of species, particularly in 
the genus Echinocereus. 

Symbols for herbaria in which specimens are deposited are as follows : Sac, 
U. S. Field Station, Sacaton, Arizona ; UA, University of Arizona, Tucson ; 
B, Herbarium of Lyman Benson, Pomona College, Claremont, California; 
3Io, Missouri Botanical Garden, St. Louis, Missouri; US, United States Na- 

[ 245 ] 



246 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th sek. 

tional Herbarium, Smithsonian Institution. "Washington, D. C. ; NY, New 
York Botanical Garden, New York City; 8, Dudley Herbarium, Stanford 
University, California; 3Iich, University of Michigan, Ann Arbor. A few of 
these specimens have been examined only in figures or photographs. The 
herbarium of the writer is now a part of the Pomona College Herbarium. 

The vegetation types referred to are as follows (ef. Benson, 1942; Shreve, 
1936, 1942; Shantz and Zon, 1924) : 

Circumboreal Flora 

Eockj Mountain Forests (spruce-fir and yellow pine) 
Eastern Forest Flora 
Sierra Madrean Flora 

Southwestern Oak Woodland and Chaparral 

Southwestern Coniferous Woodland (juniper-pinyon) 

Northern Desert (sagebrush) 
Central Prairie Flora 

Short Grass Prairie (Great Plains grassland) 

Desert Grassland 
Mexican Desert Flora (creosote bush) 

Mojavean Desert 

Sonoran Desert 

Chihuahuan Desert 

OPUNTIA 

Opuntia acanthocarpa Engelm. and Bigel. 
Opuntia acanthocarpa Engelm. and Bigel., 1856, Pac. R. E. Eept., 4: 51, pi. 18, f. 1-3. 

Typically an erect, arborescent shrub 1-2 m. high but sometimes lower 
or even sprawling; main trunk very short, usually less than one-fifth the 
height of the plant, the crown of long branches not dense; branches cylin- 
drical, the joints 1.5-3 dm. long or longer, mostly about 2-3 cm. in diameter, 
the surface bearing areoles about 18-30 mm. long and about 6 mm. broad, 
laterally compressed; spines about 10-12 at an areole, the larger ones 2.5-3.7 
cm. long, rather dense on the branches and more or less obscuring the surface 
of the stem, straw-colored, the sheaths conspicuous and persistent ; gloehids 
minute ; petals remarkably variable in color, most frequently red or yellow, 
the flower 2.5-3.7 cm. in diameter; fruit dry and more or less shrivelled at 
maturity, spiny, not giving rise to flowers at the areoles (not proliferous) and 
not persistent. 

Abundant on sandy flats and in washes in the IMojavean and Sonoran 
Deserts at 2,000 to 4,000 feet elevation. California in the southern and eastern 
parts of the Mojave Desert and in the western part of the Colorado Desert; 
southern Nevada ; southwestern Utah ; Arizona from western IMohave County 
and northern Yuma County to the Verde River and the Tucson Mountains 
and southward to Sonora. 

Type collection : "On the mountains of Cactus Pass, about 500 miles west of Santa Fe." 
Bigelow, Mo. 



Vol. XXV] BENSON: REVISION OF ARIZONA CACTACEAE 247 

Opuntia acanthocarpa var. Thornberi (Thornber and Bonker) L. Benson, 

comb. nov. 

Opuntia Thornberi Thornber and Bonker, 1932, Fantastic Clan, 133, 148, pi. opposite 134, 
upper fig. 

Bushy cliolla occurring in patches 1 ni. or more in diameter ; joints of the 
main stem 4 dm. or more in length ; tubercles 3-5 cm. long, about 6 mm. broad 
and 6 mm. high ; spines 7-10, not forming a dense mass on the stem, much less 
conspicuous than the surface of the stem, 1.5-2.2 cm. long. 

Gravelly soil of washes, flats, and alluvial fans in the Mojavean and Sonoran 
deserts at 1,500 to 3,500 feet elevation. Peach Springs, Mohave County, and 
southeastward along the edge of the desert below the Mogollon Rim to north- 
western Graham County and northeastern Pima County. Many of the plants 
occurring within a 10 mile radius of Hillside, Yavapai County, are inter- 
mediate between this and the typical variety. 

Type collection : "Opuntia Thornberi has been described only very recently. . . ." There 
is no evidence of description elsewhere. No type is given, but fortunately the line drawang 
is excellent. 

Opuntia Stanlyi Engelm. 

Opuntia Stanlyi Engelm. in Emory, 1848, Notes Mil. Eecon. p. 157. 

Corynopuntia Stanlyi F. M. Knuth in Backeb. and Knuth, 1935, Kaktus ABC, p. 114. 

Low-growing cholla consisting of a clump of short, erect or ascending 
branches 1-3 dm. high, the clumps often 3-7 m. in diameter; joints narrowed 
below, 1-2 dm. long, usually 3-5 cm. in diameter, the surface bearing con- 
spicuous, mammillate tubercules about 2.5-4 cm. long, 6-12 mm. broad, and 
about 6-8 mm. high or in an extreme form 12-18 mm. high, not tending to 
join one another or to form ridges along the joint ; spines 10-18, those of a 
single areole 1.3-5 cm. long, the larger ones commonly 1-1.5 mm. broad, but 
sometimes 2-3 mm. broad, tan or brownish or sometimes red, strongly flat- 
tened, sheaths wanting or rudimentary and adherent to the tip of the spine, 
papillate-roughened; glochids large ; petals yellow, the flower about 2.5-3 cm. 
in diameter ; fruit markedly spiny, yellow. 

Rocky or sandy mesas or arroyo banks in the Sonoran Desert at 2,500 to 
5,000 feet elevation. Arizona on the San Pedro River above Winkleman and 
from the eastern edge of Aravaipa Valley to the upper Gila River Valley 
and Clifton, Duncan, and the San Simon Valley (Graham, Greenlee, and Co- 
chise counties) ; New Mexico on the Gila River. 

Type collection: "October 22d, 1846. Abundant on the Del Norte and Gila." Type not 
preserved. On page 63, the location of the camp of Oct. 22, 1846, is given as Lat. 32° 38' 13" ; 
Long. 109° 07' 30". This is just south of the Gila River on the boundary between the present 
states of Arizona and New Mexico. 



248 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. 4th ser. 

Opuntia Stanlyi var. Kunzei (Rose) L. Benson, comb. nov. 
Opuntia Kunzei Rose, 1908, Smiths. Misc. Coll., 50: 505. 

Joints narrowed below, 7.5-15 cm. long, 2.5-3.5 cm. in diameter; tubercles 
distinct, about 1.5-2.5 cm. long, narrower and lower than in the typical 
variety; fruit (as far as known to the writer) 3.3-5 cm. long, 12-14 mm. in 
diameter. 

Plains at 1,000 to 2,000 feet elevation. ]\Iaricopa Mountains to Casa Grande 
and southwestward to the Gunsight Mountains and southward to Baboquivari 
Valley. Perhaps occurring in Sonora, 

Type collection : Pima County, Arizona, Br. B. E. Kunze. 

Opuntia Stanlyi var. Wrightiana (Baxter) L. Benson, comb. nov. 

Grusonia WrigMiana Baxter, 1935, Calif. Cactus, p. 58. 
Opuntia Wrightiana Peebles, 1937, Desert Plant Life, 9: 43. 

Joints nearly cylindrical, 1-1.5 or 2 dm. long, 2.5-3.5 or 4 cm. in diameter; 
tubercles tending to be confluent into ribs on the stem, 12-18 mm. long, 3^ 
mm. broad, 3-5 mm. high ; fruit 3.3-5 cm. long, about 12-14 mm. in diameter. 

Sand and clay soils of valley floors in the Sonoran Desert at 300 to 1,500 
feet elevation ; restricted to regions of 1-5 inches of annual rainfall. South- 
eastern edge of California; Arizona from Detrital Valley, Mohave County, 
to'Yuma County and the western edge of Pima County ; southward as far as 
St. George's Bay, Sonora. 

Type collection: "Type Specimen: Three joints, one with fruit, collected by Allan B. 
Clayton, April 15, 1934, and deposited in the Dudley Herbarium of Stanford University. 
Type Locality : Petrified forest near the Colorado Eiver, four miles west of the Quartzite- 
Yuma road, 33 miles north of Yuma, Arizona." 

Opuntia erinacea Engelm. and Bigel. 
Opuntia erinacea Engelm. and Bigel., 1856, Pac. E. E. Eept., 4: 47, pi. 13. f. 8-11. 

Low-growing prickly pear forming clumps 0.3-1 m. in diameter, commonly 
1-1.5 dm. high, the stems usually 2-4 jointed ; joints ovate to elliptic-oblong or 
oblong, flat, 5-15 or 20 cm. long, 5-7.5 cm. broad, glabrous; areoles usually 
about 8-10 mm. apart, all spine-bearing ; spines 4-7 or 9, the longer ones about 
5 cm. long, more or less stiff, somewhat flattened, elliptic in cross section, white 
or pale gray ; glochids 1.5-3 mm. long ; petals yellow, white, deep pink, or red, 
the flower about 5-6 cm. in diameter; fruit dry at maturity, more or less cylin- 
drical, about 3 cm. long, 12-14 mm. in diameter, spiny. 

Alluvial areas in the Mojavean Desert at 1,500 to 3,000 feet elevation. Cali- 
fornia in the southeastern Mojave Desert; southern Nevada; southwestern 
Utah ; northwestern Arizona in Mohave County ; Havasupai Canyon, Coco- 
nino Count}' ; near Carrizo, Apache County. 

Type collection: "West of the great Colorado near the Mojave Creek." Bigeloio in 
1853-4, Mo. 



Vol. XXV] BENSON: BEVISION OF AEIZONA CACTACEAE 249 

Opuntia erinacea var. ursina (Weber) Parish. 

Opuntia nrsina Weber in Boiss., 1898, Diet. Hort., 2 : 896. 

Opiintia erinacea var. ursina Parish in Jepson, 1936, Fl. Calif., 2 : 542. 

Similar to the typical variety; joints obovate to oblong, 1-2 em. long, 
6-7.5 cm. broad; areoles all spine-bearing; spines usually 6-14, the longer 
ones remarkably flexible and thread-like, white or pale gray, not defiexed. 

Rocky mountain slopes in the Mojavean Desert at 2,000 to 3,000 feet eleva- 
tion. California in the southeastern part of the Mojave Desert; Arizona at 
scattered stations in Mohave County. 

Type collection: Ord Mountains, southern Mojave Desert in San Bernardino County, 
California. 

Opuntia erinacea var. hystricina (Engelm. and Bigel.) L. Benson 

comb. nov. 
Opuntia hystricina Engelm. and Bigel., 1856, Pac. R. E. Rept., 4: 44, pi. 15. f. 5-6. 

Joints obovate, 5-10 or 15 cm. long, 3.7-7.5 or 9 cm. broad, areoles all or all 
but the lowest spine-bearing ; spines 4 or usually 5-8, the longest ones 4.5-7.5 
or 10 cm. long, tending to be rigid, defiexed; petals red or yellow. 

Plains and mountains in the Southwestern Coniferous Woodland, the Short 
Grass Prairie, and the Rocky Mountain Forests at 4,500 to 7,000 feet eleva- 
tion. Northern Arizona from Coconino County to Apache County; northern 
New Mexico. Flowering in June. 

Type collection : "... Abundant from the Eio Grande westward to the San Francisco 
mountains, . . .. The specimens before us were obtained at the Colorado Chiquito and on the 
San Francisco Mountains." Bigelow in 1853-4, Mo. 

Opuntia erinacea var. rhodantha (K. Schum.) L. Benson, comb. nov. 

Opuntia rhodantha K. Schum., 1897, La Semaine Hort.; 1898 Gesamtb. Kakteen. 

735. 

Joints obovate, 5-10 cm. long, 3.7-7.5 cm. broad, more or less lead-colored 
(in the other varieties green) ; usually only the distal areoles of each joint 
spine-bearing, those on at least the lower half of the joint usually spineless; 
spines 1-6 or 8, the longer ones 2.5-3.7 cm. long, tending to be rigid, defiexed ; 
flowers usually red but sometimes yellow {e.g. McGee Creek, Mono County, 
California, L. Benson 11212, TJA, B.). 

Flats and rocky hills in the Southwestern Coniferous Woodland, the North- 
em Desert, and the Rocky Mountain Forests at 5,000 to 8,000 feet elevation. 
California along the east side of the Sierra Nevada in southern Mono County 
and Inyo County (Big Trees Camp, Lake Sabrina Road above Bishop, L. Ben- 
son 6009, B) and eastward to western Nebraska; Arizona from Mt. Dellen- 
baugh, Mohave County, to Prescott, the Mogollon Rim, the northern end of 
the White Mountains and northeastward. 

Type collection: "Colorado, bei 200—2300 m Hohe: PURPUS; bliihte bei SPATH im 
Juni " 



250 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Opuntia compressa (Salisb.) Macbr. 

Plate 25, fig. A, 3 

Cactus Opuntia L. 1753, Sp. PI., p. 468. 

Cactus compressxis Salisb., 1796, Prodr., p. 348. 

Opuntia compressa Macbr., 1922, Contr. Gray Herb., (65) : 41. 

Op^mtia vulgaris of authors, not of Mill., 1768. 

Low, prostrate or nearly prostrate prickly pear; roots fibrous; joints light 
green, broadly obovate to nearly circular, 4—11 cm. long, 4—9 cm. broad ; leaves 
4r-5 mm. long, more or less appressed; spines none or solitary in the distal 
areoles; petals pale yellow, the flower about 5 cm. in diameter, "with about 
8 petals" ; fruit 2-3 cm. long. 

Dry gravelly or sandy soil in the Eastern Forest Flora at low elevations. 
Southern New England and southward along the Atlantic coastal plain to 
South Carolina. This species is represented westward by varieties, and the 
Arizona i)lants of both fibrous-rooted and tuberous-rooted types tentatively 
are referred to them. 

Opuntia compressa (Salisb.) Macbr. var. microsperma (Engelm.) 

L. Benson, comb. nov. 

Cactus humifusus Raf., 1820, Ann. Nat., p. 15. 

Opuntia humifusa Raf., 1830, Med. Fl. U. S., 2 : 247. 

Opuntia mesacantJia Raf., 1830, Bull. Bot. Seringe, p. 216. 

Opuntia caespitosa Raf., 1830, loc. cit. 

Opuntia Rafinesquei Engelm., 1856, Pac. R. R. Rept., 4: 41, pi. 11. f. 1-3. 

Opuntia Eafincsquei var. microsperma Engelm., 1856, loc. cit. 

Opuntia cymochila Engelm. and Bigel., 1856, Pac. R. R. Rept., 4: 43, pi. 12. f. 1-3. 

Opuntia stenochila Engelm. and Bigel., 1856, loc. cit., p. 44, f . 4-6, 

Opuntia mesacantlia var. microsperma Coult., 1896, Contr. U. S. Nat. Herb., 3 : 429. 

Opuntia nesacantha var. cymochila Coult., 1896, loc. cit., p. 430. 

Opuntia mesacantlia var. stenocJdla Coult., 1896, loc. cit., p. 430. 

Roots fibrous ; joints dark green, averaging a little larger than in the typical 
variety, 7-12 cm. long, 5-10 cm. broad ; leaves "6-8" mm. long, not appressed; 
spines mostly- 1-3, one much larger than the other (accessory) ones; petals 
"10-12" ; fruit 3-4 cm. long. {Fig. A 3.) 

According to Peebles, apud Kearney and Peebles, U. S. Dept. Agric. Misc. 

Publ., (423) : 611, 1942, this plant, designated as 0. Eafincsquei, occurs in 

northern Arizona from "Apache County to northern Mohave County, not 

common, 5,000 to 7,000 feet . . .." The writer has seen no Arizona specimens. 

This wide-ranging middle western variety of the eastern Opuntia compressa is 

exceedingly variable. It includes a host of minor forms variously considered 

as species and varieties, and some may be worthy of segregation as varieties. 

Type collections: (1.) C. humifusus, "From New York to Kentucky and Missouri." 
(2.) 0. mesacantlia, Kentucky to Louisiana. (3.) 0. caespitosa, Kentucky and Tennessee. 
(4.) 0. Rafinesquei, nom. nov. for the first three described species, and antedated l)y all 
of them. No type designated and only the general range given. (5.) Var. microsperma. 
No type designated and the range not distinguished from the broad range of the typical 



Vol. XXV] BENSON: BETISION OF AEIZONA CACTACEAE 251 

variety (of 0. Eafinesquci) in tlie Mississippi Valley. (6) 0. cymochila, called a subspecies, 
but the name published as a binomial, "On the Camanche plains east of Llaiio Estaeado, 
near the 100th degree of longitude; and from there to Tucumcari, 80 miles east of the 
Pecos" in New Mexico. Bigeldw in 1853-4, Mo. (7.) E. stenochila, called a subspecies but 
published as a binomial, "At the Canyon of the Zuiii," Bigelow in 1853-4, Mo. 

Opuntia compressa var. macrorhiza (Engelm.) L. Benson, comb, no v. 

Plate 25, figs. A, 1, 2 
Opuntia macrorhiza Engelm., 1850, Bost. Jour. Nat. Hist., 6 : 206. 
Opuntia fu si for mis Engelm. and Bigel., 1856, Pac. R. E. Eept., 4: 43. 
Opuntia mesacantha Eaf . var. macrorhiza Coult., 1896, Contr. U. S. Nat. Herb., 3 : 430. 
Opuntia plumbea EosE, Smiths. Misc. Coll., 1908, 50 : 524. 
Opuntia delicata Eose, 1911, Contr. U. S. Nat. Herb., 13 : 310. 
Opuntia Loomisii Peebles, 1938, Cact. and Succ. Jour., 9: 109. 

Similar to var. microsperma; roots tuberous even in young plants; spines 
commonly 2 or more per areole. 

Plains and hills in the Desert Grassland, the Southwestern Coniferous 
Woodland, the Southwestern Oak Woodland, and the Rocky Mountain For- 
ests at 4,500 to 6,000 feet elevation and (eastward) in the Short Grass Prairie 
and in the Eastern Forest Flora. Arizona from Mohave County to Apache 
County, eastern Pima Countj^, and Santa Cruz County ; eastward to Missouri 
and Texas. The Arizona form is variable. Usually it is longer-spined and often 
the joints are larger. Some forms approach the fibrous-rooted species, Opuntia 
phaeacantha. 

Type collections: (1.) 0. macrorhisa, "Naked, sterile, rocky places on the Upper Guada- 
loupe." Texas, Lindheimer, Mo. (2.) 0. fusiformis, called a subspecies, but published as a 

binomial, "Cross-timbers longitude 97°-99° " Bigelow in 1853-4, Mo. (3.) 0. plumbea, 

San Carlos Indian Eeservation, Arizona, Coville in 1894, US. This form is apparently 
identical with that on the Great Plains. A specimen (Pearce, Cochise County, Nichol 
in 1940 [pi. 25 fig. A, 1]) is growing in the Pomona College botany greenhouse. (4). 0. 
delicata, "Type U. S. National Herbarium no. 454622, collected by J. N. Eose at Calabasa, 
April 30, 1908 (no. 11951)." Arizona. (5.) 0. Loomisii, "Type, E. F. Loomis No. SF. 298, 
Prescott, Arizona, Aug. 10, 1930, represented by a plant grown under cultivation at Sacaton, 
Arizona, and material removed from the cultivated plant and deposited in the U. S. Na- 
tional Herbarium, Washington, D. C."- This is the extreme opposite the type described by 
Eose as 0. plumbea, and it approaches 0. phaeacantlia. 

Opuntia tortispina Engelm. and Bigel. 

The plant described previously as Opuntia tortispina is merely a form of 
Opuntia phaeacantha with long, twisted, light-colored spines. The description 
was drawn from plants collected four miles south of Navajo Bridge, Coconino 
County, by A. A. Nichol in 1940. 

ECHINOCEREUS 

Mostly small cacti forming low clumps 1-4 dm. or rarely about 1 m. in 
diameter and rarely more than 3 (maximum 6) dm. high or the stems not 
infrequently solitary, the length of a single branch not more than 8 (or rarely 



252 CALIFOBNIA ACADEMY OF SCIEXCES [Proc. -Ith ser. 

in a decumbent type 12) times the diameter ; stem usually branching only at 
or near the base except after injury, or sometimes unbranched, with continu- 
ous ribs, cylindrical or ovoid, leafless even on the new season's growth; areoles 
not producing glochids ; flower appearing below the apex of the branch ; the 
flower bud bursting through the epidermis above the lower edge of a spine- 
producing areole; floral tube (hypanthium?) long or at least well-developed 
above the ovary; fruit spiny, the clusters of spines readily removed at ma- 
turity. 

The plants of this genus occurring in Arizona are members of complex 
groups. Previously (Benson et al., 1940) ten described species were recog- 
nized as closely related entities, although it was clear that the group was in 
need of reorganization and that several species might better be reduced to 
varieties. Further study in the fleld and collection of additional herbarium 
and garden material emphasizes the need for the following revision. The dif- 
ferentiation of this genus from Cereus has been discussed in another paper 
(Benson, 1941). 

Key to the Species 

1. Petals scarlet or crimson (with plastid pigmentation?); areoles bearing white felt at 
maturity; flowers not closing at night, remaining open until the end of anthesis. 

1. Ecldnocereus trifflocliidiatits. 

1. Petals purple to lavender (probably pigmented with anthocyans) ; areoles not bearing 

white felt at maturity, the felt of the young areoles rarely persistent for two years; 

flowers closing at night and reopening in the morning or in hot weather withering at the 

end of a single day. 

2. Areoles vertically elongate; stems unbranched 2. Echinocereus pectinatus. 

2. Areoles circular; stems usually branched except in young plants. 

3. Central spines 2—6, approximately the same size or at least all well developed, usually 
(at least the lower deflexed ones) flattened at least at the bases. 

3. Echinocereus Engelmannii. 
3. Central spine 1, often with 1-3 shorter accessory or rudimentary centrals in some of 
the areoles of the plant, circular in cross section, not flattened. 

4. Principal central spine not both strongly curved and pointing do^^^lward ; spines 
white, gray, red, black, bro^\^l, or bicolored; flowers 5-8.5 cm. in diameter. 

4. Echinocereus Fendleri. 

4. Principal central spine strongly curved near the base, turned do^^^lward ; spines 

yellow or straw-color, uniformly colored; flowers (in the specimens available) 

3.5-5 cm. in diameter. 5. Echinocereus Ledingii 

1. Echinocereus triglochidiatus Engelm. 

Echinocereus triglochidiatus Engelm., in Wisliz., 1848, Mem. Tour. N. Mex., p. 93. 
Cereus triglochidiatus Engelm., apud A. Gray, 1849, Mem. Amer. Acad., II. 4: 50. 
Echinocereus paucispinus (Engelm.) Engelm., ex. Eiimplcr var. triglochidiatus K. Schum., 
1898, Gesamtb. Kakteen, 281. 

Older plants dense, forming clumps of few to many stems each 2^ or 6 dm. 
high, mostly 4—8 cm. in diameter, obviously and conspicuously green, not 
obscured by the spines, ribs commonly 5-8, somewhat tuberculate at the 
areoles ; spines exceedingly variable, sometimes as few as 3 or as many as 16, 



Vol. XXV] BENSON: REVISION OF ARIZONA CACTACEAE 253 

1-3 em. long; central spines 0-5 or 6, smooth and ribbed, somewhat longer 
than the radials ; radial spines 3-10; areole circular, bearing white felt ; petals 
crimson or scarlet, the flower about 3 cm. in diameter ; floral tube 2-3 cm. long 
above the ovary, the scales with tufts of white hair ; fruit more or less spiny, 
subcylindrical, about 2-3 cm. long. 

Southern Colorado to trans-Pecos Texas. A few Arizona plants approach 
the typical variety. 

Type collection: "On "Wolf Creek [near Santa Pe, New Mexico], in pine woods, flowers 
in June [Wislizenius in 1846, Mo.] ; Santa Fe (Fendler) " Mo. 

1a. Echinocereus triglochidiatus var. gonacanthus (Engelm. and Bigel.) 

Engelm. and Bigel. ex W. T. Marshall 

Cereus gonacanthus Engelm. and Bigel., 1856, Pae. E. E. Eept., 4: 33, pi. 5. f. 2-3. 
Echinocereus gonacanthus Engelm., ex Eiimpler in Forster, 1885, Handb. Cacteenk., ed. 

2, p. 806. 
Echinocereus paucispinus (Engelm.) Engelm., ex Eiimpler var. gonacanthus K. Schum., 

1898, Gesamtb. Kakteen, p. 281. 
Echinocereus triglochidiatus var. gonacanthus (Engelm. and Bigel.) Engelm. and Bigel., 

ex W. T. Marshall in Marshall and Bock, 1941, Cactaceae, p. 117. 

A robust type similar to the typical variety; spines mostly 6-8 per areole, 
twisted and somewhat curved ; central spine usually solitary but sometimes 
more than one, 3-5 cm. long, markedly angled wdth usually 6 or 7 sharp ribs ; 
radial spines 2-3 cm. long. 

High plains at the lower edge of the Rocky Mountain Forests at 6,000 to 
8,000 feet elevation. Arizona 40 miles west of Zuiii, New Mexico, and near St. 
Johns, Apache County, (Wyatt W. Jones, the specimen with 3-angled central 
spines.) New Mexico near Gallup and Zuni. Reported from the White Sands. 

Type collection: "On high sand-bluffs, covered with scattering cedars, near the natural 
well, about 40 miles west of Zuni, near the 109th degree. Only seen in that locality." Prob- 
ably Apache County, Arizona, Bigelow in 1853-4, Mo. Zuni, New Mexico, is within 10 or 
12 miles of the Arizona line, but the 109th Meridian is just east of the state boundary. 

1b. Echinocereus triglochidiatus var. polyacanthus (Engelm.) L. Benson, 

comb. nov. 

Echinocereus polyacanthus Engelm. in Wisliz., 1848, Mem. Tour. N. Mex., p. 104. 
Cereus polyacanthus Engelm. apud A. Gray, 1849, Mem. Amer. Acad., II, 4: 50. 
Echinocereus arisonicus Orctjtt, 1926, Cactography, (3) : 3. 

Clumps of stems not dense or dome-like, the stems usually 15^5, 1.5-5 dm. 
high, 6-10 cm. in diameter, elongated and cylindrical ; ribs about 9-10, promi- 
nent, not markedly tuberculate at the areoles; central spines 2-4, turned down- 
ward or perpendicular to the stem, straight, stout or slender, 2.5-5 cm. long ; 
radial spines about 10, appressed ; flower about 3.5-4 cm. in diameter ; floral 
tube about 3 cm. long above the ovary. 

Desert Grassland and Southwestern Oak Woodland at 3,500 to 5,500 feet 
elevation. Arizona in the Pinal Mountains, Pinal County, and the Galliuro 



254 CALIFORNIA ACADEMY OF SCIENCES [Proc. iiH ser. 

Mountains, Graham County, and southward to Santa Cruz and Cochise coun- 
ties. Southeastern Arizona to western New Mexico, Chihuahua, and Durango. 
Floweriiip- late in ^lay and in June. 

Type collections: (1.) E. poly acanthus, "Cosiluiiriachi," Chihuahua. Wifilizenius in 
1846-7, Mo. (2.) E. arizonicus, Pinal Mountains, Pinal County, Arizona. 

Ic. Echinocereus triglochidiatus mw. melanacanthus (Engelm.) L. Benson, 

comb. nov. 

Echinocereus coccineus Engelm. in Wisliz., 1848, Mem. Tour. N. Max., p. 93. 

Cereus coccineiis Engelm. apud A. Gray, 1849, Mem. Amer. Acad., II, 4: 51. 1849, not 

Salm-Dyck ex DC. in 1828. 
Cereus coccineus yar. melanacanthus Engelm., loc. cit. 
Cereus coccineus var. cylindricus Engelm., loc. cit. 

Cereus majavensis var. suniensis Engelm. and Bigel., 1856, Pac. R. R. Rept., 4 : 33, pi. 4, f . 9. 
Cereus Bigelovii Engelm. var. zuniensis Engelm., apud Engelm. and Bigel., 1856, loc. cit. 

(on the plate). 
Cereus phoeniceus Engelm. apud Engelm. and Bigel., 1856, loc. cit., p. 34, f . 1-3. 
Echinocereus phoeniceus Engelm., ex Riimpler in Forster, 1885, Handb. Cacteenk., ed. 2, 

p. 788. 
Echinocereus mojavensis var. zuniensis Engelm., ex Riimpler, Forster, 1885, loc. cit., p. 803. 
Echinocereus phoeniceus var. inermis K. ScHUM., 1896, ]Monatsschr. Kakteenk., 6: 150. 
Echinocereus coccineus var. inermis 3. A. Purpus, 1925, Mitt. Deutseh. Dendr. Ges., 1925 : 49. 
Echinocereus triglochidiatus var. coccineus Engelm., ex W. T. Marshall in ]SlARsnALL and 

Bock, 1941, Cactaceae, p. 117. 
Echinocereus canyonensis Clover and Jotter, 1941, Bull. Torrey Club, (58 : 417. f . 1 (4). 

Stems from a few to more than 500, cylindrical or ovoid, 0.7-1.5 dm. high, 
mostly 3-5 or 6 cm. in diameter ; ribs commonly 8 or 10, not markedly tuber- 
culate at the areoles ; central spine 1 or sometimes more than 1, perpendicular 
to the joint, straight, tapering, 1.3-3 cm. long; radial spines commonly 5-10, 
similar to the central spine but smaller; flow^er 3-4.5 cm. in diameter; floral 
tube 2.5-3 cm. long above the ovary. 

Rocky hillsides and the forest floor in the Southwestern Coniferous Wood- 
land, the Southwestern Oak Woodland and Chaparral, and the Rocky ]\Ioun- 
tain Forests at 4,000 to 8,000 feet elevation. Arizona in the mountainous 
regions from the Ilualpai Mountains to the Kaibab Forest, and southeastward 
to the Santa Catalina Mountains, Pima County, and to Greenlee County, 
thence southward to Santa Cruz and Cochise counties. Southern Utah to 
Colorado, Arizona, and New Mexico. Flowering from March until June, de- 
pending upon altitude. 

Type collections: (1.) E. coccineus, "With the foregoing, also about Santa Fe." The 
"foregoing" was E. triglochidiatus, see above. Mo. (2.) Var. melanacanthus, "Among a 
number of plants of this family Avhich Mr. Fendler sent from Santa Fe in a living state, 
but which unfortunately were all dead when they came to hand, are some specimens which 

appear to be varieties of this species " (3.) Var. cylindricus, cf . statement for var. 

melanacanthus. Mo. (4.) Var. suniensis, "It was found near Canon Diablo, on the Colorado 
Chiquitn, about 120 miles west of Zufii." Canyon Diablo, near the Little Colorado River, 
Arizona, Bigclow in 1853-4, Mo. (5.) C. phoeniceus, nom. nov. for E. coccinms. (6.) Var, 



Vol. XXV] BEN SON : BEVISIOX OF ABIZON A CACTA CEAE 255 

hterniis,". . . Heir C. A. Purpus aus Colorado an die liiesige beriihinte Gaitueiei von SPATH 
schickte . . .." (7.) E. conyonensis, "TYPE {Clover and Jotter 2317) collected in a sandy 
pocket on a steep limestone outcrop 100 yards from the [Colorado] river, Bass Cable below 
Hermit Creek Eapids, Grand Canyon, Coconino Co., Arizona. Locally abundant. Living 
specimen in the Botanical Gardens, (Gard. 16846), University of Michigan, Ann Arbor, 
Michigan." Specimen now pressed. 

Id. Echinocereus triglochidiatus var. mojavensis (Eiigelm.) L. Benson, 

comb. nov. 

Cereus mojavensis Engelm. and Bigel., 1856, Pac. E. E. Eept., 4 : 33. pi. 4. 
Cereus Bigelovii Engelm., apud Engelm. and Bigel., 1856, loc. cit. (on the plate.) 
Echinocereus mojavensis Engelm., ex. Eiimpler in Forster, 1885, Handb. Cacteenk., ed. 2, 
p. 803. 

Similar to var. melanacanthus ; ribs (of the material available) strongly tu- 
bereiilate at the areoles; spines curving, tortuous, somewhat flexible, Avhitish; 
floral tube (of the plants available) 1.7-2 cm. long. 

Rocky hillsides in the upper Mojavean Desert and the Southwestern Coni- 
ferous Woodland at 3,500 to 5,000 feet elevation. Arizona in scattered locali- 
ties in Mohave County (Black Mountains; Grand Wash Cliffs; Hualpai 
Indian Eeservation). Southeastern California to southern Utah and north- 
western Arizona. 

Type collections: (1.) C. mojavensis, "Found between the Eio Colorado and Mojave 
Creek." California east of the Mojave River, Bigelow in 1853-4, Mo. (2.) C. Bigelovii, 
apparently a discarded name once intended to be proposed for the same plant. 

2. Echinocereus pectinatus (Scheidw.) Engelm. 

Echinocactus pectinatus Scheidw., 1838, Bull. Acad. Sci., Brux., 5 : 492. 

Eehinocactus pectiniferus Lemaire, 1839, Cact. Gen. Nov. sp., p. 25. 

Ecliinopsis pectinata Fennel, 1843, Allg. Gartenz., 11 : 282. 

Echinocereus pectinatus Engelm., in WiSLiz., 1848, Mem. Tour. N. Mex., p. 110. 

Cereus pectinatus Engelm., apud A. Gray, 1849, Mem. Amer. Acad. Sci., II, 4: 50. 

Cereus pectiniferus Labouret, 1853, Monogr. Cact., p. 320. 

Stem unbranehed, erect, cylindrical, 1-2 dm. high, 3-8 cm. in diameter ; ribs 
about 20, not markedly tuberculate at the areoles, but transversely grooved 
between adjacent areoles; spines white to red, not completely obliterating the 
stem ; central spines 2-5, mostly 3, mostly 1-3 mm. long, in a single row ; 
radial spines 15-20, appressed against the stem, pectinate, markedly ap- 
pressed, 3-8 mm. long, flattened, straight, tapering; areoles vertically elon- 
gate, the growth of different years marked off by the color of the spines 
produced in early or late season; flower funnel-form; petals reddish-purple, 
tending toward lavender ; floral tube about 12 mm. long above the o.vary ; fruit 
nearly spherical, about 2.5 cm. in diameter. 

Rocky or gravelly hills in the Desert Grassland and perhaps the Southwest- 
ern Oak Woodland at 4,000 to 5,000 feet elevation. Southeastern Arizona to 
Chihuahua. The plant occurring in Cochise Count}^, Arizona, seems to be 
referable to the typical variety, but it has not been seen in flower (Perilla Mts., 



256 CALIFORNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

Harlan in 1939, Sac; near Douglas, W. T7. Jones TJA, B; 16 miles northeast 

of Donglas, Benson and Darrow 11113, UA, B). This is one of several Chi- 

huahiian Desert species occurring on limestone hills in southeastern Arizona. 

Type collection: The identity of the type is difficult to establish. According to Britton 
and Rose, Cactaceae, 3: 29-30, 1922, "This species was first collected by Galeotti who sent 
a collection to Belgium from the states of San Luis Potosi and Guanajuato, Mexico. The 
type station, Villa del Pennasco, we have not located. It was soon after figured by Lemaire 
(Icon. Cact., pi. 14 or 15) and Pfeiffer (Abbild. Beschr. Cact., 2: pi. 10), very likely from 
the type collection." According to Engelmann, loc. cit., however, the type is from Chihuahua: 
"The description of the plant (which died ^\^thout producing flowers) found in several 
works, as well as in the latest publication on Cactaceae, before me, of Foerster, Leipzig, 
1846, was made, as Prince Salm informed -me, from specimens sent from Chihuahua by 
Mr. Potts. It entirely agrees with my specimen from the same region." 

2a. Echinocereus pectinatus var. rigidissimus (Engelm.) 
Engelm. ex Kiimpler 

Cerevs pectinatus (Scheidw.) Engelm., var. rigidissimus Engelm., 1857, Proc. Amer. Acad., 

3: 279. 
EcMnocere\is rigidissimus Rose, 1909, Contr. U. S. Nat. Herb., 12 : 293. 
Echinocereiis dasyaeanthus Engelm., var. rigidissimus W. T. Marshall in Marshall and 

Bock, 1941, Cactaceae, p. 119. 

Stems 0.7-3.5 dm. high, 5-10 cm. in diameter ; areoles crowded, the spines 
completelj' obliterating the surface of the stem; central spines none; radial 
spines flat, thicker and stronger than in the typical variety, 15-18, strongly 
appressed, 6-14 mm. long ; flowers between lavender and magenta. 

Occurring on rocky sloj^es and grassy areas in the Desert Grassland and the 
Southwestern Oak Woodland at 4,000 to 6,000 feet elevation. Arizona from 
the Baboquivari Mountains to the Santa Catalina Mountains, Pima County, 
and southeastward to Santa Cruz and Cochise counties ; northern Sonora. 

Type collection : "... from Sonora . . .." According to Engelmann, in Emory, Rept. IT. S. 
and Mex. Bound. Surv., 2 : 32, 1859. "In the Sierras of Pimeria Alta in Sonora, and farther 
west (A. Schott). Flowers in June and July." Mo. 

2b. Echinocereus pectinatus var. neomexicanus (Coulter) L. Benson, 

comb. nov. 

Echinocereus dasyaeanthus Engelm., in Wisliz., 1848, Mem. Tour. N. Mex., p. 100. 
Cereus dasyaeanthus Engelm., apud A. Gray, 1849, Mem. Amer. Acad., II, 4: 50. 
Cereus ctenoides Engelm., in Emory, 1839, Rept. U. S. and Mex. Bound. Surv., 2 : 31, pi. 42. 
Cereus ctenoides Engelm., et Riimpler in Forster, 1885, Hand. Cacteenk., ed. 2, p. 819, f . 109. 
Cereus dasyaeanthus var. neomexicanus Coult., 1896, Contr. U. S. Nat. Herb., 3: 384. 
Not Echinocereus neomexicanus Standl., 1908, Bull. Torrey Club, 35: 87. 

Stems 1-3 dm. high, 5-7 or 10 cm. in diameter; areoles crowded, the spines 
usualh' practically obliterating the surface of the stem ; central spines mostly 
5-8, usually arranged irregularly (or in a single row in types transitional to 
the typical variety), 3-10 mm. long; radial spines somcAvhat flattened, 15-20 
or more, not markedly appressed, 10-15 mm. long; flowers typically yellow 
but sometimes varying to the magenta series. 



Vol. XXV] BENSON: REVISION OF ARIZONA CACTACEAE 257 

Desert Grassland and Oak Woodland at 4,000 to 5,000 feet elevation. Ari- 
zona (according to Professor Thornber) in the southern part of the Baboqni- 
vari Mountains, Pima County, and (according to A. A. Nichol) in the Perilla 
and Guadaloupe mountains, Cochise County, as well as (according to A. A. 
Nichol) in jjerhaps modified form in the Oro Blanco Mountains west of 
Nogales, Santa Cruz County, and at Altar and Pitiquito in adjacent Sonora; 
Southern New Mexico ; Trans-Pecos Texas ; Chihuahua. 

Type collections: (1.) E. dasy acanthus, "On the mountains near El Paso; in August in 
flower and fruit." Wislizenius in 1846-7, Mo. (2.) C. ctenoides, "From Eagle Pass to Santa 
Eosa (Bigelow) ; on the Pecos (Wright.)" Texas. Mo. (3.) Var. neomexicanus, "Type, 
Wright 366 in Herb. Mo. Bot. Gard. Southeastern New Mexico." The necessity for use of 
this name in varietal status is unfortunate, for the little-known epithet neomexicanus dis- 
places the well-known one, dasy acanthus, and the epithet neomexicanus was proposed later 
in specific rank for a plant of the Echinocereus triglochidiatus complex. 

3. Echinocereus Engelmannii (Parry) Parry ex Rlimpler 

Cereus Engelmannii Parry ex Engelm., 1852, Amer. Jour. Sci., II, 14: 338. 

Cereus Engelmannii var. variegatus Engelm., 1856, Pac. E. E. Eept., 4 : 35, pi. 5. f . 4-7. 

Cereus Engelmannii var. chrysocentrus Engelm., 1856, loc. cit., f . 8-10. 

Echinocereus Engelmannii (Parry) Parry ex Eihnpler in Forster, 1885, Handb. Cacteenk., 

ed. 2, p. 805. 
Echinocereus Engelmannii var. variegatus Engelm., ex Eiimpler, Forster, 1885, loc. cit., 

p. 806. 
Echinocereus Engelmannii var. chrysocentrtis Engelm., ex Eiimpler, Forster, 1885, loc. cit., 

p. 806. 
Echinocereus chrysocentrus Orcutt, 1926, Cactography, (3) : 4. 

Clumps not dense, stems erect, usually 1-25, not crowded, cylindrical, 
usually 1.5-3 dm. high, 5-7.5 cm. in diameter, firm, ribs usually 10-13, not 
markedly tuberculate at the areoles ; spines sometimes obliterating the surface 
of the stem; central spines 2-6, all large and well-developed, red, yellow, white, 
brown, or gray, or the lower one white and the others variously colored, some 
of them deflexed, tapering, 3.7-9 cm. long, stout but somewhat flexible, curv- 
ing or twisted, usually flattened at the bases ; radial spines 6-12, similar to 
the central but smaller ; areole circular or nearly so, not bearing white felt 
after the first year ; petals purple to magenta, the flower 5-7.5 cm. in diameter, 
funnel-form; floral tube about 12 mm. long above the ovary ; fruit red, nearly 
spherical to ovoid, about 3 cm. long, spiny, shedding the interlocking spines at 
maturity, edible. 

Hills, plains, alluvial bottoms, and benches in the Northern, Mojavean, and 
Sonoran deserts at 100 to 2,500 or rarely 5,000 feet elevation. California about 
the Owens River Valley and in the Mojave and Colorado deserts ; southern 
Nevada ; southern Utah ; Arizona from southern Mohave County to Yuma 
County, Pinal County, and western Pima County; Baja California to Sonora. 
Flowering in late March and in April. 

Type collections: (1.) C. Engelmannii, "Mountains about San Felipe, on the eastern 
declivity of the Cordilleras." Colorado Desert in San Diego County, California, Parry in 



258 CALIFORNIA ACADEMY OF SCIEXCES [Pkoc. 4th skk. 

1852, Mo. (2.) Var. varicgatus, "On the Cactus mountains and at the head of Williams 
Eiver; degrees 113i/^ longitude." Arizona, Bigelow in 1853-4 Mo. (3.) Var. chrysocentrus, 
"It was found where C. variegatus [var. varie gains'] disappears on the lower part of 
Williams's River, and was seen from there to Mojave Creek, and up that stream to the Sierra 
Nevada." Western Arizona and California, Bigelow in 1853-4, Mo. The species is l)arely 
to be maintained as distinct from E. Fendleri, and the separation is rather arbitrary. 

3a. Echinocereus Engelmannii v;n-. Nicholii L. Benson, var. no v. 

Plate 25, fig. B. 

Caulibus 10-30 erectis, 3-5 dm. longis, 5-7 vel 8 cm. diametro ; aculeis flavis ; 
petalis colore similibus lavendulae. 

Stems erect, in large clumps of 10-30, 3-5 dm. high, 5-7 or 8 cm. in diame- 
ter ; spines all yellow ; petals lavender. 

Common on rocky hillsides in the Sonoran Desert at 1,000 to 2,000 feet 
elevation ; not occurring on the alluvial flats between the mountains. Arizona 
in western Pima County from the Organ Pipe Cactus National Monument to 
the Silver Bell Mountains ; largely on the Papago Indian Reservation. 

Type collection: Silver Bell Mine, Silver Bell Mountains, Pima County, Arizona, L. 
Benson 10720, March 28, 1941, TYPE VA ; ISOTYPE B. Mr. A. A. Nichol has brought 
many plants of this variety to the University of Arizona on the suppostion that they Avere 
a new entity. 

3i?. Echinocereus Eng-elmannii var. decumbens (Clover and Jotter), 

L. Benson, comb. nov. 
Echinocereus decumbens Clover and Jotter, 1941, Bull. Torrey Club, 68: 417, fig. 7, 1941. 

Stems decumbent, up to 50 or more, forming dense masses, elongated and 
slender, 2-6 dm. long, 3-5 cm. in diameter; spines yellowish to white or 
pinkish, the lower deflexed and flattened central spine 2.5-3.5 cm. long, the 
other centrals and the radials 1.5-2.3 cm. long, relatively weak and flexible ; 
flowers unknown. 

Talus and ledges in precipitous canyons of the Sonoran Desert at about 
2,000 to 3,000 feet elevation. Arizona at Mile 161/2, Marble Canyon, Colorado 
River, Coconino County, and at Palm Canyon, Kofa Mountains, Yuma 
County (Benson and Darrow 10870 VA, B). The Palm Canyon plant grew 
on top of a huge boulder where almost no soil was present, and it covered an 
area three or four feet in diameter. 

Type collection : "Type on limestone ledge 30 feet from the river's edge and at the base 
of rocky talus at Mile 161/2 Marble Canyon, Coconino Co., Arizona {Clover and Jotter 2212). 
Living Hi)eeimen at the Botanical Gardens of the University of Michigan, Ann Arbor, 
Michigan (Gard. 16S70. Fig. 7.)" Specimen now pressed. 

4. Echinocereus Fendleri (Engehn.), Engelm. ex Riimpler 

Cereus Fendleri Engelm., apud A. Gray, 1849, Mem. Amer. Acad., II. 4 : 51. 
Cereus Fendleri var. paupcrculus Engelm., A. Gray, 1849, loc. cii. 

EcJiinocerevs Fendleri (Engelm.), Engelm. e.x TJiiniiiler in Forstek, 1885, Handb. Cacteenk., 
ed. 2, p. 801. 



Vol. XXV] BENSON : BETISION OF ARIZONA CACTACEAE 259 

Clumps not dense, tlie stems 1-5 or perhaps more, not crowded against one 
another, ovoid, 7-15 cm. long, 5-10 cm. in diameter, flaccid, ribs usually about 
9-10, not markedly tuberculate at the areoles ; spines not obliterating the sur- 
face of the stem; central spine 1, at first very dark brown to black, gray in 
age, turned upward from the base, tapering, 2-4.5 cm. long, stout, slightly 
curving upward the entire length, rigid, not flattened ; radial spines 9-11, 
similar to the central but shorter and white or pale gray ; areole circular, not 
bearing white felt at maturity, the felt rarely persisting two years ; petals 
magenta to purplish, the flower about 5-7 cm. in diameter; floral tube about 
12 mm. long above the ovary ; fruit ovoid, about 3 cm. long, spiny, the spine- 
clusters interlocking, falling away as a sheet at maturity. 

Grassland and open areas in the Rocky Mountain Forests at 5,000 to 7,000 
feet elevation. Northern Arizona in Coconino, Navajo, and Apache counties ; 
southern Utah ; northern New Mexico. 

Type collections: (1.) C. Fendleri, "Santa Fe, on elevated sandy plains; flowering in 
June." Fcndler, Mo. (2.) Var. pauperculus, apparently collected in the same vicinity by 
Fendler, Mo. 

4a. Echinocereus Fendleri var. rectispinus (Peebles), L. Benson, comb. nov. 
Echinocereus recfispmws Peebles, 1938, Amer. Jour. Bot., 25 : 675, f. 1, 3d. 

Stems 1-5, usually 1-2 dm. high, 6-9 cm. in diameter, firm, the ribs 8-10; 
central spine perpendicular to the stem, straight, 1.2-2.5 cm. long, in some 
areoles accompanied by small accessory centrals, rigid, rather stout; flower 
about 6-6.5 cm. in diameter. 

Eocky slopes and benches in the Desert Grassland at 3,500 to 5,500 feet 
elevation. Arizona from Nogales and the Santa Rita Mountains to Ft. Grant 
and Cochise County. 

Type collection: "Type: Peebles No. SF 905 (fig. 1), hills near Nogales, Arizona, eleva- 
tion 3,900 feet. May 5, 1935, represented by a living plant under cultivation at U. S. Field 
Station, Sacaton, Ariz., and by material deposited in the U. S. National Herbarium as 

No. 1729266." 

4b. Echinocereus Fendleri var. robustus (Peebles), L. Benson, comb. nov. 

Echinocereus rectispinus Peebles var. robustus Peebles, 1938, Amer. Jour. Bot., 25 : 675, 

f. 3c. 
Echinocereus robustus Peebles, 1940, Jour. Wash. Acad. Sci., 30 : 219. 

Stems 5-20, 1.7^.5 dm. high, mostly 8-10 cm. in diameter, ribs 8-13, firm ; 
central spine straight, perpendicular to the stem, 2.5-7.5 cm. long, in some 
areoles accompanied by small accessory centrals, rather flexible, slender ; 
flower usually 7.5-8.5 cm. in diameter. 

Rocky slopes in the Sonoran and Chihuahuan deserts and the lower part 
of the Desert Grassland at 2,000 to 4,000 feet elevation. Arizona from eastern 
Pinal County and eastern Pima County to the upper Gila River valley and 
the lower parts of Cochise County. This is the only Echinocereus in the desert 
about Tucson. Flowering in April and May. Closely related to the next variety. 



260 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th sek. 

Type collection: "Type: Peebles No. SF 896, Tucson to Sabino Canyon, Pima County, Ari- 
zona, August 27, 1935, represented by a living plant g^o^\^l under cultivation at Sacaton, 
Ariz., and by material deposited in the U. S. National Herbarium as No. 1729267." 

4c. Echinocereus Fendleri var. Boyce-Thompsonii (Orcutt), L. Benson, 

eoinb. nov. 

Echinocereus Boyce-Thompsonii Orcutt, 1926, Cactography, (3) : 4. 

Stems 1-10, 1-2.5 dm. high, 5-7.5 em. in diameter, rather firm, ribs 12 or 
commonly 14-18 or sometimes more; central spine 2.5-4 cm. long, accompanied 
in at least some areoles by 1 or 2 accessory centrals, perpendicular to the stem 
or more commonly deflexed, straight, flexible, very slender, remarkably light- 
colored, either straw-colored, reddish, or white (gray in the two preceding 
varieties) ; flower usually about 6-6.5 cm. in diameter. This variety is transi- 
tional in some of its forms to Echinocereus Engelniannii. 

Rocky slopes or grassy uplands in the upper part of the Sonoran Desert, 
the Desert Grassland, the Southwestern Coniferous Woodland, and the South- 
western Oak Woodland at mostly 2,500 to 5,500 feet elevation. Central Arizona 
from the vicinity of Seligman southeastward just below the Mogollon Rim to 
the upper Gila River Valley. Flowering from March until May. 

Type collection : "Grounds of the Boyce-Thompson Southwestern Arboretum, near Su- 
perior, Arizona, at an elevation of about 2,300 feet." The original description was reprinted 
by Peebles, Amer. Jour. Bot., 25: 677. 1938. 

4d. Echinocereus Fendleri var. Bonkerae (Thornber and Bonker), 

L. Benson, comb. nov. 

Echinocereus Bonkerae Thornber and Bonker, 1832, Fantastic Clan, p. 71-3, 85, pi. oppo- 
site p. 28, 72. 

Clumps rather dense for the group, the stems usually 5-15, 1.2-2 dm. high, 
3.5-6.5 cm. in diameter, rather firm, ribs 11 or 13-16 ; central spine 6-8 mm. 
long, accompanied in some areoles by 1 or 2 accessory centrals, perpendicular 
to the stem, rigid, white or pale gray tipped with brown ; flower tending toward 
purple, about 6-6.5 cm. in diameter, the petals and sepals more nearly rotate 
than in the other varieties, or even slightly reflexed. 

Hills and slopes in the Desert Grassland and the Southwestern Oak Wood- 
land at 3,500 to 5,000 feet elevation; best developed in shady situations where 
leaf -mold is present. Arizona from the vicinity of Globe to the Santa Catalina 
Mountains and the Graham (Pinaleno) Mountains. Specimens collected by 
A. A. Nichol in 1940 are clearly transitional between this and var. Boyce- 
Thompsonii. 

Type collection: "We are nearing the beautiful Pinal Mountains in southeastern Ari- 
zona. . . ." According to A. A. Nichol, the plant described by Professor Thornber was from 
near Oracle at the north base of the Santa Catalina Mountains. Mr. Nichol planted the 
specimens in the University of Arizona Cactus Garden, and in 1939, the writer placed the 
remains of the last individual in the University Herbarium. They are the closest approach 
to a type. Coulter, Contr. U. S. Nat. Herb., 3 : 384, 1896, described a plant probably of this 
variety from Oracle (Evans in 1891), placing the vegetative material near Cereus ctenoides 



Vol. XXY] BENSON : BEVISIOK OF ASIZOXA CAC TACEAE 261 

Engelm. (Echinocereus pectinaius var. neomexicanus above). Britton and Rose, Cactaceae, 
2: 37, 1922, published a photograpli of this plant (fig. 45) as E. Fendleri. 

5. Echinocereus Ledingii Peebles 

Echinocererts Ledingii Peebles, 1936, Caet. and Succ. Jour., 8: 35. 

Clumps not particularly dense ; longer stems cylindrical, 1.5-4 or 5 dm. 
high, 6-9 or 10 cm. in diameter, ribs 12-16, not markedly tuberculate at the 
areoles ; spines not completely obliterating the surface of the stem, light yellow 
or straw-color; central spine 1, but often accompanied by 1-3 short upper 
accessory centrals, deflexed and strongly curved downward near the base, 
about 2 cm. long, stout, rigid, tapering; radial spines about 10-12, straight, 
about 9 mm. long; areole circular or nearly so, not bearing white felt; petals 
rose-purple, the flower 3.5-5 cm. in diameter; floral tube about 9 mm. long 
above the ovary; fruit pale green, spherical, a little less than 1 inch in diam- 
eter, spiny when young. 

Slopes in the Southwestern Oak Woodland at 4,500 to 6,000 feet elevation. 
Southeastern Arizona in the Quinlan, Santa Catalina, Santa Rita, Huachuca, 
Graham (Pinaleno), and Chiricahua Mountains in Pima, Graham, Santa 
Cruz, and Cochise counties. Perhaps endemic in Arizona. The localities are 
based upon records supplied by A. A. Nichol ; specimens have been obtained 
only from the Graham Mountains. 

Type collection : "Type collected at about 4,500 feet elevation on slopes of Mt. Graham, 
Pinaleno Mountains, Graham Co., Arizona, Louis Wankum, July 11, 1935, U. S. Nat. Herb. 
No. 1,634,004." 

ECHINOCACTUS 

In this genus only a few changes are proposed now, and other problems are 
deferred pending accumulation of additional data. 

Echinocactus acanthodes Lemaire 

Echinocactus acantJiodes Lemaire, 1839, Cact. Gen. Nov. Sp., p. 106. 

Echinocactus viridescens var. cylindraceus Engelm., 1852, Amer. Jour. Sci., II : 338. 

Echinocactus LeContei Engelm., 1856, Pac. E. R. Eept., 4: 29. 

Echinocactus cylindraceus Engelm., 1857, Proc. Amer. Acad., 3 : 275. 

Echinocactus Wislisenii Engelm. var. LeContei Engelm. in Eothrock, 1878, Eept. U. S. 

Geogr. Surv., 6 : 128. 
Ferocactus LeContei Britt. and EosE, 1922, Cactaceae, 3 : 129. 
Ferocactus acanthodes Britt. and Eose, 1922, loc. cit. 
Ferocactus Bostii Britt. and Eose, 1922, loc. cit., p. 146. 
Echinocactus Eertrichii Weinberg, 1929, Desert, 1 : 40. 
Ferocactus acanthodes var. Bostii W. T. Marshall in Marshall and Bock, 1941, Cactaceae, 

p. 148. 

The common barrel cactus in western and central Arizona does not seem to 
have any points of consistent difference from the type common in the Cali- 
fornia deserts. Previously (Benson et al, 1940) it was treated as a species 
pending further investigation, but with the following statement, "Probably 
merely a form of Echinocactus acanthodes." 



262 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Type collections: (1.) E. acanthodes, type collected in California. (2.) Var. cylindraceus, 
"Found near San Felipe, on the eastern slope of the California Mountains." Colorado 
Desert, San Diego County, Parry, Mo. (3.) E. LeContei, "This gigantic .species was first 
noticed by Dr. John L. LeConte, on the lower Gila, where also Dr. C. C. Parry saw it. . . . 
Subsequently, Dr. Bigelow met with this remarkable plant, abundantly from the Cactus 
Pass, at the headwaters of Williams' River down this stream to the Colorado, and Avest of 
it till E. polycephalus took its place." (4.) F. Bostii, "The tj-pe is based on a plant collected 
in Lower California, 40 miles south of the International Boundary Line (Eost No. 327)." 
(5.) E. ncrtrichii, "Type specimen: Huntington's Botanischer Garten und Museum, San 
Marino, California." "Type — Weinberg in Henry E. Huntington's Botanic Garden, San 
Marino, Calif. Locality — Tortilla and Gila mountains, Arizona." 

Echinocactus polycephalus Engelm. var. xeranthemoides Coulter 

Echinocactus polycephalus Engelm. var. xeranthemoides Coult., 1896, Contr. U. S. Nat. 

Herb., 3 : 385. 
Echinocactus xeranthemoides Engelm. ex Coult., 1896, loc. cit., as syn. 
Echinocactus xeranthemoides Engelm. ex Eydb., 1917, Fl. Eocky Mts. and adj. Plains, p. 579. 

Stems rarely more than 12, the largest ones 1-2 dm. in diameter, the clump 
usually asymmetrial, that is, the tallest stem on the edge ; spines glabrous, 
yellow or pale gray tinged with red or sometimes pink or light red ; central 
spines spreading at right angles to the stem. (In the tj^pical variety: stems 
commonly 10-30, the larger ones 2-3 dm. in diameter, the clump symmetrical, 
that is with the largest stem in the center; spines densely covered with felt 
when young, red ; central spines spreading each in a low" arc.) 

Rocky, south-facing ledges in the Southwestern Coniferous Woodland at 
4,000 to 7,000 feet elevation. Arizona in the northern edge of Mohave County 
and along the Colorado River system from Lee's Ferry and Canyon Diablo 
to the Grand Canyon region ; southern Utah. 

Type collection: "Type, Siler of 1881 and 1883 in Herb. Mo. Bot. Card." 

MAMMILLARIA 

As in Echinocactus and Opuntia, although a number of problems exist, data 
for solution of some of them are inadequate. Only the following changes are 
proposed. 

Mammillaria vivipara (Nutt.) HaAv. var. aggregata (Engelm.), L. Benson, 

comb. nov. 

Mammillaria aggregnia Engelm. in Emory, 1848, Notes Mil. Eeconn., p. 146. 
Cereus aggregatus Coult., 1896, Contr. U. S. Nat. Herb., 3 : 396. 
Echinocereus aggregatus Eydb., 1906, Bull. Torrcy Club, 33 : 146. 
Coryphantha aggregata Britt. and Eose, 1923, Cactaceae, 4 : 47. 

Low-growing plant, the stems at first solitary but later forming mounds 
2-6 dm. high and up to 6 dm. in diameter ; stems 4—5 cm. in diameter at ma- 
turity, tubercles cylindrical, about 8-10 mm. high, about 6 mm. in diameter, 
the surface of the stem obliterated by spines; central spines usually 5-6, white 
tipped with brown, straight, the upper ones turned upward, 1.5-2 cm. long, 
the lower ones spreading at various angles from the stem, about 6-9 mm. long. 



Vol. XXV] BEN SON : BEVI S lOX OF ABIZON A CACTACEAE 263 

m 

tapering", slender but rigid, not flattened, the centrals appearing after the 
radials and not present in yomig stems; radial spines 15-20, similar to the 
centrals, white, slightly brownish at the extreme tips, 12-18 or 25 mm. long, 
somewhat flattened; petals usually lavender or pink, oblanceolate, rather 
abruptly acuminate, 4-6 mm. broad, the flower about 3.5^ cm. in diameter ; 
fruit green with a dull purplish tinge, ellipsoid, about 2.5 em. long, 1.8 mm. 
in diameter. 

River bottoms, the desert floor, or most frequently on grassy plains or rocky 
slopes in the upper part of the Sonoran Desert, the Desert Grassland, and the 
Southwestern Oak Woodland and Chaparral at 3,000 to 5,000 or 6,000 feet 
elevation. The eastern half of southern Arizona; largely south of the Mogollon 
Rim ; New Mexico on the Gila River drainage ; Sonora. Flowering in May 
and June. 

Type collection: "Oct. 18, 1846. Head waters of the Gila, 6,000 feet above tlie sea." 
Interpretations of the type have alternated between this plant and Echinocereus triglocM- 
diatus var. melancanthus. Engelmann, loc. cit., first interpreted the plant as a member of 
the Mammillaria vivipara complex, and (as pointed out by Peebles apud Kearney and 
Peebles, U. S. Dept. Agric. Misc. Publ., (423) : 605, 1942, the original drawing shows 
crossed spirals indicating tubercles on the stem. The type was not preserved, or was lost. 

Mammillaria vivipara var. arizonica (Engelm.) L. Benson, comb, no v. 

Mammillaria arizonica Engelm. in Brew, and Wats., 1876, Bot. Calif., ed. 1, 1: 244. 
Cactus radiosus (Engelm.) Coult. var. arizonica Coult., 1894, Contr. U. S. Nat. Herb., 

3: 121. 
Mammillaria radiosa Engelm. var. arizonica K. Schum., 1898, Gesamtb. Kakteen, p. 481. 
Con-ypliantlia arizonica Britt. and Eose, 1923, Cactaceae, 4: 45. 

Stems solitary at first, later forming great mounds ; spines predominantly 
dark brown; lower central spines 2, one of these spreading perpendicularly 
(to the areole), the other turned downward; petals deep pink, linear-lance- 
olate, gradually acuminate, 2.5-4 mm. broad. 

Rocky places in the Southwestern Coniferous Woodland and the Rocky 
Mountain Forests at 4,000 to 8,000 feet elevation. Southwestern Utah ; north- 
ern Arizona from Mohave County to Apache County and southward to the 
Mogollon Rim and the White Mountains, and perhaps farther in the higher 
mountains. 

Type collection : "On sandy and rocky soil in Northern Arizona, from the Colorado east- 
ward (Coues, Palmer, F. BiscJioff), and into Southern Utah (/. E. Johnson). . ." 

Mammillaria vivipara var. deserti (Engelm.), L. Benson, comb. nov. 

Mammillaria chlorantha 'Enge\m.apud Eothr., in Wheeler, 1878, U. S. Geogr. Surv. W. of 

lOOthMerid., 6: 127. 
Mammillaria deserti Engelm. in S. Wats., 1880, Bot. Calif., 2 : 449. 

Cactus radiosus (Engelm.) Coult. var. deserti Coult., 1894, Contr. U. S. Nat. Herb., 3 : 121. 
Cactus radiosus var. chloranthus Coult., 1894, loc. cit. 
Cactus radiosus var. Alversonii Coult, 1894, loc. cit., p. 122. 
Mammillaria Alversonii Zeissold, 1895, Monatsschr. Kakteenk., 5 : 70. 
Mammillaria radiosa Engelm. var. deserti K. Schum., 1898, Gesamtb. Kakteen, p. 481. 



264 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. rtXH skr. 

Mammillaria radiosa var. chlorantha K. Schum., 1898, loc. cit. 

Mammillaria radiosa var. Alversonii K, ScHUM., 1898, loc. cit. 

Coryphantha chlorantha Britt. and Rose, 1923, Cactaceae, 4: 43. 

Coryphantha arisonica Britt. and Rose, 1923, loc. cit., p. 45. 

Coryphantha deserti Britt. and Rose, 1923, loc. cit., p. 45. 

Mammillaria arisonica Engelm. var. deserti Engelm. ex Davidson and Moxley, 1923, Fl. 

S. Calif., p. 244. 
Mammillaria arisonica var. Alversonii Engelm. ex Davidson and Moxley, 1923, loc. cit. 
Coryphantha Alversonii Orcutt, 1926, Cactography, (3) : 3. 

A robust type with the stems usually solitary or sometimes few, these 7-20 
cm. high, 5-7.5 cm. in diameter ; central spines 3-14, usually white tipped with 
brown, straight, spreading at all angles, 12-18 mm. in length, averaging long, 
tapering, stout, rigid, not flattened; radial spines about 20, similar to the 
centrals but more slender, white ; petals straw-colored, yellow, pink, or purple, 
the flowers 2.5-3.5 cm. in diameter. 

Rocky slopes in the Mojavean Desert at 1,500 to 3,500 feet elevation. Cali- 
fornia in the ranges of the eastern and southern parts of the Mojave Desert; 
southern Nevada; southwestern Utah; northwestern Arizona in northern and 
western Mohave County. A variable plant possibly to be segregated into more 
than one variety with accumulation of additional data. The characters of the 
proposed segregates seem inconsistent in their association. 

Type collections: (1.) M. chlorantha, "Southern Utah, east of Saint George, Dr. Parry. 
I. E. Johnson." (2.) M. deserti, "At Ivanpah, 30 miles northeast of San Bernardino, in one 
of the mountain ranges stretching into the desert, S. B. Parish." Mo. Ivanpah is in the New 
York Mountains, about 140 miles northeast of San Bernardino, California. (3.) Var. 
Alversonii, "Type, Alversou's specimens in the Herb. Mo. Bot. Gard. and Herb. Coulter . . .. 

SOUTHERN CALIFORNIA (A. H. Alverson of 1892) The decidedly pink flowers were 

sent by Mr. S. B. Parish from specimens growing in cultivation in San Diego, and are not 
from the original collection of Mr. Alverson." 

Mammillaria Heyderi Miihlenpf ordt 

Mammillaria Heyderi Muhlenpfordt, 1848, Allg. Gartenz., 16: 20. 

Cactus Heyderi Kuntze, 1891, Rev. Gen. et Sp. PI., 1: 260. 

N eomammillaria Heyderi Britt. and Rose, 1923, Cactaceae, 4: 75. 

Obscure depressed cactus arising only 1-5 cm. above ground level; stem 
solitary, shaped like a turnip, flat-topped, half or two-thirds subterranean, 
5-8 cm. long, 7-12 cm. in diameter, the tubercles conical, 6-10 mm. high, 5-7 
mm. in diameter; juice of the plant milky; central spines 2-3, white tipped 
with dark brown, straight and subequal, 6-8 mm. long, tapering, slender, 
rigid, not flattened; radial spines 10-15; petals pink; sepals not fringed; 
flower 10-12 mm. in diameter. 

Limestone hills in the Chihuahuan Desert or the Desert Grassland at 3,500 
to 5,000 feet elevation. Along the Mexican Border in Cochise County near 
Bisbee {Peebles SF 922, Sac, JJA) and in the Peloncillo Mountains (L. Ben- 
son 10272, TJA, B). One of several Chihuahuan Desert plants barely reaching 
southeastern Arizona and there restricted to favorable limestone soils. 

Type collection : none given. 



Vol. XXV] BENSON: BEVISION OF ARIZONA CACTACEAE 2Go 

Mammillaria Heyderi var. MacDougalii (Rose) L. Benson, comb. nov. 

Mavimillaria MacDougalii Eose in Bailey, 1916, Stand. Cyclop. Hort., ed. 1. 4: 1982. 
Neomammillaria MacDougalii Britt. and Eose, 1923, Caetaceae, 4: 74. 

Stems 5-7.5 or 12.5 cm. long, 7.5-17.5 cm. in diameter; central spines 
straight or the principal one curving gradually outward ; petals cream-colored 
or light yellow ; sepals fringed ; flower about 17-20 mm. in diameter. 

South exposures of rocky hill crests in the Desert Grassland and the South- 
western Oak Woodland at 4,000 to 6,000 feet elevation; often in crevices of 
rocks or in rocky soil. Arizona from the Baboquivari Mountains eastward 
to the Santa Catalina Mountains and to southern Cochise County ; probably in 
adjacent Sonora. Often associated with Echinocereus pectinatus var. rigi- 
dissimus, the Arizona rainbow cactus. Flowering late in March and in April. 

Tj^pe collection : "Common in the mountains about Tucson, Ariz., where it 
was collected by D. T. MacDougal, for whom it is named." 

The dimorphic fruits of some of the typical Mammillaria species, including 
If. Heyderi and the variety MacDougalii have been well-known for some time, 
and Peebles, Cact. and Succ. Jour., 13 : 143-5, 1941, has reported and illus- 
trated similar types (clavate-red and spherical-green) in Mammillaria micro- 
carpa. They occur also in Mammillaria fasciculata (15 miles west of Silver 
Bell, Pima County, Arizona, L. Benson 10606, VA, B). 

The plants described previously as Mammillaria microcarpa Engelm., M. 
fascicidata Engelm., 31. Wilcoxii Tourney, M. viridiflora (Britt. and Rose) 
Boed., and M. Mainae K. Brandegee are in need of further study, but at 
present data are inadequate. 

LITERATURE CITED 

Benson, Lyman 

1939. Notes on taxonomie techniques. Torreya, 39 : 73-75. 

1941. Taxonomie studies. American Journal of Botany, 28: 358-364. 

1942. The relationship of Ranunculus to the North American Floras. American Journal of 
Botany, 29 : 491-500. 

Benson, Lyman, J. J. Thornber, A. A. Nichol, and Lucretia Breazale Hamilton 

1940. The cacti of Arizona. University of Arizona Biological Science Bulletin No. 5. 134 
pp. 52 pi. Tucson, Arizona. 

Peebles, Eobert H. 

1941. Variation in the fruit of Mammillaria microcarpa. Cactus and Succulent Journal, 
13: 143-145. 

1942. Preservation of cactus material. Cactus and Succulent Journal, 14: 3-8. 

Shreve, Forrest 

1936. The plant life of the Sonoran Desert. Scientific Monthly, 42 : 195-213. 
1942. The Desert Vegetation of North America. Botanical Eeview, 8: 195-246. 

Shantz, Homer Leroy and Eafael Zon 

1924. Natural vegetation. Atlas of American Agriculture, U. S. Dept. of Agriculture, 
Washington, D. C. 



EXPLANATION OF PLATE 

Plate 25 

Opunlia and Echinocereus : A, Opuntia, 1, Opitnlia covipressa var. macro- 
rhiza, collected at Pearce, Arizona, by A. A. Nichol in 1940, 2, Opuntia com- 
pressa var. macrorhiza, from sand hills near Guion, Arkansas, Demaree 23490, 
3, Opuntia compressa, from the Atlantic seaboard, received from cultivation in 
Massachusetts; B. Echinocereus Engelmannii var. Xicholii, growing in the 
Ajo Mountains, Arizona. (Photograph A by Eobert A. Darrow; B by Walter 
S. Phillips.) 



[ 2<H' ] ^ 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 10 



[BENSON] PLATE 25 




[267] 



PROCEEDINGS 

OF THE -£ 

CALIFORNIA ACADEMY OF SCIENCES l-^ 

Fourth Series 
Vol. XXV, No. 1 1, pp. 269-290, 4 text figs. November 10, 1944 




ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 11 
ENDEMISM IN CREPIS 

BY 

ERNEST B. BABCOCK 

Professor of Genetics 
University of California ' 

I. INTRODUCTION 

ONE RESULT of the exteiisive investig-ations on the genus Crepis has been the 
formulation of a fairly definite hypothesis concerning the origin, devel- 
opment and migrations of this essentially monophj^letic group of plants. This 
hypothetical history of the one hundred and ninety-six species of Crepis has 
been found to be consistent with a great mass of other evidence concerning the 
origin and migrations of floras in the northern hemisphere. A review of this 
evidence. together with a detailed discussion of the data on Crepis will be 
presented in a monograph which is in press. 

The criteria of phylogenetic relations in Crepis represent five different dis- 
ciplines. The primary basis for classifying all the species into twenty-seven 
sections and arranging the sections in an approximate phyletie series is com- 
parative morphology. Second in importance have been the data on chromo- 
some numbers in one hundred and thirteen of the species (cf. Babcock and 
Jenkins, 1943). Thirdly, the cytogenetic analysis of certain interspecific 
hybrids has confirmed the conclusions reached concerning the phyletie rela- 
tions of the parent species as determined on the basis of comparative morphol- 
ogy and chromosome numbers, and has revealed the evolutionary process by 
which reduction in chromosome number in the series 6-5-4-3 has been accom- 
plished (cf. Babcock, 1942, pp. 176-178; Tobgy, 1943). Fourthly, purely 
genetic data on many interspecific hybrids have substantiated the grouping of 

[ 269 ] 




Fig. 1. Crepis geracioides Hausskn. a, plant, XV2; i, flower-head, x 2 ; c, a floret lacking 
tlic ovary, x 4 ca.; C, trielionie from the corona-tube, x 100 ca.; d, anther-tube slit and opened 
out, x8 ca.; e, detail of anther-appendages, x 32 ca.; f, g, two achenes and a pappus-seta 
from each, x 8 ca.; h, W, two inner involucral bracts showing outer and inner faces respec- 
tively, X 4 ca.; i, haploid set of chromosomes, x 1500. 




Fig. 2. Crepis senecioides Delile. a, b, c and 1-, plants and parts, x i^ ; d, m, florets lacking 
the ovary, x4 ca.; e, n, anther-tubes slit and opened out, x 8 ca.; f, o, details of anther- 
appendages, X 32 ca.; g, h, p, q, aehenes and single pappus-setae, X 8ca.; r, haploid set of 
chromosomes, x 1500. 



272 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

the species into sections on a morphological basis ; and the genetic analysis 
of hybrids has shown the importance of grene mutation as a cause of speeiation 
in Crepis. Fifthly, a study of geographic distribution in the genus as a whole, 
taken section by section, when considered in the light of well-establislied paleo- 
botanical evidence, has led to the formulation of a phylogenetic history of 
the genus. 

In order to give the reader a concrete notion of the morphological contrasts 
between the most primitive and most advanced types of Crepis species, one of 
each is illustrated (figs. 1 and 2). The differences in size and habit of the 
plant, in size of the heads, florets, fruits, etc., are so obvious, it seems hardly 
necessary to point them out. The primitive species, C. geracioides, is a montane 
perennial which is endemic in the southern Balkan Peninsula. The advanced 
species, C. senecioides, is a precocious, desert annual which has been collected 
at a number of stations scattered along the Egyptian-Tunisian littoral. Be- 
cause of its wide distribution it is not considered to be an endemic species in 
the restricted sense in which that term is used in the present paper. But, since 
it is the most advanced species in the genus on morphological grounds, it has 
been used to illustrate the extreme reduction in size which has accompanied 
development of the annual habit, extreme precocity, and remarkable longevity 
of the tiny seeds (achenes) , all of which adapt it so well to desert conditions. 
The fact that these two species, which represent the earliest and latest stages 
in a very long developmental history, now exist on opposite sides of the Medi- 
terranean Sea, which is only some 600 kilometers (400 miles) wide in this 
region, is sufficient to pique one's curiosity as to whence and how they came 
to be there. 

Stated as briefly as possible, the answer is as follows. Crepis originated in 
Central Asia early in the Tertiary period. Some, at least, of the most primitive 
species existed at low elevations, under mesophytic conditions. But migration 
into higher altitudes, accompanied by speeiation, must have occurred during 
the early development of the genus and before the extensive migrations took 
place to the northeast, southeast, northwest and southwest from Central Asia. 
The migrations to the southwest affected a much larger number of Crepis 
species than the other three migrations. The general trend of the southwesterly 
migration passed through Turkestan and Iran and then followed one of three 
main routes: (1) through northern Asia Minor into the Balkan Peninsula 
and southern Europe; (2) through southern Asia Minor, Iraq, Syria and 
Palestine into Egypt; and (3) through southern Arabia into Abj^ssinia and 
tropical Africa. These migrations were accompanied or followed by increas- 
ing dessication in southwestern Asia and the Mediterranean region and this, 
together with the many topographical changes Avhich occurred, resulted in 
increased speeiation throughout that general region. Thus C. geracioides 
almost certainly migrated from Central Asia around the south end of the 
Caspian Sea, through northern Asia Minor, into the Balkan Peninsula where 
it still exists in tliree localities. But C. senecioides almost certainly developed 



Vol. XXV] BABCOCK: ENDEMISM IN CEEPIS 273 

where it now exists from some less advanced ancestor which was unable to 
tolerate the increasing aridity of the climate. That ancestor, in turn, probably 
developed from a still more primitive type somewhere in the Iran-Iraq region. 
A presentation of the general background of evidence for these rather dog- 
matic statements must await the publication of the monograph. 

One of the most interesting aspects of the geographic distribution of Crepis 
is the large amount of endemism present and the distribution of the endemic 
areas. It is the purpose of the present paper to examine this evidence in order 
to determine to what extent it agrees with the phyletic history briefly outlined 
above, which was worked out for the genus as a whole. At the same time it may 
be possible to derive some conclusions concerning endemism in Crepis which 
will have general significance. 

The present study is confined to the Old World species inasmuch as the 
data on endemism in the New World species have already been reported (Bab- 
cock and Stebbins, 1938). With reference to the history of the genus as a 
whole, the conclusion was reached that all of the American species either 
migrated from Asia across the Beringian land-bridge before the Pleistocene 
glaciation could prevent such migrations, or that they were derived from 
species of Asiatic origin. 

The term endemic, in its usual biological connotation, is applied to a plant 
or animal which is restricted in its distribution to a single country, region or 
area. This definition becomes much more definite as soon as the size of the 
region or area is stated. During the years spent in accumulating data of vari- 
ous kinds concerning Crepis species, the writer came to realize that a great 
many of them are endemic in the general sense defined above ; and that a con- 
siderable number of them are also of very restricted distribution. Some of 
these restricted species are insular, being confined to one or other of such 
islands as Crete, Sicily, Majorca, Madeira or Fuerteventura and Lanzarote 
of the Canary Archipelago. Others are restricted to a single limited district 
on a continent, as in the case of the 22-chromosome forms of C. monticola and 
C. pleurocarpa, two of the oldest endemic American species of Crepis (cf. 
Babcock and Stebbins, 1938) . Still others are similarly restricted by ecological 
factors, but they occur in two or more small districts and these may be either 
near together or widely separated within a given endemic area. An example of 
the latter is C. geracioides; but the three localities at which C. geracioides is 
known to occur are separated by a maximum distance of approximately 150 
kilometers (90 miles) . Hence this species can hardly be described as a narrow 
endemic except in a purely relative sense, as when compared with such a wide- 
spread species as C. senecioides. 

In order to obtain more definite information as to the amount and possible 
significance of endemism in Crepis, it was decided to set arbitrarily a lower 
limit of fifty kilometers (30 miles) and an upper limit of 150 kilometers (90 
miles) ; and to classify all the species whose maximum straight-line distance 
between known localities falls within the lower limit as "narrow A" ; all those 



274 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

whose maximum distance between known localities exceeds the upper limit as 
"wider" ; while those falling between the two limits were classed as "narrow 
B." It was found that there were so few species in the "narrow B" group that, 
for the sake of simplicity, the "narrow^ B" and "wider" groups were lumped 
together, thus recognizing only two categories, "narrow" and "wider," with 
the dividing line at 50 kilometers. 

On this basis it was found that there are forty-eight species of Crepis for 
which the available data indicate a "narrow" distribution. Twenty-eight of 
these are known from only one locality' while the other twenty are known from 
two or more localities. Of the twenty-eight that are known from a single lo- 
cality, twentj'-f our are known to the writer from just one collection ; but the 
other four are known to have been collected several times at the one locality. 
Three of these four localities are mountain peaks in Greece ; the other is the 
eastern promontory of Madeira Island. The twenty-four localities at which 
only a single collection has been made are mostly in regions w^hich have been 
less thoroughly explored botanically, such as Asia Minor, the Caucasus, 
Turkestan and tropical Africa. Many of these localities are in mountainous 
countrj'^ and some are mountain peaks. Since many of the "wider" endemics 
are known to be very local in their distribution, and half of the "narrow" ones 
are known from two or more collections, it is fairly probable that the majority 
of the species known from only one collection will turn out CA'cntually to be 
actually "narrow" as here defined. At any rate the purposes of the present 
paper can be served better by recognizing them as "narrow" than by ignor- 
ing them. 

Classification of the remainder of the Crepis species as endemic or not en- 
demic was accomplished by defining a series of endemic areas and then by 
accepting as endemic only those known to be restricted to a single area. Many 
of these areas are widely recognized as rich in endemic species of plants. By 
this method ninetj^-one species were selected as "wider" endemics. This does 
not mean that these species are continuously distributed over a district more 
than 150 km. (90 miles) in diameter. On the contrary these "wider" endemics 
are all more or less local in their distribution ; although a few are fairly 
abundant in certain districts. Thus in the case of C. geracioides it happens 
that the three known localities are about 150 kilometers distant from one an- 
other ; whereas C. terglouensis, another very primitive endemic, is known from 
at least twenty high alpine stations ranging from south-central Switzerland 
to central Tirol and Upper Austria. In marked contrast with these are such 
insular species as C. cretica and C. canariensis which are abundant on the 
islands where they exist. A few species have been omitted which occur mostly 
in one endemic area but are also represented in another and hence, on a less 
restricted criterion, might have been considered as endemic. Interesting ex- 
amples are C. pygmaea, a very primitive 12-chromosome species of Spain, the 
Pyrenees, the southwestern Alps and northern Italy; C. smyrnaea, another 
12-chromosome primitive species known from only three localities in southern 



YoL.XXV] BABCOCK: EXDEMISM IN CEEPIS 275 

Greece and western Asia Minor; C. pontana, the most primitive 10-chromo- 
some species of Crepis, which occurs from the eastern Alps to the western part 
of the Balkan Peninsula ; and C. albida, the next most primitive 10-chromo- 
some species which is polymorphic and is distributed from the Grand Atlas 
Mts., in Morocco through Spain, the mountains of southern France and the 
Maritime Alps. It becomes difficult to know where to draw the line, if one 
begins to go beyond the limits of a single endemic area. But, on this restricted 
basis, there are still one hundred and thirty-three endemic species which is 
72 per cent of the one hundred and eighty-five Old World species. 

II. THE DISTRIBUTION OF THE ENDEMIC 
REGIONS AND AREAS FOR CREPIS 

The endemic regions and areas for Crepis are defined in Table 1 and shown 
on the accompanying map (fig. 3). On this map the large size of the tropical 
African area (no. 19) will be one of the first things to catch the attention. It 
should be emphasized that, in the latter area, all of the eighteen endemic 
species found there are extremely local in distribution. All but three occur 
on only one or two mountains or in a single mountain range. These mountains 
and ranges are shown as solid circles or narrow ellipses on the map. The other 
three species are knoAvn from only single stations which are at low elevation 
and these are shown as solid squares on the map. Following is a key to the 
symbols used to designate these locations in area No. 19 and three others in 
area No. 10. 

The long winged-shaped ellipse in area No. 10 is the Pamir-Ferghana-Altai 
system of mountains. 

KEY TO MAP SYMBOLS 

Regions are indicated by Roman numerals. Areas are 
indicated by Arabic numerals. 

Mountains shown hy solid circles and ellipses: 

AT. Ala-tau Mts., Central Asia. 

CH. Mt. Chirinda, southeastern Africa. 

DA. Darvas Mts., eastern Kussian Turkestan. 

CU. Congo-Uganda plateau north of Lake Kiwu, Africa. 

HI. Himalaya Mts., Kashmir region, high alpine. 

KE. Mt. Kenya, eastern tropical Africa. 

KI. Mt. Kilimanjaro, eastern tropical Africa. 

KU. Mountains south of Lake Ki'\vu, tropical Africa. 

MC. Mt. Cameroon, western tropical Africa. 

ME. Mt. Meru, adjacent to Kilimanjaro. 

NY. Mountains of Nyasaland and adjacent Congo. 

EM. Mt. Kungwe and Mt. Mbeya, tropical Africa. 

UK. Urundi Province, Kisozi, tropical Africa. 

UL. Uluguru Mts., Tanganyika Prov., Africa. 

UT. Ulu-tau Mts., northern Eussian Turkestan. 



176 



CALIFOBXIA ACADEMY OF SCIEXCES [Proo. 4th ser. 



Single lowland stations in Africa: 

EC. Elizabethville, southeastern Belgian Congo. 
FU. Fort Portal, Uganda. 
CA. Cului, Angola. 

TABLE 1 
Endemic Regions and Areas for Old World Crepis 



Hegions 
I. Southwestern Europe; Alps; north- 
western Morocco 

II. Balkan Peninsula 
III. Asia Minor ; Syria-Palestine ; Cri- 
mea 

IV. Caucasus ; Iran 

V. Turkestan-Pamir-Altai region 

VI. Northwestern India 

VII. Southeastern Asia 

VIII. Eastern Siberia 
IX. Northern Japan 
X. Abyssinia ; Eritrea ; southwestern 
Arabia 

XI. Africa, tropical 
XII. Lybia- Morocco -Cadiz (littoral) ; 
southern and western maritime 
Italy and southern France; Ma- 
deira and Canary Islands 



Areas 

1. S. Spain and N.W. Morocco 

2. N.E. Spain and "W. France 

3. European Alps 

4. Balkan Peninsula 

5. Asia Minor 

6. Syria and Palestine 

7. Crimea 

8. Caucasus region 

9. Iran 

10. Turkestan (E. and N. Eussian) and the 
Pamir-Altai mountains 

11. N.AV. Himalaya Mountains, high alpine 

12. N.W. India-Baluchistan, low montane 

13. S.E. Asia (= S.E. Tibet, S.W. China, N. 
Burma, Laos, Annam) 

14. E. Siberia (Okhotsk district) 

15. Hokkaido-Sakhalin 

16. S.W. Arabia in Yemen Province 

17. Eritrea 

18. N. Abyssinia, mountains 

19. Africa, tropical (isolated localities) 

20. Lybia (littoral) 

21. Tunisia (middle and southern) 

22. E. Algeria (littoral) 

23. S. Italy and Toscana, Sicily, Sardinia, 
Corsica ; S. France 

24. W. Algeria, N. Morocco, Cadiz (mostly 
littoral) 

25. Madeira and Canary Islands 



Tlie distribution of the twelve endemic regions defines the east-west distri- 
bution of the genus as a whole almost completel}', the gaps between the en- 
demic regions being occupied more or less by the various widespread spet^ies. 
In this connection it should be repeated that the total evidence on geographic 
distribution in Crepis, considered in relation to the evidence on phylogeny, 
definitely indicates that the center of origin and early development of Crepis 
was in Central Asia, in the Altai-Tian Shan mountain region in the north- 
eastern part of region V (fig. 3) ; and that the present wide distribution of 
the genus was accomplished through migration in four general directions 
from the center. One of these trends was towards the northwest, into what is 
now Russia. Only five species exist today in northern Europe as a result of that 



Vol. XX Y] 



BABCOCK : ENDEMISM IN CBEPIS 



277 



trend of mig-ration and none of these is endemic. Another migration trend 
was to the east and northeast, resulting in two endemics in Japan, one in the 
Okhotsk district of Siberia, (region VIII, fig. 3), and all of the American 
species. Still another trend was to the southeast, resulting in the nine endemic 
species found in region VII. Most important of all the migration trends from 
Central Asia, in terms of the number of species involved, was the southwest- 




Fig. 3. Map of the Old World showing the endemic regions (Eoman numerals) and areas 
(Arabic numerals) for Crepis, (In regions V and XI solid circles indicate single mountains 
and ellipses, mountain ranges or systems; solid squares indicate single lowland stations. 
See key to symbols in text.) 

erly trend across Turkestan and Iran. From that region the three branches 
already mentioned led to northwest and tropical Africa; to the Caucasus, 
Asia Minor and southern Europe; and to the eastern Mediterranean and 
North African littoral. The point to be emphasized here is that the distribu- 
tion of the endemic regions for Crepis is entirely consistent with the generic 
history briefly outlined above. 

III. PHYLOGENY AND ENDEMISM IN CREPIS 

Based on the evidence from comparative morphology and cytogenetics it 
has been possible to classify all the species of Crepis on a relative phylogenetic 
basis with some degree of assurance. For purposes of the present discussion 
they have been grouped into three classes, the more primitive, the interme- 
diate, and the more advanced species. In Table 2 are shown the relations of 
the endemic species in these phylogenetic groups to the size of area occupied 



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VoL.XXV] BABCOCK: ENDEMISM IN CBEPIS 279 

by each species (narrow or wider), to its altitude class (alpine, montane or 
low), its moisture classification (as occupying a moist, sub humid or arid en- 
vironment), its life-duration class (perennial or annual-biennial), and its 
variability class (monomorpliic or polymorphic). 

Considering first the totals in the right hand column, it will be noted that 
twenty-nine of the endemic species are primitive and twentj^'-nine are ad- 
vanced, whereas seventy-five, or more than one-half of them, are intermediate. 
Evidently Crepis may be described as a conservative genus and this becomes 
more patent when it is realized that of the twenty-nine advanced endemics 
only some half-dozen desert or montane annuals can be considered as very 
highly specialized species. There is apparently no correlation between the 
phj^logenetic grouping and extent of distribution (narrow vs. wider), since 
about one-third of all the endemics are narrow and two thirds are wider and 
these same proportions hold roughly for each of the three phylogenetic classes. 
In altitudinal distribution there is positive correlation between primitive type 
and alpine altitude and very strong positive correlation between advanced 
type and low altitude. In the moisture classes we find that, of the one hundred 
and four endemics in the primitive and intermediate groups taken together, 
just one-fifth occur under arid conditions, M'hereas four-fifths of the advanced 
endemics occur under arid conditions. Hence there is strong positive correla- 
tion between advanced type and arid environment. In the life-duration classes 
the differences are most striking — all the primitive and intermediate endemics 
are perennial whereas nearly all of the advanced endemics are annual or (in a 
few cases) biennial. In regard to variability there is a stronger tendency 
towards polymorphism in the advanced than in the intermediate or primitive 
endemics. All of the primitive and intermediate endemics are perennial species 
and more than four-fifths of them are monomorphic and occur at montane or 
alpine elevations and under suhhumid or moist conditions. Most of the ad- 
vanced endemics are annual species and two-fifths of them are polymorphic; 
none occur at alpine elevations and about three-fourths of them are found at 
low elevations and under arid conditions. 

Having noted the lack of correlation between extent of distribution (as 
expressed by the arbitrary classes "narrow" and "wider" (and the three 
phylogenetic groups, it may be of interest to examine the relations between 
extent of distribution and the other four classifications of endemic Crepis 
species. In Table 3, in order to reveal the relations involved more clearly, the 
actual numbers, given in the upper two rows, are expressed in the lower two 
rows as percentages of the respective totals, given in the right-hand column. 
The totals given in the middle row are merely for the purpose of checking 
against the other tables. Comparing these percentages, it is clear that there is 
little or no positive correlation between narrow and wider distribution and 
phylogenetic grouping. But it is equally clear that there is positive correlation 
of wider distribution with low altitude and arid climate. As for duration of 
life and variability, about four-fifths of both the narrow and the wider en- 



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VoL.XXV] BABCOCE: EXDEMISM IN CEEPIS 281 

demies are perennial ; wliereas all of the narrow endemics are monomorphic, 
but one-third of the wider endemics are polymorphic. 

Considering- these facts in the light of the conclusions derived from Table 2, 
it may be inferred that the advanced, annual endemic species which are poly- 
morphic are mostly of wider distribution and occur at low altitude, under arid 
conditions. This inference has been verified by reexamination of the original 
data. There are twenty -five annual endemic species (Table 2) of which twenty- 
three occur in an arid climate and all but one of these at low altitude. Only 
eighteen are known to have a wider distribution ; but, of the other seven, six 
are known as yet only from the type locality and further collections may show 
that some of these also have a wider distribution. 

The contrast between this group of advanced endemics and the strictly al- 
pine endemics is most striking. Of the thirty-one alpine endemics, all are 
perennial ; twenty-nine are monomorphic and only two, polymorphic ; while 
all but one occur under subhumid or moist conditions. Six of them are primi- 
tive, aiid twenty -five intermediate — there are no advanced alpine species. This 
means, of course, that the alpine endemics are all relatively old species and 
adapted to mesophytic conditions; whereas the lowland endemics of advanced 
type are relatively young species and adapted to xerophytic conditions. 

But the contrast between the two gToups does not hold for size of distribu- 
tional area. Of the thirty-one alpine endemics twenty-one are wider and only 
ten narrow in distribution. Furthermore, there are ten other alpine species of 
Crepis which are so extensive in geographic distribution that they are not 
considered to be endemic. For example, C. conyzaefolia is a primitive alpine- 
subalpine species distributed from the Pyrenees to the Balkan Peninsula and 
in northern Asia Minor, Transcaucasia, northern Persia and the Altai region. 
Such evidence, together with the prevalence of wide distribution among the 
alpine endemics, certainly indicates that the alpine endemics were formerly 
more widely distributed and have become restricted to their present locations 
presumably through radical changes in the environment. 

The evidence on chromosome numbers is in good agreement with the forego- 
ing generalizations. In Table 4 fifty -three diploid endemic species are classified 
opposite their numbers 6, 5, 4, or 3, and in relation to phylogenetic grouping- 
and the other classifications used in the preceding tables. Phylogenetically, 
the 6-paired endemics are all primitive or intermediate, none are advanced ; 
whereas tlie 5-paired endemics are about evenly divided among the three 
groups and the 4-paired endemics are mostly intermediate or advanced. The 
one 3-paired endemic, C. fuUginosa, is advanced, widely distributed, montane, 
xerophytic, annual, and polymorphic. In distribution all of the 6-paired en- 
demies are wider as well as most of the 5's and 4's ; but the small number of 
narrow endemics that have been examined eytologically may be partly due to 
difficulties in obtaining them in living condition. The data on altitudinal dis- 
tribution indicate no correlations except in the 4-paired species which are 
mostly of low elevation. As for moisture relations, most of the fifty-three 



282 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. 4th ser. 

species are about equally divided between subliumid and arid and this holds 
for each of the chromosome number classes. Under duration of life we find 
that all of the 6-paired endemics are perennial, whereas about one-third of 
both 5's and 4's are annual. Obviously reduction in length of life has gone 
along with reduction in the number of chromosomes. As for variability, there 
is little difference between the chromosome number classes in the proportion 
of monomorphic and polymorphic species. The most significant facts derived 
from this analysis are : (1) The 6-paired species are mostly primitive and all 
perennial, hut they are about equally distributed among the altitude and mois- 
ture classes. (2) The 4-paired species are mostly intermediate or advanced 
and of low elevation, but they are about equally divided between mesophytic 
and xerophytic environments. (3) These facts seem to indicate that adapta- 
tion from mesophytic to xerophytic cofiditions has been going on in this genus 
over a very long period of time. 

IV. THE DISTRIBUTION AND PHYLOGENY OF 
CREPIS ENDEMICS 

In Table 5 each of the endemic Crepis species is tabulated according to th< 
endemic region in which it occurs and its phyletic, altitude and life-duratioi 
class ; also the fifty-three endemics that have had their chromosomes countec 
are tabulated according to chromosome number as well as the region where 
they occur. 

Considering first the primitive endemics, it will be noted that two grades 
are recognized, A and B. The thirteen species in the A group are definitely 
more primitive than those of the B group on morphological grounds. Eleven| 
of these most primitive endemics are either alpine or montane, and restricte( 
to southwestern Europe or the Balkan Peninsula or to a few high mountains| 
in tropical Africa. The other two are alpine species, one in the western Hima- 
laya Mountains and the other in the northern islands of Japan. Thus the most 
primitive Crepis endemics are now mostly distributed at great distances froi 
the assumed center of origin for the genus. Since most of the advanced en^ 
demies are closer to Central Asia than southwestern Europe, the distributionj 
of the most primitive and most advanced endemics in Crepis conforms ii 
general with Matthew's principle of radial distribution of older types from 
common center accompanied by the development of more advanced types 
nearer the center (IMattliew, 1915). This conformity with Matthew's principle 
is also apparent from the distribution of all the species (including the en-| 
demies) in several of the more primitive sections of Crepis. These facts provide 
a sound basis for the hypothetical history of Crepis which starts with the 
origin and early development of the genus in Central Asia. 

Considering next the totals for the twelve regions (Table 5. left-hand coin 
umn), it is clear that the greatest concentrations of endemics exist in thai 
Balkan Peninsula, in Asia Minor and adjacent areas, in tropical Africa, and 
in the Mediterranean littoral. But these four regions differ as to the phylo- 



VoL.XXV] BABCOCK: ENDEMISM IN CBEPIS 283 

genetic status of their endemics. In the Balkan Peninsula close to one-half of 
the endemics are intermediate, whereas, one-fourth are primitive (A+B) and 
one-fourth, advanced. These twenty-six species are similarly divided between 
montane, alpine and low elevations ; the seven advanced species are all annuals. 
Evidently the highly diversified conditions in that region have favored the 
persistence of both primitive and intermediate endemics as well as the develop- 
ment of advanced endemics. In Asia Minor and the Mediterranean littoral, 
however, the endemics are all either intermediate or advanced. But the fact 
that there are no primitive endemics in Asia Minor is probably due to the gen- 
erally more arid climate of that region as compared with the Balkan Peninsula 
as a whole. In tropical Africa, on the other hand, we find just the opposite 
situation. The eighteen endemics are all either intermediate or primitive ; and 
this is also true of four other tropical African species which are so widely dis- 
tributed that they are not included here as endemics. This absence of advanced 
species in tropical Africa probably indicates that the more uniform and favor- 
able conditions of that region have favored the persistence of more primitive 
types without inducing the development of very advanced types. 

Compared with the four regions that are richest in endemics, the following 
regions have only one-third to one-half as many in each region : southwestern 
Europe, the Caucasus-Iran region, Abyssinia and adjacent small areas, and 
southeastern Asia. One of these, the Abyssinian region, differs from the other 
four in having all the phyletic groups represented. The t^n endemics of the 
Abyssinian region are all montane except the two primitive species which are 
alpine ; and yet four of these montane species are advanced. It will be recalled, 
however, that certain other advanced species are montane. This is true of the 
one 3-paired endemic, C. ftiliginosa, which occurs in southern Greece, and of 
its 4-paired close relative, C. cretica. The three other regions contain only 
perennial endemics, none of w^hich is advanced, some being intermediate and 
some primitive. Southwestern Europe and tropical Africa are important 
regions for the most primitive endemics since nine of the twenty-six endemics 
found in those two regions are classed as primitive-A. Thus, at the western 
and southwestern extremes of distribution for Crepis, we find the greatest 
concentration of the most primitive endemics in the genus. 

In order to give another picture of the morphological contrasts between 
these most primitive endemics and the most advanced endemics in the genus, 
C. tergloucnsis, an alpine species found only in the European Alps, is shown 
in fig. 4, a-f, and C. ftiliginosa, a lower montane species of southern Greece, 
in fig. 4, a'-f '. The plants and their parts are all drawn at comparable scales. 
It will be noted that, in their haploid sets of chromosomes, the two species rep- 
resent the extremes of variation found in Crepis in both number and size of 
the chromosomes. 

With further reference to the chromosomes of fifty-three of the endemic 
species (Table 5) , it is clear that the 6-paired endemics are restricted to regions 
I to IV except for one species, C. hashmirica, of northwestern India. In fact 






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Vol. XXV] 



BABCOCK: EXDEMISM IX CEEPIS 



285 



rffT{\ 



0)7Ur 




Fig. 4. a-/, Crepis ierglouensis (Hacq.) A. Kern, a, plant, x % ; l», floret lacking the 
ovary, x3; c, anther-tube slit and opened out, x6; cZ, detail of anther-appendages, x24; 
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above. 



two-thirds of them occur in the mountains of southwestern Europe and the 
Balkan Peninsula. But the 5-paired endemics are widely distributed, and the 
4-paired species still more widely, with a strong representation in the Medi- 
terranean littoral. The evidence from chromosome numbers is consistent with 
the morphological evidence in showing that the oldest endemics are mostly in 
the mountains of southern Europe whereas the j^oungest endemics are mostlj^ 
in the Mediterranean littoral and closely adjacent areas. This evidence from 



286 CiLIFOHXIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

chromosome numbers supports the general hypothesis, based on morphology 
and geographic distribution in the genus as a whole, concerning the origin 
and distribution of the genus. This hypothesis includes the assumption that 
two of the important causes of increased speciation in the genus were the 
climatic and topographic changes which occurred in southwestern Asia and 
the Mediterranean region during late Tertiary and early Quarternary time. 
In Crepis the endemic areas most affected by tliese changing conditions are 
lto9, 21,22and24(cf.fig. 3). 

V. THE GENERAL SIGNIFICANCE OF ENDEMISM IN 

CREPIS 

In addition to the support found in this study of endemism in Crepis for 
the hypothetical history of the genus which had been previously formulated, 
it appears that some of the evidence on Crepis endemics has a bearing on im- 
portant problems of plant distribution. One such problem concerns the so- 
called age and area hypothesis (Willis, 1940) which assumes that the age of 
a species or group is proportional to the area it occupies. 

The "age and area" concept. From the foregoing review it is clear that 
among the endemic species of Crepis there are two extremely diverse groups : 
(1) ihe primitive, perennial, alpine endemics which are relatively old, mono- 
morphic, and adapted to mesophytic conditions ; and (2) the advanced, annual 
lowland endemics M^hich are relatively young, more highly variable, and 
adapted to xerophytic conditions. 

Now the present distribution of these two groups does not conform to the 
"age and area" concept. The old, alpine endemics, however, are believed by 
most students of floristics to have been more widespread before the Pleistocene 
epoch and to have moved into the narrow niches that they now occupy from 
glacial refugia (Braun-Blanquet, 1923, pp. 260-261). The fact that many of 
them are of relatively wide distribution, although they exist only in a special 
type of environment, supports this idea. On the other hand, the young, low- 
land, annual endemics are just as widely distributed as the alpine species are 
at present; and there are a number of lowland, annual species, not endemics, 
for example, C. joetida and C. pulchra, which are about as widely distributed 
as the forerunners of the alpine endemics ever could have been. Hence the 
"age and area" concept breaks down completelj^ with respect to these tM'o 
groups of endemic species. As has been pointed out by Stebbins (1942) the 
concepts of "senescence" and "competition" are no more satisfactory than 
"age and area" in attempting to account for such diverse groups of species. 
But the concept that the monomorphic, alpine species are genetically homo- 
geneous whereas the variable lowland species are genetically heterogeneous, 
i.e., composed of several or many biotypes, provides an acceptable explanation 
of their differences in variability and distribution. 

The nature of the "narrow" Crepis endemics. This genetic approach to the 
problem of the differences between the widely distributed alpine and low- 



YoL.XXY] BABCOCK: ENDEMISM IX CEEPIS 287 

laud endemics also provides a satisfactory explanation of the nature of the 
"narrow" Crepis endemics. Of the forty-eig'ht Old World species which were 
classified as "narrow" many are known only from their type locality or else 
the information about their distribution is vague ; but there are several con- 
cerning which it is possible to make fairly definite statements. These are listed 
below in their phylogenetic groups, together with pertinent data. 

Narrow Endemics — All Presumably Monomorphic 

Primitive 

1. C. Mlimandscliarica O. Hoffm. Section 8. Known only from Mt. Kilimanjaro at four 
stations, 2700-3500 meters, in forest. 

2. C. cameroonica Babe. Section 8. Known with certainty only from Mt. Cameroon at three 
stations, 2000-4000 meters, in meadows. 

Intermediate 

3. C. taygetica Babe. Section 10. Known from five collections, all made at a place called 
"Porta" at about 2300 meters altitude on Mt. St. Elias in the Tavgetus Mts., of southern 
Greece. Exposed rocks. 

4. C. crocifolia Boiss. et Heldr. Section 10. Knol^^l from three or four collections made at a 
place called Megala Zonaria, at about 2400 meters altitude, on the same mountain as 
the preceding. Exposed rocks. 

5. C. athoa Boiss. Section 10. Seven collections, all on Mt. Athos, Hagion Oros Peninsula, 
northern Greece. Exposed rocks. 

Advanced 

6. C. eritreensis Babe. Section 20. Three or four localities in southern Eritrea from 800 
to 1400 meters elevation. One collection was from "a comparatively moist, rocky for- 
mation." 

7. C. tyhalciensis Vierh. Section 20. Known from one specimen, collected in southern Crete. 
Since Crete has been botanized by numerous collectors, it is very probable that this 
species would be represented by more collections if it were widely distributed on the 
island. Certainly insular. 

8. C. divaricata (Lowe) F. Schultz. Section 25. Known with certainty from four collections 
made on the isolated eastern promontory of Madeira Island where it has been nearly 
exterminated by grazing. (Eeported in 1837-1850 from the tiny Desertas Islands near 
Madeira.) Certainly insular. 

9. C. Noronhaea Babe. Section 25. Six collections, all from Porto Santo, a small island in 
the Madeira Archipelago. Certainly insular. 

10. C. Forskalii Babe. Section 27. Two localities in Yemen Province, southwestern Arabia, 
between 1300 and 2500 meters elevation, in the coffee belt. 

It is noteworthy that three of these species, C. divaricata, C. Noronhaea and 
C. tyhakiensis, are insular and of very restricted distribution. However, they 
are relatively advanced species. Therefore their morphological homogeneity 
(assuming that C. tyhakiensis is actually monomorphic) must be due to genetic 
homogeneity rather than to senescence. Five others in the above list, C. kili- 
mandscJiarica, C. cameroonica, C. tay'getica, C. crocifolia and C. athoa, are each 
found on only a single mountain. They are almost certainly "depleted" species 
(Stebbins, 1942) and they probably became isolated through widespread 



288 CALIFORNIA ACADEMY OF SCIENCES [Proc. -Ith ser. 

changes in topography and climate during late Tertiary and Pleistocene times. 
In these also the population must be relatively small and homogeneous. The 
other two species may also be "depleted" species, although it must be admitted 
that the available information on their distribution and variability is scanty. 

SUMMARY 

1. The twelve endemic regions for Crepis, comprising twenty-five areas, 
define the east -west distribution of the genus almost completely. 

2. Seventy -two per cent of the Old World Crepis species are endemic in the , 
sense that thej^ are restricted to a single subcontinental geographic area. 
Among these endemic species are two very diverse groups, namely the most 
primitive, perennial species, half of which are alpine relics, and the advanced,! 
annual species, most of which occur at low altitudes under arid conditions. 
The remaining endemics comprise an intermediate series connecting these tw( 
extremes. But comparatively few of the alpine perennials and of the lowlanc 
annuals have narrow distributions, i.e., their distributional areas, as at present 
known, are mostly over 150 kilometers in extent. From this evidence it is in-| 
f erred that the alpine relics were formerly of wider distribution and that thej 
moved into their present niches from glacial refugia ; whereas the lowland, 
annual endemics have become adapted to a more arid climate but under con- 
ditions favoring genetic heterogeneity. From the evidence on morphology and 
distribution as well as on chromosome numbers it is inferred that the process 
of adaptation from mesophytic to xerophytic conditions went on in the genus 
as a whole over a long period of time. 

3. The distribution of the most primitive and most advanced endemic species 
of Crepis supports the conclusion, derived from the evidence on phylogeny 
and distribution of the genus as a whole, that the center of origin was in Cen- 
tral Asia. 

4. The "age and area" concept of Willis breaks down completelj^ in the light 
of the evidence on the two diverse groups of endemic species mentioned above. 

5. The concept of "senescence" fails to explain the existence of narrow en- 
demics in Crepis since this group includes some of the most primitive and 
most advanced species in the genus. Since they are either insular or terrestrial, 
"depleted" species, the population is more or less restricted in size and rela- 
tively homogeneous. 



VoL.XXV] BABCOCE: ENDEMISM IN CBEPIS 289 

- REFERENCES 
Babcock, E. B. 

1942. Systematies, cytogenetics and evolution in Crepis. Bot. Eev., 8:139-190. 

Babcock, E. B., and G. L. Stebbins, Jr. 

1938. The American Species of Crepis. Carnegie Inst. Wash., Pub. No. 504:1-199. 

Babcock, E. B., and J. A. Jenkins. 

1943. Chromosomes and phylogeny in Crepis III. The relations of one hundred and thir- 
teen species. Univ. Calif. Publ. Bot., 18:241-292. 

Braun-Blanquet, J. 

1923. tyber die Genesis der Alpenflora. Verh. Naturf. Ges. Basel, 35:243-261. 

Matthew, W. D. 

1915. Climate and Evolution. Ann. N. Y. Acad. Sci., 24:171-318. 

Stebbins, G. L., Jr. 

1942. The genetic approach to problems of rare and endemic species. Madrono, 6:241-258. 

TOBGY, H. A. 

1943. A cytological study of Crepis fuliginosa, C. neglecta, and their Fj hybrid, and its 
bearing on the mechanism of phylogenetic reduction in chromosome number. Jour. 
Genetics, 45 : 6 7-1 11. 

Willis, J. C. 

1940. The Course of Evolution. 207 pp. Cambridge University Press. 



PROCEEDINGS 
OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 



;««r^ t 







Vol. XXV, No. 12, pp. 291-306, pis. 26-27 



November 10, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 12 

NICOTIANA ARENTSII— A NEW, NATURALLY 
OCCURRING AMPHIDIPLOID, SPECIES 

BY 

T. H. GOODSPEED 

Department of Botany 
University of California* 



IN 1917, Winge pointed out that the arithmetic progression observed in the 
chromosome numbers of some genera might be the product of interspecific 
hybridization followed by chromosome doubling. Eight years later, Clausen 
and Goodspeed showed that such doubling had occurred in a fertile F^ hybrid 
between Nicotiana glutinosa and N. Tdbacum. This initial verification of 
Winge's hypothesis served to direct interest toward the incidence, causes and 
artificial induction of amphidiploidy (cf. Goodspeed and Bradley, 1942). It 
also led to an evaluation of the applicability of Winge's hypothesis in pictur- 
ing evolutionary relationships in the genus Nicotiana. 

Fundamentally, Nicotiana is a New World genus and, with the exception of 
5 species showing 9 or 10 pairs of chromosomes, all its North and South Ameri- 
can species possess either 12 or 24 pairs. Since there is no evidence that any of 
the 24-paired species is autopolyploid, it has been assumed that they were 
amphidiploid in origin (Goodspeed, 1934). Proof of such origin would con- 
tribute materially to the systematic botany of the genus and to an interpre- 
tation of its species distribution in the two Americas. For two of the 24-paired 
species, such proof is at hand. Thus, iV. Tahacuni (n = 24) has been shown to be 

* Contribution No. 112 from the University of California Botanical Garden. 

Acknowledgment is made of grants received from the Committee for Inter-American 
Artistic and Cultural Eelations, the American Philosophical Society, the American Academy 
of Arts and Sciences, the Coolidge Foundation, and to a number of private donors, in sup- 
port of the studies reported upon. The author is indebted to Dr. H-M. Wheeler and M. C. 
Thompson for assistance in the preparation of this article. 

[291] 



292 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

au amphidiploid product of hybridization between progenitors of the modern 
species N. sylvesiris (n = 12) and N. tomentosa (n = 12) or its relatives (Good- 
speed and Clausen, 1928) . These two species are peculiar to the Andes of Peru, 
Bolivia, or Argentina, and one of the products of the three University of Cali- 
fornia Botanical Garden Expeditions to the Andes has been the demonstration 
that today the distribution of the putative parental species overlaps. Simi- 
larly, progenitors of two other present day Peruvian-Bolivian-Argentine 
species, N. paniculata (n=12) and N. undnlata (n=12) , were undoubtedly the 
parents of the amphidiploid species N. rustica, the other well-known 24-paired 
species of commerce (Goodspeed, 1934, 1936). 

Despite cytogenetic evidence in favor of amphidiploid origin, it has not been 
possible to reproduce artificially a duplicate of modern N. T aha cum, or of 
N. rustica, nor is there evidence of tlie existence of at least the former species 
in a wild state. In other words, although there is good reason to believe that 
the 24-paired American species of Nicotiana are of amphidiploid origin and 
while amphidiploidy has been induced in at least 27 experimentally produced 
Fj intersiT^cific hybrids (Bradley and Goodspeed, 1943), no naturally oc- 
curring American species has hitherto been found for which such origin could 
be shown.^ It now appears that the recently discovered N. Arentsii is such a 
species. 

In November 1938 one of the members of the Second University of Cali- 
fornia Botanical Garden Expedition to the Andes, Dr. Cesar Vargas C, Pro- 
fessor of Botany and Director of the Botanical Museum in the University of 
Cuzco, Peru, found N. Arentsii in southeastern Peru, north of Lake Titicaca, 
near the Bolivian border, in and about the village of Limbani. Limbani is 
situated in the Dept. Puno, Prov. Sandia, at an altitude of about 3500 m., 
in a narrow box canyon on the eastern watershed of the Cordillera. The region 
is remote, climatically inhospitable, and little known botanically. Nearby, 
at somewhat lower altitudes, is the beginning of the Amazonian rain forest. 
The following- extract from Dr. Vargas' notes is included here, in part because 
of its general botanical interest. Below Limbani, "the river of the same name 
runs swiftly and precipitately, a turbulent and foam-flecked stream which 
springs from the thawing snows of the nearby Cordillera. The village mostly 
consists of hovels and cabins nearly all made of wliitish granitic rock with roofs 
of straw. The only street is the tortuous highway, lined on both sides by the 
principal habitations while the rest are scattered here and there on contiguous 
slopes. On the walls of the canyon are a series of shelves or terraces, now in 
ruins. Nobody knows how many hundreds of years ago they were constructed 
through the laboriousness and perseverance of a people who disappeared long 

^ Kostoff (1939) has commented upon the origin of a 32-paired race of the Australian 
species N. suavcolms (n = 16) or its close relatives whieh was first grown in the University 
of California Botanical Garden (Wheeler, 1935). For tliis race Kostoff has suggested the 
name N. Eastii (but does not formally describe it as a new species) and considers it an 
amphidiploid product of hybridization between N. suaveolens and N. maritima. We have 
grown more than one 32-paired race from Australia and their morphological distinctions 
appear to complicate the problem somewhat. 



VOL.XXV] GOODSPEED : NICOTIANA ARENTSII SP. NOV. 293 

ago and for wliom the soil and the plant constituted the most cherished gifts 
of nature. . . . However, what is most cheering ... is the abundance of vegeta- 
tion . . . herbaceous and shrub forms as far as my eyes reach ; even the rocky 
cliffs offer a varied flora. Along the road there are found several species of 
the families Melastomaceae, Ericaceae, Canipanulaceae, etc. . . . beautiful and 
graceful clusters of Bomarea hang among Baccharis, Cantua, and other 
bushes ; under the rocks and on the demolished terraces grow two species of 
Begonia, Solanum, Centropogon, Monnina and, in the dank turf, species of 
Viola and Calceolaria. The air was humid, the soil moist." The Quechua name 
for the new Nicotiana species was "Cjamasairi." 

Four years later, Mr. Roy D. Metcalf, a member of the Third University of 
California Botanical Garden Expedition to the Andes, during an exploration 
of the Inambari and Tambopata drainage basins, collected N. Arentsii again. 
He found it restricted to the immediate vicinity of the village of Limbani and 
commonly around habitations, in some cases growing on walls and fences. 
According to his report, it prefers semi-shade, moist but not wet surroundings 
and gravelly soil among rocks. The preference of N. Arentsii for disturbed 
ground and the partial protection provided by habitations is characteristic 
of many South American species of Nicotiana. In Limbani the Indians did 
not make any use of the plant. Although Mr. Metcalf 's itinerary northward 
from Chucuito (Dept. and Prov. Puno) at the northwest end of Lake Titicaca, 
took him as far as the Santo Domingo mine (immediately north of the Inam- 
bari river) in the province of Sandia and as far east as the towns of Sandia 
and Cuyocuyo, he did not again encounter this species. He states that N. un- 
dulata was also called Cjamasairi by the natives. 

From the original seed of N. Arentsii collected by Dr. Vargas, 100 plants 
have been grown in the University of California Botanical Garden at Berke- 
ley, California. Of these, 25 were grown in 1940, 75 in 1941. In the former 
year, all plants were set out in open ground; in the latter year, 35 were planted 
out, 38 held outdoors in large pots, and two allowed to flower in pots in the 
greenhouse. Potted plants from seed sown on January 10, 1941, began to 
bloom August 28, 1941. Those plants transplanted to open ground from seed 
sowai on February 16, 1940, and January 10, 1941, began to bloom in the 
middle of the next October. To test their reported perennial capacities the 
field culture was not discarded at the close of the 1940 growing season. Al- 
though the winter (of 1940) was moderate in temperature, the plants died 
after setting seed. All plants held in large pots failed to develop lower 
branches. Their inflorescences were narrower than those of field plants. 
Finally, plants blooming in the greenhouse lacked the rose-red pigmentation 
in calyx and corolla, characteristic of field grown plants. 

In all these garden cultures the branching was variable and frequently 
asymetric. There was also a strong tendency toward fasciation of elements of 
the inflorescence. Generally, the union did not extend beyond the first leaf of 
the axillary shoot. The branch thus affected would depart from the main axis 



294 CALIFORNIA ACADEMY OF SCIENCES [Pkoc. 4th ser. i 

at a decidedly more acute angle than that of a normal shoot. Faseiation did 
not appear in a population of 13 plants grown, under equivalent conditions, 
in 1943 from selfed seed of a 1942 plant. Despite certain definite but inter- 
grading variations in leaf shape and inflorescence type, the appearance of the 
original populations was as uniform as might be expected from seed of wild 
plants. 

On the basis of herbarium and garden studies, N. Arcntsir was found 
clearly to combine prominent morphological features of two other species of 
Nicotiana — i.e., N. wigandioides Koch and Fint" and N. undulata R. and P." 
The former is a subarborescent species, less massive than N. tomentosa but 
attaining a height of several meters and capable of living a number of years 
in the Berkeley climate. Especiall}^ on younger stem parts, long straight 
silvery hairs stand out prominently ; there being, however, a glabrous region 
below the insertion of the leaf. The length, and to some degree the direction, 
of these soft, glistening hairs is modified in N. Arentsii. The red-purple colora- 
tion of N. Arentsii manifested in stem, leaf, calyx, and a pinkish blush to the 
corolla is derived from N. wigandioides. Plants of both species blossoming in 
the greenhouse instead of outdoors, lack it. Slow maturing, despite early 
sowing, is characteristic of both, N. undulata on the other hand, contributes 
to N. Arentsii relativelj^ low stature and annual to limited perennial habit, 
greater leafiness in the inflorescence region, and a large adaxial calyx lobe. In 
certain plants it modifies the calyx coloration to a blue-black. 

Generally speaking, N. wigandioides is a tropical or subtropical species as 
yet known only from central (western) Bolivia. It may be associated with the 
eastern headwaters of the Beni river for it has been collected a short distance 
beyond the edge of this drainage system (Cardenas 2800, Inca Corral, Dept. 
Cochabamba, Prov. Chapare, 1700 m. ; Steinhach 5754, Cerro de Ineachaca, 
Dept. Cochabamba, Prov. Sacaba,* 2500 m.) and well within it in valleys in 
Dept. La Paz (Rushy 75a, Mulford Biol. Explor. Amazon Basin, Caiiamina, 
Prov. Inquisivi, 1100 m. ; Troll 2812, Lambate, Prov. Sud Yungas, 3400 m.; 
Buchtien 3905, Unduavi, Prov. Nor Yungas, 3200 m. ; Bushy 2434, near 
Yungas, 1200 m.). It may also occur to the north and it is unfortunate that 
we have been unable to explore the tributaries of the Rio Beni in Bolivian 
territory adjacent to the Peruvian area where N. Arentsii was found. 

Speaking of the vegetation at his last cited locality, H. H. Rusby (1888, p. 
181) writes: ''Descending to 7000 feet, we enter the great Andean forests 
which become heavier and heavier, though scarcely denser, as we descend. The 
trunks and greater branches are scarcely to be seen for the epiphytes upon 
them, chief of which are orchids, bromeliads, ferns, mosses, and aroids. At 
5500 feet, we strike the coca and cinchona belt, and at 4000 feet we find the 
heat becoming oppressive and the air sultry. From 3500 to 5500 feet is prob- 

- A detailed description of this new species appears at the end of this report where photo- 
graphs of it and of N. undulata and N. wigandioides ■will be found. 

^ Undoubtedly a mistake and should read "the canton of Saeaba, in Prov. Chapare." 



Vol. XXV] GOODSPEED : NICOTIANA ABENTSII SP. NOV. 295 

ably the region of greatest rain-fall." Beyond general pictures such as this 
there is little description of habitat for N. wigandioides except the remarks 
of Dr. Martin Cardenas on his collection of this species from Inca Corral. He 
refers to the region as "Ceja de Monte ; the 'Ceja' is a kind of beginning of the 
Yungas." The term Yungas is used here as a collective designation for Bolivian 
tropical valley terrain. He states that N. ivigandioides grows "on wet soil." 

N. undidata is a widely distributed species which, by contrast, thrives in 
high, cold, bleak situations and generally fails to compete successfully as the 
climate becomes less rigorous and the region can support a richer vegetation. 
Apparently, it does not descend the eastern flank of the Andes but every- 
where finds a congenial habitat in the Puna zone to the west. Its range is 
continuous from northern Argentina (Univ. Calif. Bot. Card. Exped. Andes 
6323, coll. West, 20 km. w. of Humahuaca, Prov. Jujuy, 3600 m.) to central 
Peru (Univ. Calif. Second Bot. Card. Exped. Andes 10985, coll. Stork, 
Muilo. 10 km. w. of Tarma, Dept. Junin, 3300 m.) and it has even been 
found in northern Peru (Univ. Calif. Third Bot. Card. Exped. Andes 
30781, coll. Metcalf, road from Cajamarca, 3200 m.). Near Oroya, Peru, it 
forms almost pure colonies where only it and Bidens can tolerate the fumes 
from nearby smelters (Univ. Calif. Second Bot. Card. Exped. Andes 10986, 
coll. Stork) ; between Candarave and Puno it occurs in the hard-packed road- 
bed (Univ Calif. Third Bot. Card. Exped. Andes 30387, coH. Metcalf) ; in 
Chucuito, Huancane, Putina, Asillo and other windswept altiplano towns of 
the Dept. Puno (ibid. 30677, 30704, 30718, 30424) it is common in the streets, 
on refuse dumps, and in the protection of houses and fences. Metcalf reports : 
"While crossing the altiplano, the one plant that one expects to find is this 
Nicotiana. "When there is nothing else, there is always this . . . while never 
growing in great abundance in any one site, it none the less covers sparsely 
huge areas. Koughly, N. undulata occurs from 10,000 feet to over 14,000 feet." 

Although N. wigandioides has not been found near the limited known range 
of distribution of N. Arentsii, the other species, N. undulata, which appears 
to have entered into its amphidiploid origin, occurs within 30 km. of it. In the 
La Paz region of Bolivia, N. undulata and N. wigandioides have been col- 
lected within this distance and in Dept. Cochabamba they are found within 
10 km. of each other. It is, therefore, not too much to say that in past time 
these two species have been in contact, if they are not so at present. In the 
dissected eastern edge of the Peruvian-Bolivian Andes, alps and tropics are 
remarkably juxtaposed and wherever a cleared pathway opens, whether it be 
by diastrophic movement, streams, or man-made paths, Nicotiana species will 
seek to colonize the disturbed ground to the limit of their climatic tolerance. 

Above, it has been pointed out that N. Arentsii is morphologically a mosaic 
of the distinctive characters peculiar to two other species of Nicotiana. Atten- 
tion has also been called to the fact that, at one time, the range of distribution 
of these two species undoubtedly overlapped and that today one of them is 
known to be practically in contact with N. Arentsii. These two lines of evi- 



296 CALIFOSXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

dence suggest that this new species originated as a hybrid between progenitors 
of the two other species, chromosome doubling then occurring in the hj^brid. 
The following cytogenetic observations largely confirm this suggestion. 

It is, first, to be noted that the chromosome number of N. Are7itsii is 24 
pairs while both N. undulata and N. wigandioides possess 12 pairs. In an 
earlier report (Goodspeed 1934), it was shown that the somatic karyotype of 
N. undulata consists of chromosomes with median or submedian centromere, 
one of the submedian pairs satellited. In the case of N. wigandioides there are 
8 pairs with median or submedian centromere and 4 with subterminal, one of 
the subterminal satellited. As regards size, there is little difference between 
the two complements; less, indeed, than between the largest and smallest 
chromosomes within a complement. Both karyotypes belong to the larger, but 
not the largest chromosome size groups in the genus. Detailed study of its 
chromosome morphology shows that the complement of N. Arentsii represents 
a summation of the two species types, with mostly median or submedian cen- 
tromere but a few with subterminal. The scant size distinctions between the 
N. undulata and N. wigandioides complements do not, however, make it feasi- 
ble to homologize full complements but it may be said that the morphological 
equivalents of the satellited pair characteristic of each of the two species has 
been identified in N. Arentsii. 

Meiotic configurations (pollen mother cells) of each of the three species 
were studied to determine whether any characteristic N. undulata or N. 
wigandioides features reappear in N. Arentsii. In each of the 12-paired 
species, two IM bivalent types w^ere selected as sufficiently distinctive to be 
identified : in iV. undulata (1) 'a completely terminalized one-ehiasma bivalent 
with oblong partners and (2) a completely terminalized thick ring bivalent; 
in N. wigandioides (1) an incompletely terminalized one-chiasma bivalent in 
which two droplet partners are separated by a central (generally small) bead 
and (2) a not quite terminalized, two-chiasma "diamond" bivalent. Each of 
these types reappeared in N. Arentsii. 

In N. undidata and N. wigandioides, meiotic chromosome behavior is normal 
but in N. Arentsii pairing was sometimes found to be irregular, even in plants 
which produced only good pollen grains. It is not possible to say that this 
condition prevailed in every plant, as only a few were examined in first 
metaphase, but the fact that it was true of every plant examined is significant. 
It was noted that from 2 to 6 chromosomes (often in more than 50 per cent of 
the PMC) either failed to pair or formed trivalents. In 20 second metaphase 
counts of one plant, this irregularity reduced to 85 per cent the PMC showing 
24-24 distribution of chromosomes, the remaining 15 per cent showing 23-25. 

Beyond a determination of chromosome number, the cytology of N. Arentsii 
had not been studied at the time hybrids of this new species with N. undulata 
and N. wigandioides were made and grown. A few plants each of both F^ 
hybrids were secured and inasmuch as the preponderance of meiotic material 
available proved to be from monosomic hybrids made with the same N. Arentsii 



Vol. XXV] GOODSPEED : NICOTIAN A ABENTSII SP. NOV. 297 

parent, it is clear that the particular N. Are7itsn plant involved in the crosses 
was itself monosomic. Apart from somewhat complicating the determination 
of extent of pairing, this situation did not alter the essential evidence, briefly 
summarized as follows : 



Fi Hybrid 


PMC 

counted 


Plant 
number 


Chromosome 

Constitution 


Pairs per . 
Range 


PMC 
Average 


N. undulata 


14 


F-341 P54: 


36 


11-12 


11.71 


X >■ 
N. Arentsii 


25 


F-385 P51 


35 


11-13 


11.96 


N. Arentsii 


25 


F-386 P55 


36 


11-13 


11.92 


X > 


25 


r-386 P58 


35 


11-13 


12.08 


N. wigandioides 


25 


F-386 P59 


35 


11-13 


11.96 



In the case of F^ N. Arentsii x N. wigandioides, somewhat more than 50 per 
cent of the counts represent 12 bivalents only. The remainder involve combina- 
tions ranging from 8n + Sm to 13n with the largest class lOn + liv and the next 
llii + liii- Almost the same types of association occurred in the case of F^ N. 
undulata x N. Arentsii except that the proportion of 12ii was approximately 
40 per cent. In view of the extent of multivalent formation in the two hybrids 
just mentioned, the occurrence of some pairing in the hybrid which represents 
the third side of the triangle, F^ N. undulata x N. wigandioides, was not un- 
expected. Thus, a range of from 2 to 9 pairs was observed in 100 pollen mother 
cells while the average number of pairs was approximately 5. 

For our present purpose, it is sufficient to call attention to the fact that the 
cytogenetic evidence shows that the gametic sets of both N. undulata and N. 
wigandioides are represented by a full set of homologous chromosomes in the 
gametic set of N. Arentsii and that the amount of pairing in F^ N. undulata x 
N.~ wigandioides is low. In other words, the cytogenetic findings are consistent 
Avith the' evidence from external morphology and from distribution which 
indicate that N. Arentsii represents an amphidiploid product of hybridization 
between those other two species of Nicotiana. 

Nicotiana Arentsii Goodspeed, sp. nov. 

Nicotiana herbacea; caule crasso, conspicue pubescente; foliis petiolatis, 
late ellipticis, rotundo-ovatis vel subcordatis; floribus in paniculas foliosas 
dispositis ; calyce magno, lobis inaequalibus ; corolla dilute straminea v. prope 
alba, obconico-clavata, limbo plus minusve zygomorpho, lobis obtusis; stami- 
nibus inclusis, abrupte curvatis, ad partem tubulatam corollae inferne affixis. 

Type. — University of California Second Botanical Garden Expedition to the Andes, 
1938-39, 9666 (U. C. Herbarium 665156). Coll. C. Vargas C, Nov. 21, 1938, Limbani, Dept. 
Puno, Prov. Sandia, Peru. Alt. 3400 m. Close to the road and houses. "Cjamasairi." Other 
authentic material: University of California Third Botanical Garden Expedition to the 
Andes, 1942^3, 30529. Coll. E. D. Metcalf, May 16, 1942, in town of Limbani, Dept. Puno, 
Prov. Sandia, Peru. Alongside huts, north end of town. Alt. 3350 ni. 

Slowly maturing annual or limited perennial 1-2 m. high. Stem commonly 
reddish purplish, rather densely clothed with distinct but not long, weak, 



298 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

silvery-white hairs, ultimately oldest parts glabrate, thinly corky, woody 
within ; main axis stiffly erect, 2-2 cm. thick, chief branches few, irregular in 
length and distribution, usually some long and rapidly ascending. Leaves 
pubescent ; blade to 25 or 30 cm. long and about 20 cm. wide, rotund-ovate or 
subcordate or the uppermost often lance-ovate, base variable, apex acute or 
acuminate, veins evident, sometimes indenting upper surface a little, margin 
slightly undulate; petiole %-^/'io as long as blade; frequentlj^ red coloration 
in petiole, midrib, veins, margin or even hairs. Inflorescence paniculate, 35-50 
cm. long, narrow or broad, lower limits indistinct by very gradual reduction 
of leaves to sessile, broadly lanceolate bracts ; central axis erect, thick, pubes- 
cent but seldom viscid ; laterals numerous, frequently rather crowded, variable 
in leng-th, generally rigid and spreading, paniculately or occasionally cymosely 
branched, ±: viscid; ultimate pedicels about 2 mm. long, later 6 mm. Flowers 
abundant, odorless or nearly so. Calyx green, green suffused with red-purple, 
or green blended with blue-black, pubescent, 10-14 mm. long, broadly cylindric 
or subcampanuiate ; membranous area below each sinus narrow, short ; lobes 
shorter than tube, broadly triangular-ovate, plane, unequal, largest never 
extending to corolla mouth. Corolla distinctly pubescent exteriorly except in 
the partially glabrous limb; tubular part 15-20 mm. long, 7-9 mm. wide, 
broadly ob conic from short, narrower base, apically somewhat swollen im- 
mediately below the slightly contracted mouth, pale greenish yellow with pink 
or purplish tingeing above; entire limb (including rimmed, hexagonal mouth) 
to 17 mm. across, in bud spirally plicate and often bright pink, in flower 
spreading, pale straw color frequently conspicuously suffused with pink 
outside, rarely also some pink inside ; lobes broad, obtuse, 3 upper ones 
smaller, commonly a little deflexed. Stamens inserted about 5 mm. from very 
base of corolla (at or near where the obconieal expansion begins) ; filaments 
thick, pale greenish, 4 equal or subequal, wooly and erect for several mm. 
above insertion, then glabrous and curving outward abruptly to rest against 
corolla, extending to about 1 mm. below corolla mouth, 1 sigmoid, not wooly 
above insertion, commonly appearing somewhat shorter ; anthers yellow-green. 
Pollen pale straw color. Hypogynous disk thick, red or red-orange. Ovary 
rotund-ovoid, greenish or purplish; style thick, faintly greenish; stigma 
green, discoid-pulvinate, median groove shallow. Capsule included, 7-9 mm. 
long, elliptic-ovoid, brown, on dehiscence upper half split into 2 divergent, 
notched valves. Seeds broadly elliptic-ovoid, 0.7-0.8 mm. long, rather dark 
brown, surface shallowly reticulate, meshes moderately large, deeply lobed 
by bounding ridges, embryo straight. Somatic chromosome number 48. 

Distinguishable from N. undulata R. & P. by later maturity ; longer, more 
distinct, glistening hairs; larger, rotund-ovate leaf; larger flower; longest 
calyx lobe plane, not boat-shaped in flower or fruit ; corolla limb sometimes^ 
pinkish, never greenish ; from .¥. ivigandioides Koch & Fint. by shorter life 
span ; smaller leaf ; markedly unequal calyx segments ; smaller corolla limb. 

Named in honor of Mr. George Arents, to whom botany is indebted for a| 
comprehensive compilation of the history and culture of tobacco. 



Vol. XXV] GOODSPEED : NICOTIANA ARENTSII SP. NOV. 299 

SUMMARY 

A new species of Nicotiana, N. Arentsii (n = 24) , from southeastern Peru is 

described, with comments upon its morphology, distribution and cytology. 

Similar comments are made in the case of two other Nicotiana species, natives 

of Peru and Bolivia — N. undulata (n = 12) and N. tvigandioides (n = 12). The 

morphological and distributional evidence points to an amphidiploid origin 

for N. Arentsii in which the other two species are involved. This hypothesis is 

confirmed by the cytology of the three species and of the F^ hybrids made 

between them. 

(Since the above ■was written, it has been possible to reexamine the herbarium material 
of two Bolivian collections of H. H. Rusby which, before Nicotiana Arentsii was recognized, 
were identified as N. wigandioides. It is now clear that they were taken from plants of N. 
Arentsii. Both are Rusby's No. 822 and both are from Mapiri (Dept. La Paz, Prov. Lare- 
caja) but one is labelled "5000 ft., April, 1886" and the other "2500 ft.. May, 1886." Mapiri 
is approximately 200 km. southeast of the Limbani area in which the collections of N. 
Arentsii described above were obtained.) 



LITERATURE CITED 

Bradley, Muriel V. and Goodspeed, T. H. 

1943. Colchicine-induced alio- and autopolyploidy in. Nicotiana. Proc. Nat. Acad. Sci. 29 : 
295-301. 

Clausen, R. E., and Goodspeed, T. H. 

1925. Interspecific hybridization in Nicotiana. II. A tetraploid glutinosa-tabacum hybrid, 
an experimental verification of Winge's hypothesis. Genetics 10:278-284. 

Goodspeed, T. H. 

1934. Nicotiana phylesis in the light of chromosome number, morphology, and behavior, 
Univ. Calif. Publ. Bot. 17:369-398. 

1936. Phylesis in Nicotiana. Abstract, Proc. VI Internat. Bot. Congr. 1:164-165. 

Goodspeed, T. H., and Bradley, Muriel V. 
1942. Amphidiploidy. Bot. Rev. 8:271-316. 

Goodspeed, T. H., and Clausen, R. E. 

1928. Interspecific hybridization in Nicotiana. VIII. The sylvestris-tomentosa-tabactim 
hybrid triangle and its bearing on the origin of tabacum. Univ. Calif. Publ. Bot. 
11:245-256. 

KOSTOFF, DONTCHO 

1939. The origin of the tetraploid Nicotiana from Bathurst. Curr. Sci. 8:110-111. 

RusBY, H. H. 

1888. An enumeration of the plants collected by Dr. H. H. Rusby in South America. 1885- 
1886. I. Bull. Torr. Bot. Club 15:177-184. 

Wheeler, H-M. 

1935. Studies in Nicotiana. II. A taxonomie survey of the Australasian species. Univ. 
Calif. Publ. Bot. 18:45-68. 



i 



EXPLANATION OF PLATES 



PLATE 26 

Left: Nicotiana Arentsii, grown from seed of Univ. Calif. Second Bot. Gard. 
Exped. Andes 9666, Limbani, Dept. Puno, Peru, coll. C. Vargas C. 

Center: N. vndvlata, grown from seed of Univ. Calif. Bot. Gard. Exped. 
Andes 7173, Pisacc, Dept. Cuzco, Peru, coll. James West. 

Right : N. wigandioides, grown from seed collected by Jose Steinbach, near 
Cochabaniba, Bolivia. 



i 



{ 3U2 J 



ID 
CM 

UJ 
h 
< 

Q. 

Q 
III 
Lli 

a. 
w 
D 



o 





PLATE 27 

Upper row : Nicotiana ujidulata, from seed collected by A. Weberbauer, at 
Huancavelica, Peru. The species is polymorphic in corolla shape and size. The 
race figured is relatively large flowered with the tubular part of its corolla 
broad and little dilated below the mouth. 

Middle row : N. Arentsii, source as in plate 26. 

Lower row: N. wiyandioides, source as in plate 26. 

(Black paper has been inserted in the corolla tubes of the sectioned flowers 
and one or two stamens removed to show the pistil. In N. undulata and N. 
Arentsii those removed included a short stamen ; in N. wigandioides the two 
removed are no shorter than those remaining.) 



[304] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 12 [GOODSPEED] PLATE 27 




[ 305] 



PROCEEDINGS 

OF THE ^ 

CALIFORNIA ACADEMY OF SCIENCESX 

Fourth Series 
Vol. XXV, No. 13, pp. 307-322, figs. 1-13 November 10. 1944 




ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 13 

THE CYTOGENETICS OF HYBRIDS IN BROMUS II. 
BROMUS CARINATUS AND BROMUS ARIZONICUS 

BY 

G. L. STEBBINS JR., H. A. TOBGY, and JACK R. HARLAN 

University of California 

INTRODUCTION 

THE FACT that Bromus carinatus H. and A. and its relatives constitute a 
complex of polyploid entities, mostly if not entirely of allopolyploid or 
hybrid origin, has been brought out in the first paper of this series (Stebbins 
and Tobgy, 1944). In an earlier survey of chromosome numbers (Steb- 
bins and Love, 1941) the octoploid number of chromosmes (2n=56) was 
counted in all representatives of this complex collected in California. Counts 
made by us of collections from Oregon, Washington, Idaho and British Colum- 
bia have revealed the same number, so that we can state with some assurance 
that the Pacific Coast members of this complex are predominantly if not en- 
tirely octoploid, with the exception of a type originally collected by one of 
us (Harlan) in Central Arizona. The 28 collections of the B. carinaius com- 
plex made in this state were found to be divided into two series on the basis 
of morphological and ecological characteristics. The first series, found mostly 
in desert washes, roadsides, and waste places at lower altitudes consists of 
strictly annual plants with relatively long- awns ; while the second, found pre- 
dominantly at altitudes of 5000 feet (1500 meters) or higher, consists of 
perennials, with broad g'lumes ; usually many-flowered spikelets, and shorter 
awns. The first series agree with the original description of B. carinatus var. 
arizonicus Shear (1900), and would be identified as B. carinatus according to 

I 307 1 



308 CALIFORXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

the key of Swallen (1942) in the Arizona flora of Kearney and Peebles. The 
second, since all but one of the specimens collected have essentially glabrous 
foliage, "svould be considered B. polyanthus Shear, according to the treatment 
of Swallen (Joe. cit.). Chromosome counts on five different collections of this 
second series revealed 56 chromosomes in them, as in Pacific Coast members 
of the complex. A detailed description of one collection of it (no. 116), with 
an account of its crossing relationships with other octoploid species, is pre- 
sented elsewhere (Stebbins and Tobgy, 1944). 

The annual, long-awned series, on the other hand, was found to possess 
uniformly the somatic number 2n=84, or 42 bivalents at meiosis, in material 
from 21 different collections (see map, fig. 4). Furthermore, the same chromo- 
some number was found in a similar annual form from Tehachapi, Kern 
County, California. When these duodecaploids were grown in the same field 
at Berkeley along with a large series of octoploids from California as well as 
Arizona, it was found that certain morphological characteristics made them 
readily distinguishable from the perennial or subperennial, long-awned octo- 
ploid types from lower altitudes in California, as well as from the short-awned 
octoploids found at higher altitudes in both states. For this reason, the specific 
distinctness of the 84-chromosome type was suspected, and the testing of this 
hypothesis by means of hybridization seemed desirable. The results of this 
hybridization and their implications in regard to the status and possible origin 
of the 84-chromosome type are here presented. 

MATERIAL AND METHODS 

Seeds from the Arizona collections were all planted in the field in the sum- 
mer of 1941. Although each progeny from a single collection was with a few 
exceptions remarkably uniform, there was a great deal of variation between- 
collections (Harlan, in press). Some of the seedlings behaved like desert 
ephemerals, producing only one or two tillers, which flowered in a few weeks 
when only 10 or 15 cm high, and produced only 2 or 3 spikelets, dying when 
the seed matured. Others behaved like winter annuals, and in the spring pro- 
duced very tall plants with several tillers and large panicles. It was from one 
of the latter type that the parent stock for hj^bridization with octoploid B. 
carinatus was selected. The particular collection, no. 125, came from south of 
Lake Meade, near Boulder Dam, Mojave County. The octoploid strain used 
for hybridization was a typical B. carinatus strain from Berkeley, represented 
in the University of California Herbarium by the collection Stebbins no. 2677, 
and described elsewhere as strain no. 5 (Harlan, in press ; Stebbins and Tobgy, 
1944). The hybridization was made in the field in May 1942, by Mr. Henry 
Moser, according to the method described earlier (Stebbins and Tobgy, loc. 
cit.). The cytological techniques used are described in the same paper. 



Vol. XXY] STEBBINS-TOBGY-HAELAX : HYBRIDS IN BROMUS II 300 

MORPHOLOGY AND SYSTEMATIC DESCRIPTION OF 
THE SPECIES AND THE HYBRID 

The morphological and cytological data presented below leaves no doubt 
that the 84-chroinosome type is a species sharply distinct from any now recog- 
nized in the B. carinatus complex, even though its diagnostic characteristics 
are somewhat meager. Therefore, a taxonomic description of it will be given 
at the outset, so that it may thereafter be referred to by its correct name. 
Although the type specimen of Shear's B. carinatus var. arizonicus has not 
been seen, and the original description is somewhat scanty, nevertheless there 
is little doubt that it belongs to the 84-chromosome species described here. All 
of the collections made by Harlan from situations similar to Tucson, its type 
locality, are of this species, and it is the only member of the complex known 
from lower altitudes in Arizona. One collection, no. 105, in which 84 chromo- 
somes were counted, came from the Santa Cruz Valley only 35 miles from the 
tj^pe locality. It may be described as follows (the taxonomic section is the 
work of the senior author) . 

Bromus arizonicus (Shear) Stebbins n. comb. 

Planta annua; culmi humiles vel alti; paniculae parvae spiculis paucis, 
vel ampla spiculis numerosis; glumae elongatae, secunda infimam lemmam 
aequans; lemmata ad marginem hirsuta, dorso scabra vel glabra, lemmata 
superiores conspicue bidentata. 

Plants annual ; culms 15-140 cm high, with 3-5 nodes. Leaves somewhat 
glaucous, glabrous or pubescent. Panicle sometimes much reduced, but in well 
developed plants ample, the branches spreading, in groups of 5-6 at the lower 
nodes, the longest bearing 5-8 spikelets ; branches often finely hirsute, the 
hairs more slender than in B. carinatus. Spikelets mostly 5-7 flowered ; glumes 
subequal or distinctly unequal, elongate, the upper glume ahotit equalling in 
length the lowest lemma. Lemmas of medium length, mostly 11-14 mm long, 
sparsely or densely hirsute near the edges, short-scahrous on the hack, iiden- 
tate, the teeth rather prominent, those of the upper lemmas often icith the 
lateral nerves extending into them. 

Bromus carinatus var. arizonicus Shear, U. S. Div. x\grost, Bull, no, 23 : 
62.1900. Type in the U. S. National Herbarium, from near Tucson, Arizona, 
C. G. Pringle in 1884. B. carinatus calif amicus Shear, U. S. Dept. Agr. Div. 
Agrost. Bull. 23 :60, at least in part. 

Central and southern Arizona to southern and central California, in desert 
areas and open, disturbed ground of valleys ; an introduced weed in the north- 
western portion of its present range. The following specimens, in the herbaria 
of the University of California (UC) and the California Academy of Sciences 
(CAS), are typical: CALIFORNIA: Mark West Creek, jBoZancZer 3985 (UC) ; 
Davis, Yolo County, Stelhins 3356 (UC) ; Alcalde, Fresno County, Eastwood 
13492 (CAS) ; Tulare, Davy 3055 (CAS); Kettleman City, Kings County, 



310 



CALIFORNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 



Hoover 442 (UC) ; Bakersfield, Kern County, Davy 1896 (UC) ; San Emigdio 
Caiion, Kern County, Davy 1985 (UC) ; Tehachapi, Kern County, Harlan 
128 (UC) ; Paso Robles, San Luis Obispo County, Barher in 1900 (UC) ; Santa 
Maria, Santa Barbara County, Eastwood 344 (CAS) ; Pasadena, Los Angeles 
County, Grant 117, 117a (UC, CAS) ; Claremont, Los Angeles County, Davy 
in 1902 (UC) ; San Bernardino, Parish in 1889 (UC) ; Colorado Desert, Riv- 
erside County, Hall 5980 (CAS) ; Fall Brook, San Diego County, Jones 3108 




Fig. 1. Bronins carbial us, sti'ain no. 5. 
Fig. 2. Bromus arizonicus x carmatus, no. 306.5. 
Fig. 3. Bromus arizonicus, no. 311.1 (progeny of Harlan no. 125). Figs, la, 2a, 3o, spike- 
lets, X 1. Figs, lb, 2b, 3b, lower glumes; Ic, 2c, Sc, upper glumes; Id, 2d, 3d, lemmas and 
paleae, all x 2. Figs. Ic, 2e, 3e, apices of lemmas, x 4. Figs. If, 2f, of, portions of surface of 
lemmas, with midrib at right and margin at left, showing pubescence, x 8. 



(CAS) ; San Diego, Orciitt 1178 (UC) ; between Del Mar and Lower Cali- 
fornia, Lemmon in 1888 (CAS) ; Santa Catalina Id., Branclegce in 1890 
(UC) ; Santa Cruz Id., Brandegee in 1888 (UC) ; Santa Rosa Id., Hoffmann 
in 1929 (CAS) ; San Clemente Id., Murlarger 56 (UC) ; San Nicolas Id., 
J. T. Howell 8226 (CAS). BAJA CALIFORNIA : San Quintin, Epling and 
Stewart in 1936 (CAS) ; 24.3 miles east of Rosario, Wiggins 5291 (UC, CAS). 
ARIZONA: Wickenburg, Maricopa County, W. W. Jones in 1921 (UC) ; 
Tempe, Maricopa Count)'-, Gillespie 5519 (UC) ; Fort Lowell, Thornher 535 
(UC) ; 25 miles southeast of Lake Meade, Mohave County, Harlan 125 (UC) ; 
east of King-man, Mohave County, Harlan 123 (UC) ; Pinal Mts., southwest 
of Globe, Gila County, Harlan 112 (UC) ; Salt River Canyon, Gila County, 
Harlan 115 (UC) ; Sacaton, Pinal County, Harlan 103 (UC) ; east of Flor- 



Vol. XXV] STEBBINS-TOBGY-HAELAN : HYBRIDS IN BEOMUS II 311 

ence, Pinal County, Harlan 107 (UC) ; Red Rock, Pinal Comity, Harlan 
105 (UC). 

As stated preYioiisl.y, the chromosome number 2n=84 has been determined 
in all of the Harlan collections cited. 

Without having seen the type specimen, the writer cannot determine 
whether B. arizonicus is the same as B. carinatus calif ornicus Shear, or B. 



^7^^. 




Fig. 4. Map showing distribution of Harlan collections of Bromus arizonicus (2n =84), 
solid circles, and B. ijolyanthus vel aff. (2?i = 56), hollow squares, in Arizona. The 5000 foot 
contour level is drawn. 

calif ornicus Nutt. Nuttall's specific name is a nomen nudum (cf. Shear 1900), 
and his specimen, according to Shear, "is a mere scrap," of which the locality 
of collection is indefinite. Nevertheless, the specimens cited by Shear under 
B. carinatus calif ornicus, since all come from San Diego or Lower California, 
are very likely B. arizonicus. 

The characteristics which distingiiish B. arizonicus from all other members 
of the subgenus Ceratochloa in the Western United States are the strictly 
annual habit (the plants always die when the seeds have matured), the elon- 
gate glumes, the hirsute pubescence near the margin of the lemmas, and the 
rather prominent lobes of at least the upper lemmas (fig. 2) . All of the typical, 
octoploid members of the B. carinatus complex as grown in Berkeley pro- 
duced some new tillers after the seed had ripened, except when the plants 



312 



CALIFORNIA ACADEMY OF SCIEXCES [Proc. 4th ser. 



were obviously diseased. In all of them the lowest lemma is distinctly longer 
than the upper glume ; its pubescence, whether long or short is more or less 
evenly distributed over the surface ; and the lobes are very short, with the 
latei-al nerves ending far below their base (fig. 1). 

The relative distributions of B. arizonicus and the octoploids referred to 
B. folyanthiis are shown on a map (fig. 4) giving the localities from which 
chromosome counts were obtained. Only one collection from California has 



TABLE 1 

Comparisons of the Diagnostic Characteristics of B. carinatus, 
B. arizonicus, and Their Fi Hybrid 



Number of nodes of culms 

Pubescence of top node 

Pubescence of upper culm leaf 

Maximum number of branches at node 

of panicle 

Maximum number of florets per spikelet 

Length second glume 

Length second lemma 

Ratio, glume/lemma 

Pubescence of second lemma 

Lobes of upper lemmas 



B. carinatus 
(no. 5) 



5-6 
+ below 

+ lower 



5-8 

8-9 
14.3 
16.4 

0.87 
+ + 
throughout 
small 



Fi hybrid 
(no. 125 X no. 5) 



5 

+ + 
throughout 
+ + upper 
+ lower 

4-5 

7-9 

16.0 

15.2 

1.05 

+ + margins 

+ back 
intermediate 



B. arizonicun 
(no. 125) 



4-5 

+ 

throughout 

+ upper 

lower 

4-6 

6-7 

12.4 

12.3 

1.01 

+ + margins 

0-+ back 

prominent 



been counted, that from Tehachapi (no. 128), but the external morphology 
of the other California specimens cited leaves no doubt as to their specific 
identity. At Davis, California, the northern limit of its known range, B. 
arizo7iicus is obviously a recent introduction along the railroad tracks, and it 
is very likely introduced at the San Joaquin Valley localities also. 

The particular strain of B. arizonicus used for hybridization was one of the 
most vigorous and latest in flowering of all. As recorded elsewhere (Harlan, 
1942; Stebbins and Tobgy, 1944), the no. 5 strain of B. carinatus is also a 
very vigorous one. It is about intermediate or somewhat late in flowering 
compared to other Berkeley strains, but earlier than most of the montane 
types. It is distinguished from most other B. carinatus strains from Berkeley 
by its rather large spikelets and lemmas; the average length of the lemmas 
and awns in B. carinatus as a whole is not very different from that in B. 
arizonicus. The chief distinguishing characteristics of the two parental strain* 
are given in Table 1. 

No hybrid seeds were obtained out of 42 florets emasculated when B. ari' 
zonicus was used as the pollen parent on B. carinatus. In the reciprocal cross, 



Vol. XXV] STEBBINS-TOBGT-HABLAN : HYBRIDS IN BBOMUS II 313 

B. arizo7iicus 5 x -S. carinatus ^, 37 seeds were obtained out of 40 florets 
emasculated. Of these, 10 were sown in September, 1942. Four were lost 
through an accident, and six were planted in the field in November. Of these 
4 turned out to be hybrids. Morphologically, they were intermediate between 
their parents, except that in number of florets per spikelet they were nearer 
their B. carinatus parent, and in relative size of glumes and lemmas, like B. 
arizonicus (Table 1). They proved to be completely pollen and seed sterile, a 
condition which would be expected from the cytological situation described 
below. Three of them died during May, 1943, even before their annual B. 
arizonicus parent. A fourth showed some tendency to grow out after being ciit 
back, as does B. carinatus, but died during the fall of 1943. . 

CYTOLOGY OF THE PARENTS AND HYBRID 

1. B. carinatus: The meiotic chromosome behavior of strain no. 5 of B. cari- 
natus used in this study has been reported earlier (Stebbins and Tobgy, 
1944). The main features may be outlined here. At diakinesis and I-meta- 



♦ftt*J^»V 






: >.«• • 






^^ ^^ 



,-'' 



Figs. 5-7, chromosome pairing at first metaphase in: 5, B. carinatus; 6, B. arizonicus ; 
7, B. arizonicus x carinatus. All reproduced X 1300. 



314 



CALIFORNIA ACABEMY OF SCIENCES [Pkoc. 4th 



SER. 



8 






•To 5 






% % 



• • 






U^ \ 



10 



m ' 



< • I 



w > 



'rv 









1 1 




^ 



^.<| 



f; 



2u 









13 






Figs. 8-13. Photomicrographs showing chromosome behavior, all reproduced X 800. 
8, B. carinatus, I metapliase; 9, B. arisoiiicus, diakinesis; 10 and 11, B. arisonicus x cari- 
natuti, I mctaphase: 12 and 13, B. arisonicus x carinatiis, I anapliase. 



Vol. XXV] STEBBINS-TOBGY-HARLAN : HYBEIBS IN BEOMUS II 315 

phase there were always 28 bivalents ; no univalents or multivalents were ever 
seen. Seven bivalents could be distinguished by their conspicuously large size, 
while the remaining 21 bivalents were uniformly smaller and may therefore 
be termed medium-sized (figs. 5 and 8). At I-metaphase most of the bivalents 
had two chiasmata, one in each arm, and occasionally some of the large biva- 
lents had three chiasmata. The majority of the bivalents, large and medium- 
sized, were therefore pairing in both arms, although rarely one or two of them 
were pairing in one arm only. In 9 per cent of the I-anaphase cells and in 1 per 

TABLE 2 
Chromosome Pairing at Meiosis in Bromus arizonicus X carinaius 



Combination 


No. of cells* 


Per cent 










niiii 


mjj 


m, 


ii 






7 


14 


14 


7 


12(2) 


33.5 


6 


15 


15 


7 


6 


16.2 


5 


16 


16 


7 


6 


16.2 


4 


17 


17 


7 


9(2) 


24 3 


3 


18 


18 


7 


2(1) 


5.4 


2 


19 


19 


7 


2(1) 


5.4 


Total . . . 








37 


100.0 













' The numbers in parentheses refer to 6 cells in which two of the chromosomes regularly forming a mji ap- 
peared instead as 2mi. 



cent of the Il-anaphase cells one inversion bridge and a fragment were seen. 
Almost 99 per cent of the tetrads had 4 normal microspores ; in the remaining 
1 per cent, one or two micronuclei were also present. There was 81.0 per cent 
stained pollen. 

2. B. arizonicus: At diakinesis and I-metaphase of B. arizonicus there were 
always 42 medium-sized bivalents (figs. 6 and 9) ; univalents and multivalents 
were significantly missing. There was no differentiation into large and small 
bivalents and none of the bivalents could be classified in the large size category 
characteristic of seven bivalents in B. carinatus. In almost half of the I-meta- 
phase cells all the bivalents were paired in both arms and in the other half 
one, two, or three bivalents were paired in one arm only. There was less in- 
version hyljridity exhibited in the B. arizonicus plant examined than in that 
of B. carinatus. Only 1 per cent of the I-anaphase cells and 0.4 per cent of the 
Il-anaphase cells showed one chromatid bridge and one fragment. About 2.5 
per cent of the tetrads had one or two micronuclei in addition to the 4 normal 
microspores, but the remaining 97.5 per cent had 4 large microspores only. 
There was 90 per cent stained pollen. 

3. B. arizonicus x B. carinatus. The F^ hybrid plants (2n = 70) between B. 
arizonicus (2n = 84) and B. carinatus (2n = 56) received 42 medium-sized 



31G 



CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 



chromosomes from B. arizonicus and 21 medium + 7 large chromosomes from 
B. carinatus. Thirty-seven cells at I-metaphase were completely analyzed in 
the hybrid (Table 2). 

Pairing of the parental chromosomes followed a comparatively simple 
scheme which may be described as follows : 

1. The 7 large chromosomes from B. carinatus were always seen as 7 unpaired univalents 
(li in the table). 

2. At least 2 sets of 7 medium chromosomes from B. arizonicus also appeared as 14 un- 
paired univalents (m, in the table). 

3. At least 14 medium bivalents were almost always seen (mn in the table) ; these being 
formed from the consistent pairing of 2 sets of 7 medium chromosomes from B. arizonicus 
with 2 corresponding sets from B. carinatus. 

4. A maximum number of 7 trivalents was formed (mm in the table) ; these resulted 
from partial or complete homology between one set from B. arizonicus, another from the 
same species, and a third from B. carinatus. When the maximum number of 7 trivalents 
was formed, there were 14 medium bivalents, 14 medium univalents and 7 large univalents 
in the cell. With decreasing numbers of trivalents, the number of bivalents and univalents 
was correspondingly increased. A typical I-metaphase cell is shoA^Ti in figure 7, and two 
other cells in figures 10 and 11. The percentage of cells with these verying numbers of 
trivalents is given in the last column of Table 2. A slight reduction in the normal pairing 
in a few cells was found (cf. footnote. Table 2). 

TABLE 3 
Sporad Analysis in B. arizonicus X carinatus 



Number of cells 


Number of micronuclei 


Totals 


Per cent 


in sporad 


7-11 


12-16 


17-21 


4 cells 

5 cells 


29 

15 
8 
2 


51 
23 
13 

4 


27 
6 
4 

1 


107 
44 
25 

7 


58.5 
24.0 


6 cells 


13.7 


7 cells 


3.8 






Totals 

Percentages 


54 
29.5 


91 
49.7 


38 
20.8 


183 


100.0 



There was good evidence of extensive inversion hybriditj' in the F^. This 
was shown by bridge-fragment configurations in 48.5 per cent of the cells at 
I-anaphase and by 16.5 per cent of the cells at Il-anaphase. Usually only one 
bridge and occasionally two were seen at I-anaphase, and only once was a cell 
with three bridges encountered (fig. 13). All the bridges at Il-anaphase were 
genuine second division bridges and not relics from the first division. There- 
fore the extent of inversion hybridity in the hybrid may be represented as 
one bridge in 65 per cent of the pollen mother ceUs. It could not be always 
determined whether the inversions were confined to the bivalents or were 
found in the trivalents also, but evidence was obtained of bridges in both 
types of configurations. Therefore it is likely that the two groups of parental 



Vol. XXY] STEBBINS-IOBGY-EABLAN : HYBBIDS IN BEOMUS II 317 

chromosomes whicli regularly pair differ by inversions as well as the two 
partly homologous sets £rom B. arizonicus which enter into the trivalents. 
Since heterozygosity for inversions was found to a small degree in both 
parents, it is reasonable to suppose that most of the pairing sets differ by at 
least a few inversions. 

The majority of the univalent chromosomes divided equationally and the 
two daughter univalents separated to the two poles at I-anaphase, but occa- 
sionally some of the univalents, though already divided, would fail to undergo 
the anaphasic separation and would pass entirely to one pole or the other 
(figs. 12 and 13). There was excessive chromosome lagging at the second 
division, which, together with that at the first division, may best be repre- 
sented by the very complex condition of the sporads (Table 3) . 

"With such conditions prevailing in the sporads the repeated observation of 
complete sterility of both pollen and ovules is the expected condition. 

DISCUSSION 

Specific distinctness of B. arizonicus. The experimental evidence presented 
here shows that, because of the extreme abnormality of meiosis in their Fi 
hybrid, and the complete inviability of both pollen and ovules, B. arizonicus 
and B. carinatus are completely incapable of interbreeding, that is of ex- 
changing genes. Therefore, they would be recognized as distinct species under 
any modern definition of this entity (cf. Dobzhanskj^ 1941, p. 373). Further- 
more, the cytological evidence indicates that B. arizonicus would be equally 
intersterile with any other octoploid member of the B. carinatus complex 
(cf. Stebbins and Tobgy, 1944). The two species clearly are not made up of 
the same chromosomal material. Out of the haploid complement of 42 in B. 
arizonicus, 21 chromosomes are nearly or quite homologous to 21 of the haploid 
set of 28 in B. carinatus, 7 are partly homologous with 7 of these 21 carinatus 
chromosomes and 14 are entirely different. The resemblance between B. ari- 
zonicus and B. carinatus, therefore, lies principally in their common posses- 
sion of 21 pairs of medium-sized (m) chromosomes. 

Cytological constitution and possiMe origin of B. arizonicus. In another 
paper (Stebbins and Tobgy, 1944) it was shown that the 21 medium-sized 
chromosomes (m) iiiB. carinatus are almost completely homologous with the 
21 chromosomes which form the entire haploid complement of the South 
American species, B. catharticus Valil (B. unioloides HBK). In that paper, 
these sets were designated AABBCC, so that the genome formula of the various 
octoploid members of the B. carinatus complex was written as AABBCCLL, 
LL representing the sets of 7 large chromosomes. The present evidence shows 
that B. arizonicus possesses the AABBCC set, perhaps somewhat modified, but 
lacks the 7 pairs of large, or the LL chromosomes. Of its other three sets, one 
may be designated as Co C2, because of its similarity to one set of the catharti- 
cus group, this partially homologous set being arbitrarily designated as the 
CC one. The remaining two sets of B. arizonicus may be designated DDEE. 



318 CALJFOKXIA ACADEMY OF SCIENCES [Proc. 4th ser. 

Thus the g-enome formula of B. arizonicus may be written AABBCjCiCaCo 
DDEE. It should be clearly understood that this- formula is used only to 
express pairing relationships ; the actual genie content of the genomes prob- 
ably varies considerably from species to species, and they undoubtedly differ 
in the various species by minor structural variations. 

This cytological constitution indicates that B. arizonicus is an allopoly- 
ploid derived bj^ chromosome doubling from a hybrid between B. catharticus 
or a close relative of it and some other species with 21 pairs of medium-sized 
chromosomes. The identity of this other parent is not yet established, but 
based on the morphological differences between B. arizonicus and B. cai'inatus, 
its main features can be postulated. They are as follows : 

1. It should be an annual, preferably found in desert or at least semiarid situations. 

2. Its panicle branches should be slender, but should be glabrous or finely short-pubescent, 
rather than coarsely scabrous as in B. catharticus. 

3. Its glumes should be elongate, definitely exceeding the lower lemmas. The outer glume 
should have chiefly 3 nerves. 

4. Its lemmas should be rounded on the back or weakly keeled, should be long-hirsute 
near the margins, conspicuously bidentate at the apex, and long-awned. 

5. It should form a maximum of 7 bivalents in hybrids with B. catharticus or B. carinatus. 

The morphological characteristics listed here are found in only one species 
of Bromus known to us, B. Trinii Desv. This species also has the required 21 
pairs of medium-sized chromosomes, as determined by Knowles (1944) as 
well as by the senior author. Its pairing relationships with B. catharticus and 
B. carinatus have not yet been tested. The hypothesis may, however, be ad- 
vanced that B. arizonicus is an allopolyploid derived from B. catharticus and 
B. Trinii or close relatives of these species. It is hoped that this can be tested 
in the near future by appropriate hybridization experiments. The hypothesis 
involves some difficulties from the historical point of view, since both of the 
putative parents are generally believed to be native only to South America, 
and to be recent introductions into the area now occupied by B. arizonicus. 
This problem, however, had best be considered after the evidence in favor of 
the proposed hypothesis has become more definite. 

There is one cytological discrepancy found in the pairing behavior of the 
chromosomes of B. arizonicus in the pure species as compared with their be- 
havior in its hybrid with B. carinatus. The trivalents found in the hybrid 
indicates definitely a partial homology between two of the sets in B. arizonicus. 
That they could not possibly consist of two sets from B. carinatus and one 
from B. arizonicus is evident from the fact that in another hybrid, B. carina- 
ius X B. mollis, the chromosomes nearly all remain unpaired, showing that 
there is little or no homology between any of the four sets found in B. carinatus 
(Knowles 1944) . The difficulty is that these two partially homologous sets of B. 
arizonicus should occasionally form quadrivalents in the pure species. How- 
ever, in several different strains, examined by each of the three authors, the 
presence of 42 bivalents was observed with strikingly consistent regularity, 



Vol. XXV] STEBBINS-TOBGY-HARLAN : HYBRIDS IN BROMUS II 319 

and the partial homology between two of the sets was not even suspected until 
the cytology of the hybrid was studied. Two explanations for this phenomenon 
are suggested. The first is that advanced by Miintzing and Prakken (1940) 
for a similar behavior in Phleum pratense, which was also found in the same 
species by Myers (1941) . They assume that in Phleum and other genera there 
is a "special genotypically controlled tendency to bivalent formation" (p. 494) . 
The other explanation is that of differential affinity, or preferential pairing, 
advanced by Darlington (1927, p. 198) to explain the lack of multivalents in 
the allopolyploid Primula kewensis, and by Skirm (1942), to explain the 
difference in pairing found in two different tetraploicls of Tradescantia, both 
obtained from the same diploid hybrid. From the data now at hand, there is 
no way of concluding which is the best explanation for the situation in Bromus 
arizonicus. 

In conclusion, the following points have arisen from this study which are 
of particular interest to students of the taxonomy of the Gramineae. In the 
first place, whatever the second parent of B. arizonicus is, it most certainly 
belongs neither to the section Ceratochloa nor to the section Bromopsis, as 
postulated for the second parent of the octoploid B. carinatus and its relatives 
(Stebbins and Tobgy, loc. cit.) . The morphological characteristics listed above 
would immediately remove the species from the former section, while in the 
latter the chromosomes are large, like the LL set of B. carhiatus (Stebbins 
and Love, 1941). Both B. carinatus and B. arizonicus, therefore, are allopoly- 
ploids from intersectional hybrids, but the two, sections thus connected by 
allopolyploidy are different in each case. Therefore the cytogenetic evidence 
entirely supports the recognition of Bromus as a single genus, rather than the 
separation of Ceratochloa as generically distinct, as has been done by a num- 
ber of agrostologists. It speaks even more emphatically against the splitting 
of Bromus into several different genera, as was proposed by Nevski (1934). 

In the second place, the particular type of relationship which exists between 
B. carinatus and B. arizonicus will probably be found to be rather frequent 
when more genera of this family are studied cytogenetically, since polyploidy 
is so prevalent in them. They are a typical example of species which are mor- 
phologically very similar, and cytogenetically partly very similar and partly 
very different. Their similarities are due to their common possession of the 
B. catharticus set. Their morphological differences represent, in a much di- 
luted form, the differences between two different sections of the genus. Both 
of these sections diverge from typical Ceratochloa in their much less strongly 
keeled lemmas and much longer awns ; hence both B. arizonicus and B. carin- 
natus diverge from B. catharticus in the same respects. B. carinatus and 
B. arizonicus may therefore be spoken of as allosyngenic amphidiploids (or 
allopolyploids), since they differ in respect to one of their original parents, 
and have the other in common. The likelihood is that in all genera possessing 
large polyploid complexes, allosyngenic amphidiploids form one of the chief 
sources of taxonomic difficulty. 



320 CALIFORNIA ACAVEMY OF SCIEXCES [Pkoc. 4th ser. 

SUMMARY 

1. A new species, B. arizonicus, is described, based on B. carinatus variety 
arizonicus Shear. This species has the somatic chromosome number 2n = 84, 
as compared with 2n = 56 for B. carinatus. 

2. Tlie hybrid, B. arizonicus x B. carinatus, can be obtained only when B. 
arizonicus is the female parent. It is morphologically intermediate between 
its parents and is completely sterile. 

3. At meiosis this hybrid regularly shows a maximum of 7 trivalents, with 
the rest of the chromosomes existing as bivalents and univalents. The 7 large 
chromosomes obtained from B. carinatus have no homologues in B. arizonicus, 
and therefore are always unpaired, as are also 14 medium-sized chromosomes 
from B. arizonicus. 

4. The presence of bridge-fragment configurations in either the first or the 
second division of meiosis in 65 per cent of the pollen mother cells indicates 
that the chromosomes of two or more of the pairing sets differ hy a number of 
inverted segments. 

5. The cytogenetic evidence amply demonstrates the specific distinctness of 
B. carinatus and B. arizonicus. Their genomic formulae may be written as 
follows : for B. carinatus AKB^GS^J-iJj^, for B. arizonicus AABBC1C1C2C2 
DDEE. The A, B, and Ci sets constitute the chromosomal complements of B. 
catharticus. It is possible that B. arizonicus possesses this complement in a 
slightly modified form. 

6. The hypothesis is advanced that B. arizonicus is an allopolyploid derived 
from doubling of the chromosome number in a hybrid between B. catharticus 
or a close relative of that species and some other species with 21 pairs of 
medium-sized chromosomes, not belonging to the subgenus Ceratochloa. 

7. The cytogenetic evidence here presented supports the retention of 
Bromus, sens, lat., as a single genus. 

8. The relationship of B. carinatus and B. arizonicus is described as that of 
allosyngenic amphidiploids. This type of relationship is believed to be a com- 
mon source of taxonomie difficulty in genera which have highly developed 
polyploid complexes. 



Vol. XXV] STEBBINS-TOBGT-HARLAN : HYBRIDS IN BEOMUS II 321 

LITERATURE CITED 

Darlington, C. D. 

1937. Eecent Advances in Cytology, 2nd ed. 671 pp. Philadelphia, Blakiston's. 

Harlan, J. R. 

1942. A survey of the genetic variability in the Bromus carinatus complex. Thesis (Ph.D.), 
Univ. of California. 

Knowles, p. F. 

1944. Interspecific hybridizations of Bromus. Genetics 29: 128-140. 

MiJNTZiNG, A., and R. Prakken 

1940. The mode of chromosome pairing in Phleiim twins with 63 chromosomes and its 
cytogenetic consequences. Hereditas 26:463-501. 

Myers, W. M. 

1941. Meiotie behavior of Phleum pratense, Pldeum siCbulaium, and their Fi hybrid. Journ. 
Agr. Res. 63:649-659. 

Nevski, S. a. 

1934. Sched. Herb. Flor. Asiae Mediae, in Acta Univ. Asiae Mediae, Tashkent, Ser. VIII 
b, Bot., Fase. 17:14-20. 

Shear, C. H. 

1900. Studies on North American grasses. A revision of the North American species of 
Bromus occurring north of Mexico. U. S. Bept. of Agric. Div. Agrost. Bull. no. 23. 
66 pp. 

Skirm, G. W.' 

1942. Bivalent pairing in an induced tetraploid of Tradescantia. Genetics 27 : 635-640. 

Stebbins, G. L. Jr., and E. M. Love 

1941. A Cytological study of California forage grasses. Am. Jour. Bot. 28 : 371-382. 

Stebbins, G. L., Jr., and H. A. Tobgy 

1944. The cytogenetics of hybrids in Bromus. I. Hybrids within the section Ceratochloa. 
Am. Jour. Bot. 31:1-11. 

Swallen, J. R. 

1942. Gramineae, in Kearney and Peebles, Flowering Plants and Ferns of Arizona. U. S. 
Dept. Agric. Misc. Publ. no. 423. 1069 pp. 



PROCEEDINGS 

OF THE 



Uj 



CALIFORNIA ACADEMY OF SCIENCES V. 

FoLirtli Series 

Vol. XXV, No. 14, pp. 323-356, pis. 28-3G, 3 charls, 3 hislogianis 

November iG, 1944 




ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 



No. 14 

THE IMPORTANCE OF FIELD HYBRIDS IN 
DETERMINING SPECIES IN THE 

GENUS CEANOTHUS 

BY 

HOWARD E. McMINN 

Professor of Botany, Mills College 

«VERY FIELD BOTANIST, liaviiig tlie faculty of careful observation, is repeat- 
edly impressed v^ith the number of apparent hybrids which occur in 

I certain genera of seed-bearing plants. The genus Ceanothus is one of these 
genera. According to Katherine Brandegee, Dr. C. C. Parry was the first to 
call attention to the prevalen'ce of natural hybrids in this genus. Mrs. Brande- 
gee in her "Studies of Ceanothus" (1894) discussed the various hybrids which 

^she had observed in the field and herbaria. She was of the opinion that most 
of the hybrids between members of the same section were fertile but she gave 
no supporting evidence. She stated that, "The only infertile hybrid, within 
its section, known to me is No. 69, C. incanus x papillosus. In this the ovaries 
are more or less abortive and no fruit was formed." If we accept the sterility 
test for distinct biological species, the two parents, C. incanus and C. pajyil- 

Hosus, in this cross belong to two different species complexes (cenospecies of 
the experimentalist). John Thomas Howell (1940a) "presents the case of hy- 
bridization between certain species of the section Cerastes. His conclusion 
is worthy of note : "My general conclusion is that anyone, interested in the 
taxonomy and the possible evolution of entities in the subgenus Cerastes of 
Ceanothus, cannot disregard the probable effect of hybridization, and that 
some of our puzzling entities can be more definitel}^ limited as taxonomic 
units with the proper study and interpretation of intermediates of suspected 

[ 323 ] 



324 



CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 



hybrid orijiin." ^lany botanists have been reluctant to accept the importance 
of hybridization in the evolution of the numerous forms of Ceanothus and 
some ignore it entirely. However, anyone who has done mue'h serious field 
study of this genus will come to agree with Howell's general conclusion. 

It is the opinion of the writer tluit the number of true biological species 
(ecospecies of the experimentalist) existing at a given time in any genus of 
plants cannot be known until extensive experimental studies for testing the 
fertility of hybrids can be carried out. Until more logical methods for the 
determination of true biological or distinct species are set forth, botanists 
will do well to test out the numerous variants w'ithin a genus by putting 
them through the biological sterility sieve advocated by Clausen, Keck, and 
Hiesey (1939) shown in table 1. 

Table 1. Tlir conci))! of sprrirs. 



Degree (if 
separation 


Hylirids fertile, 

second generation 

vigorous 


Hybrids partially 

sterile, 

second generation 

weak 


Hybrids sterile 
or none 


\ Internal 

\ 
External 


In different 
environments 


Distinct subspecies 
ECOTYPES 


Distinct species 

ElCOSPECIES 


Distinct 

species cn))iplc.res, 
CENOSPECIES 


In the same 
environment 


Local variations 
of one species 

BIOTYPES 


Species overlapping 

in common territory 

(with hybrid swarms) 



Experimental methods require much time and labor, and it is evident to 
botanists who study woody plants that the adoption of the experimental 
method will be slow, and until such adoption other methods now employed 
by workers will continue to be used. However, by being aware of the impor- 
tance of experimental methods, field workers will become more alert to the 
existence of natural hybrids and to geographic and ecological variations. 
By recognizing field hybrids and applying to them the same principles used 
by Clausen, Keck, and Hiesey in determining experimentally cenospecies, 
ecospecies, ecotypes, and biotypes, it has been possible to gain much helpful 
information in determining species complexes (cenospecies), distinct or bio- 
logical species (ecospecies), subspecies (ecotypes), and local variations (bio- 
types) in the genus Ceanothus. 

In this genus there are two rathei- well-defined sections or sub-genera, 
Euceanothus with leaves alternate and fruit not horned, and Cerastes with 
typically opposite leaves and horned fruit ( Plate 28) . In a recent publication, 
by Van Rensselaer and the writer (1942) the Euceanothus section includes 
thirty-three binomials (so-called species), seventeen trinomials (varieties), 
many minor variations, four named natural hybrids, and many other hybrids. 



Vol. XXV] McMIKN: FIELD HTBEIDS IN CEAXOTHUS 



325 



The Cerastes section includes twenty-three binomials, nine trinomials, many 
minor variations, four named natural hybrids, and other hybrids. Three 
named natural hybrids between members of the two sections are also included. 
The writer believes that the number of existing distinct species (ecospecies) 
is much fewer than represented by the number of binomials given in manuals, 
and that more subspecies (ecotypes) and local variations (biotypes) will be 
recognized when sufficient knowledge of the behavior of the hybrids is known. 
Some of this behavior can be learned by careful observation in the field. 

INTEKSECTIONAL HYBRIDS 

If observed hybrid plants fail to set fruit year after year, one may assume 
with some degree of certainty that they are sterile, and that their parents 
belong to distinct species complexes (cenospecies). This situation is partially 

CHART 1 

Wfttural HTbridB betireen Snaciea 
of the Sections Ceraa tea and Sueeajiothus 



45, C. rlgidufl 

43. C, purpureus 

t?. C. prostratus 

7ar. 1 




61. X C. Vei t ch ianus 

T C. serrulatua 
b^. X C. rugosus 



ames Roof- 



-3-^-1 



". griaeus 25» 

3. cordulatus 14. 

Tolutlnu3 1. 

C. sninosiis 11. 

j^Ji^-C. oli^anthus 21. 

" C, 8orediatU9 22. 

~- - -C. thyrslflorus 26. 



exemplified in the intersectional cross between C. prostratus and C. cordulatus 
which has been given the designation x C. scrrulatus. This hybrid was first 
collected in 1926 from a small colony of not more than four or five plants in 
a shallow draw on a ridge between Emerald Bay and Cascade Lake, Eldorado 
County, California. To my knowledge it has never been found at any other 
locality. It appears that a single hybrid plant may have been formed and that 
the other plants have resulted from natural layering. The evidence for con- 
sidering the two parents of this hybrid as belonging to different species com- 
plexes would be more conclusive if more sterile hybrid plants were known. 
Associated with this hybrid is C. prostratus, C. cordulatus, and C. velutinus. 
The prostrate habit and the presence of sunken stomatal pits on the underside 
of the leaves relate it to C. prostratus, but the predominantly thinner and 
alternate leaves, the small deciduous stipules, and short racemose flower clus- 
ters relate it to C. cordulatus. The presence of both opposite and alternate 
leaves and the very pale blue to nearly white flowers seem to be intermediate 
characters. The plants have never been known to set fruit. 

Two other field hybrids and six garden hybrids have been observed by the 
writer between members of the two sections (Chart 1). Only one of these 
hybrids, the garden hybrid C. James Roof, has ever been known to set fruit. 
The seeds of this hybrid when tested failed to germinate. The parents were 
C. purpureus and probably one of the following : C. oliganthus, C. spinosus. 



326 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

C. thyrsiflorus, or C. sorediatus. The two other observed field hybrids between 
members of the two sections are x C. rugosns and x C. Veitchianus. x Ceano- 
thus rugosns appears to be a hybrid between C. prostratns and C. velntinus. 
Specimens have been obtained from two localities, one from the lava beds near 
Egg Lake, Modoc County, California, and the other from near Truckee, Ne- 
vada County, California. M. S. Baker and F. P. Nutting collected the speci- 
mens from INIodoc County in 1894. Baker in a note states that the plant was 
a ''decumbent form of C. vehitinus — probably a hybrid of C. prostratns and 
C. vehitinus." The specimens from Truckee were collected by C. F. Sonne in 
1890. Sonne reported that the plant sets flowers sparingly and that it does 
not fruit at all. It would seem then, that the two parents, C. ijrostratus and 
C. velntinus, definitely belong to different species complexes. 

X Ceanothus Veitchianus appears to be a hybrid between C. grisens and 
probably C. rigidus. To my knowledge it was first collected by William Lobb 
in California, probably near Monterey where C. grisens and C. rigidus grow. 
Lobb probably collected seeds from C. grisens and C. rigidus and from this 
collection appeared the hybrid x C. Veitchianus in the Veitch and Sons nurs- 
eries of Exeter and Chelsea, England. I have seen no specimens collected from 
the field. The few garden hybrids of x C. Veitchianus which I have observed 
have never set fruit. It would appear from the sterility test that C. rigidus 
and C. griseus belong to different species complexes. 

The genetic barrier between the two sections must be something more than 
chromosome number, because the diploid number for all species examined in 
the two sections by Nobs (1941) is 24 (Plates 29 and 30) . Nobs also examined 
six hybrid plants of different parentage and all were found to have the same 
number of chromosomes as the parents, namely 2n = 24. 

INTRASECTIONAL HYBRIDS 

If obser\'ed hybrid plants are partially sterile and set fruit which produce 
seeds that germinate poorly or whose seeds produce a weak second generation, 
we may assume that the hybrid plants were produced by the crossing of dis- 
tinct species (eeospecies of the experimentalist). If the hybrids proved to be 
fertile and the second generation vigorous, we may then assume that their 
parents belonged to distinct subspecies (ecotypes of the experimentalist) or 
that they were local variations of one species (biotypes of the experimentalist ) , 
depending upon whether they occurred in different environments or in the 
same environment. Examples of such hybrids have been observed many times 
in the field between members of the same section of the genus wherever tliey 
overlap in their distribution. 

Section Euceanothus 

In the section Euceanothus, the evidence from hybridization points toward 
the existence of more cenospecies and eeospecies than in the section Cerastes. 
In tliis section {Euceanothu.s) hybrids liave been observed between most 



Vol. XXV] McMIXX: FIELD EYBBIDS IX CEAXOTHUS 



327 



of the species that occupy the same area and that flower at the same time. 
In some cases where two species, as C. cordulatus and C. diversifolius or as 
C. thyrsiflorus and C. incanus, occur together only an occasional hybrid plant 
is formed and that plant usually appears to be totally sterile. In these cases 
we may assume that C. cordulatus and C. diversifolius belong to different 
species complexes (cenospecies) and also that C. thyrsiflorus and C. inca- 
nus belong to different species complexes. Since neither C. cnrdulatiis nor 



1. C. velutinus 
la. var. laevigatu: 

2. C, enericanus 

3. C. ovatUB 

4. C. sangulneaB 

5. C. serpyllifoliua 

6. C. microphjllus 

7. C. integerrimu? 
(riB. white or "D 

8. C, parrlfolius. \ 
(FlB. ■blue) 

9. C. Kartlnil 

10. C. Faljneri 

11. C. spinosus 
(TIb. pale blue)^ 

1?. C. 1-eucoderiiis 

13. C. iBcanus 

14. C. cordulatus 

15. C. buxlfclius 

16. C. rendleri 

17. C. dopressua 



CHAET 2 

K„f„rpl I'yhri.^^.. '?r»---^':" ^^'■cie- of the Sectlog F.u:canothu3 

yio/rerg blu9 
__— C. arboreua 




, diTersifoliuj 33» 



C. diversifolius are ever found in nature growing with either C. thyrsiflorus 
or C. iyicanus, no evidence from field hybrids has been obtained regarding 
their interrelationships. From morphological characters it would seem that 
G. cordulatus and C. incanus belong to the same species complexes, and that 
that complex is different from the one to which C. thyrsiflorus or the one to 
which C. diversifolius belongs. 

In the natural hybrids observed between C. cordulatus and C. velutinus 
and between C. thyrsiflorus and C. papillosus on the other hand, many varia- 
tions occur which appear to be only partially sterile and which appear to be 
members of a hybrid swarm, thus indicating that probably these two entities 
are distinct species (ecospecies). If these hybrids were quite fertile and their 
second generation vigorous, more local variations would be expected than are 
now found, and then these four entities would be considered subspecies (eco- 



328 CALIFOEXIA ACADEMY OF SCIEXCES [Proc. 4th ser. 

types) or local variations of one species (biotypes). depending upon whether 
they normally occurred in different environments or in the same environment. 
Most of the other crosses observed in the Euceanothus section, and recorded 
in chart 2, appear to follow the same general patterns as given in the examples 
just cited. 

A summary of the recorded natural hybrids in this section shows a total of 
thirty, involving twenty-one species. Ceanothus thyrsiflorus, which in its dis- 
tribution contacts more species than any other entity in the Euceanothus 
section, has entered into nine crosses, whereas the other species with fewer 
contacts have been involved in from one to five crosses. Three of the thirty 
crosses have been given hybrid binomial designations because they have been 
considered as species or varieties by other workers. One other hybrid has been 
named because of its presence in the nursery- trade. Since much confusion 
exists in the literature about these hybrid entities, it seems worth while to give 
a brief account concerning them. 

Named Hybrids of the Section Enccanoihus 

X Ceanothus mendocinensis appears to be a hybrid between C thyrsiflorus 
and C. velutinus var. laevigatus. Plants of this cross have been collected in 
several scattered locations in Mendocino County, California, and in adjacent 
counties where the two parents overlap in their distribution. They usually 
occur as individual plants with one or both parents, never in colonies. The 
fertility of the seed has not been tested, but since so few plants have been 
found where the parents occur together, and since the hybrids appear to be 
rather uniform in those characters which are intermediate between those of 
the parents, it would appear that the plants were first-generation hybrids and 
that the parents belong to distinct species. 

X Ceanothus Lorenzenii has been considered to be a variety of C. velutinus 
by some botanists. It occurs in the Sierra Nevada of California from Kern 
County northward to Siskiyou County and eastward in Nevada. Apparently 
this hybrid is the result of a cross between C. velutinus and C. cordulatus 
(Plate 31). Field studies made about five miles north of Markleeville, Alpine 
County, California, in the summers of 1941 and 1942, furnished the most 
convincing evidence. On an extensive old burned-over area there occurred 
C. velutinus, C. cordulatus, and C. prostratus. The hybrid plants were found 
only in areas where C. velutinus and C. cordulatus were closely associated. 
In areas where one species occurred to the exclusion of the other no hybrids 
were observed. Most of the hybrids resembled the species C. velutinus more 
than C. cordulatus. The stiff branches and smaller leaves without varnisli 
above, however, gave evidence of the part C. cordulatus had in its origin. 
A few plants were found which resembled C. cordulatus more than C. velu- 
tinus. It appeared from the field studies that the hybrid jilants were mostly 
sterile. If they were quite fertile one would expect more individuals inter- 
mediate between the two parents and the hybrid (Histogram 1 ) . I believe most 



Vol. XXV] McMINX: FIELD HYBRIDS IN CEANOTHUS 



329 



of the specimens of x C. Lorerizenii are first generation hybrids. Nobs, after 
field and eytological studies, also concluded that C. velutinus var. Lorenzenii 
Jepson is a hybrid between C. velutinus and 0. cordulatus (Plates 32 and 33). 
X Ceanothus Lohhianus is probably the result of a cross between C. griseus 
and C. dcntatus. It was first grown in England from seed collected by William 



no. of 
plants 



Histogram 1 




1 2 3 4 5 
C. velutinus Dougl. 



index 
6 7 8 9 10 ^^'^^^ 

C. cordulotus Kellogg 



X C. Lorenzenii (Jepson) McMinn 

Histogram 1 represents the distribution of jDlants into 11 groups based upon an index 
value determined by studying 10 characters of C. cordulatus. The plants were taken from 
a random sampling in an area where C. cordulatus and C. velutinus overlapped in their dis- 
tribution. Plants A\ith all 10 characters were classified as C. cordulatus; plants with none 
of these characters were classified as C. velutinus; those with 4, 5, and 6 characters were 
classified as x C. Lorenzenii; and those with 1, 3, 7, or 8 characters are unnamed hybrids. 
No plant in the sampling was found with 2 or 9 characters. As the groups approach the 
plants more closely resemble C. velutinus. 

(x = hybrid) 

Lobb in California, probably in the region of Monterey where C. griseus and 
C. dentatus occur. A few plants collected as natural hybrids in the Monterey 
region and now growing in California gardens approximate the type specimen 



330 



CALIFOEXIA ACADEMY OF SCIENCES [Proc. 4th ser. 



of the original C. Lohhianns. Seeds have been collected from these hybrids 
but their fertility is not known. Since so few plants of this hybrid occur either 
in the wild or in any known garden collection it would seem that the hybrid 
is only partially fertile and therefore the parents, C. griseus and C. dentatus, 
belong to distinct species. 

X Ceanothus regius is a hybrid between C. ihyrsiforus and C papillosus. 
It has been observed several times wherever the two parents overlap in their 



no. of 
plants 



Histogram 2 



40- 




12 3 4 
0. papillosus T a G. 



6 7 8 9 10 
C. thyrsiflorus Esch, 



index 
value 



X C. regius 

Histogram 2 represents the distribution of plants into 11 groups based upon an index 
value detorniined by studying 10 characters of C. tliyi'sifloni.'^. The plants were counted from 
a random sampling in an area Avhere C. thyrnflorufi and C. papillosus overlapped in their 
distribution. Plants with 10 characters were classified as C. thyrsiflorus; those with none 
of these characters were classified as C. j^opillosus ; those with 4 or 5 characters were classi- 
fied as X C. rcyius ; and those with 2, 6, or 8 characters were classified as unnamed hybrids. 
No plant in this sampling was found with 1, 3, 7, or 9 characters. 



distribution in San Mateo and Santa Cruz counties, California. The hybrid 
give evidence of being only partially sterile, consequently the parents belong 
to distinct species (Plate 34, and Histogram 2). 

Section Cerastes 

In the section Cerastes (those plants with typically opposite leaves) there 
appears to be more evidence for a greater number of subspecies and fewer 



I 



Vol. XXV] McMINN: FIELD HYBRIDS IN CEANOTHUS 331 

species and species complexes. In this section, C. ciineatus, C. Greggii, and 
C. prostratus have Avider ranges of distribution than any of the other entities 
and they have entered into more crosses than any of the others. Wherever 
C. cuneatus and C. Greggii and its vars. vestitus and perplexans overlap in 
their distribution, many intermediates occur which appear to be fertile. But 
since their degree of fertility is not known it is not possible to state whether 
the parents are in the nature of subspecies or species. In the crosses observed 
between C. cuneatus and C. prostratus the hybrid plants appear to be partially 
sterile, wherefore, their parents appear to be distinct species. At the present 
time no localities are known to the writer where C. prostratus and C. Greggii 
occur together. However, there is evidence in the southern middle Sierra 
Nevada of California that they may have overlapped in their distribution 
sometime in the past. In this area, particularly in Tuolumne and Calaveras 
counties, entities occur which seem to possess characters originating from 
both species. Two of these entities appear in some manuals as species binomials 
and are worthy of further consideration. 

The apparently fertile entity known as C. fresnensis (Fresno Mat), occurs 
on the western slope of the Sierra Nevada between 3000 and 6500 feet eleva- 
tion, from Fresno County northward to Tuolumne County or probably to 
Eldorado and Placer counties, mostly at the southern extension of the range 
of C. prostratus. In 1939 the writer gave the distribution of this rather baffling 
entity as extending from Fresno County northward to Plumas County. How- 
ever, after becoming more aware of the presence of hybrids in this genus, 
extensive field studies were carried on during the summers of 1940, 1941, and 
1942, and I am now of the opinion that some of the herbarium specimens 
studied from the more northerly localities had been taken from hybrid plants 
of C cuneatus x C. prostratus and its variety Jaxus, and from C. arcuatus. 
Ceanothus fresnensis is entirely prostrate, while C. arcuatus and most hybrids 
between C. cuneatus and C. prostratus are usually intermediate between the 
two in habit of growth. That C. Greggii var. vestitus has entered into the 
origin of C. fresnensis is evidenced by the character of the pubescence and 
the nature of the fruit. Ceanothus Greggii var. vestitus now occurs in Kern, 
Inyo, and Mono counties and in the past may have overlapped in its distribu- 
tion with C. prostratus. John Thomas Howell (1940) also considers the prob- 
ability that C. fresnensis "is descended from the offspring of a fertile cross 
between C. prostratus and C. vestitus" (C. Greggii Yar. vestitus). If we accept 
an early hybrid origin of C. fresnensis, then C. prostrat\is and G. Greggii var. 
vestitus must have belonged to distinct subspecies. 

The other apparent fertile entity previously referred to is C. arcuatus. It 
occurs in scattered localities from about 3000 to 7600 feet elevation on the 
western slope of the Sierra Nevada of California from Madera County north- 
ward to Plumas County. Near the summit on the south slope of Robb's Peak, 
Eldorado County, elevation 6500 to 6700 feet, it is one of the dominant shrubs 
associated with Arctostaphylos patula and Ceanothus prostratus. A few pros- 



332 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

trate plants Avliieli appear to be Ceanothus fresnensis also occur in this asso- 
ciation. These may be hybrids of C. prostratus and C arcuatus. 

Ceanothus arcuatus seems to have segregated out of a hybrid population 
formed by the crossing of C. cuneatus with C. fresnensis. The habit of growth, 
character of fruit, and color of flowers are intermediate between the two 
probable parents. The gi-ayish bark resembles that of C. cuneatus while the 
leaves resemble more those of C. fresnensis. In the Sugar Pine Ridge region 
along Highway 108, Tuolumne County, occur plants of C. cuneatus and 
C. fresne7isis as well as hj^brids between these two entities. At higher elevations 
along Highway 108 the hybrid form alone occurs. In Eldorado, Nevada, 
Placer, and Plumas counties, plants scarcely distinguishable from these hy- 
brids occur in the Upper Transition and Canadian Life Zones far above the 
distribution of C. cuneatus. Since these plants are scarcely distinguishable 
from the hybrids of Tuolumne County and since they occur in a different 
geographical area in the absence of both C. fresnensis and C. cuneatus, I 
believe there are sufficient reasons for regarding them as belonging to a dis- 
tinct subspecies or probably to a distinct species. It seems to me we have here 
a subspecies or a species which has "just arrived" and is now migrating north- 
Avard from its center of hybrid origin. 

The group of entities including C. pinetoruni, C. postratus and its vars. 
laxns and occidentulis, C. pmnilus, C. confusus, C. divergens, and C. purpii- 
reus has so many forms which appear to grade into each other that the entities 
may well exist as subspecies of one large polymorphic species. These entities 
are not only very closely related to each other but also to another smaller 
group composed of C. rigidus, C. Masonii, C. gloriosus and its var. exaltatus. 
The two groups differ slightly from one another in the character of the leaf 
margins. The former normally has leaves with serrate or spinulose margins, 
and the latter has leaves with dentate to quite entire margins. In the foothills 
of the Hood Mountain Range in Sonoma County, California, members of these 
two groups overlap in their distribution and have produced numerous hybrids, 
many of which appear to be fertile. Since the barrier between most of these 
entities appears to be that of geographic or ecologic isolation rather than 
genetic, it would seem that the entities are probably in the nature of subspe- 
cies. They seem to hybridize freely wherever they meet, but appear quite 
distinct in their own geographic ranges. 

Ceanothus sonomensis of the Hood Mountain Range, Sonoma County, Cali- 
fornia, may be the result of the crossing of one of these hybrids with C. cunea- 
tus, which occurs in the same area. This unit now occupies a small area 
somewhat isolated from the other forms o^ Ceanothus growing in that general 
region. When C. cuneatus is brought into the picture it brings in another 
series of forms which seems to hybridize with the others, and until experi- 
mental methods are used it would be unwise to make predictions as to the 
exact nature of the entities. That C. sonomensis, C. arcuatus, and several other 
somewhat similar entities in the section Cerastes might be the results of inter- 



Vol. XXV] McMINN: FIELD HYBRIDS IN CEANOTEUS 



333 



specific hybridization cannot be entirely ruled out. G. Ledyard Stebbins 
(1942a) lias pointed out in his paper on the "Role of Isolation in the Differen- 
tiation of Plant Species" that occasionally recombinations should occur in 
the progeny of a partially sterile interspecific hybrid involving "small units 
genetically independent of morphological differences." He further states that 
"Some of these new recombinations could be fertile types which would be 
different in appearance from either of their original parents and exhibit 
partial intersterility with both of them. They would thus become the progeni- 
tors of new species." 

CHAET 3 
Katviral Hybrlda 'betwcBn gpeclea of the Section Cerastes 
FlowerB white Flowers blue 

34. C. verrucosus -> — -> > — ^. ri(;idu3 ''.5, 

35. C. insularia ,C. Masonll 46. 
26. C. megacarpus • 
07. C. crassifollus^ 



/ p. gloriosus 47. 
/ -^ var. exaltftua 




38. C. Oreggii 

yar. vestltus 
yar. perplexano 

39. C. lanuginosus^ 

40. C. cuneatu 

41. C. Ferrisae — -^ 
yiowere tlua cr lavender ^ 

4S. C. arcuatu; 

43. C. fresnensls'^ 1 

■? -»(t. Vision 

44. C. T-«nni1nel1c y 



A summary of the recognized natural hybrids in the section Cerastes, as 
given in the book Ceanothus, gives a total of twenty-one, of which four are 
named and seventeen unnamed (Chart 3). The four named hybrids represent 
binomials which have appeared as distinct species in some publications. Eight 
so-called species of this section have entered into the hybrid complex. Ceano- 
thus cuneatus, C. prostratus, and C. Greggii are the most widely distributed 
named entities and they have entered into crosses more often than any of the 
other named entities. Ceanothus verrucosus, a species with relatively few 
variations, to my knowledge has not entered into any natural cross. This 
species has a rather limited range of distribution in San Diego County, Cali- 
fornia, which does not overlap with the distributional ranges of other species 
in the same section. According to the concept of genetic homogeneity as 
reviewed by Stebbins (1942), we might consider that C. verrucosus, because 
of its limited distribution, is genetically homogeneous and that it consists of 
relatively few biotypes which in turn are relatively homozygous. On the other 
hand we might consider C. cuneatus, C. prostratus, and C. Greggii, siuce they 



334 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. 

are widespread species, as being heterogeneous in their genetic make up. This 

heterogeneity is well exemplified by the numerous biotypes which occur in 

these species. 

Named Hybrids of the Section Cerastes 

X Ceanothus otayensis in known only from Otay Mountain in San Diego 
County, California, and is here considered to have originated as the result of 
a cross between C. crassifolius and C. Greggii var. perplexans. Both of these 
entities occur together on Otay Mountain. The hj^brid plants bear good fruit 
and the seeds appear to be fertile. If they are found to produce strong second 
generation plants, then the two parents, C. ci-assifolius and C. Greggii var. 
perplexans, must be considered to belong to different subspecies. One evidence 
for their fertility is the amount of variability which occurs among the hybrid 
plants. 

The name x Ceanothus flexilis has been given to those hybrid plants occur- 
ring in northern Shasta, Lassen, and southwestern Modoc counties of Cali- 
fornia. In these localities C. cuneatus and C. prostr-at^is var. laxus overlap in 
their distribution and they are considered to be the parents of x C. flexilis. 
The habit of growth, many leaf characters, size of fruit, and flower color are 
usually intermediate between the two parents. The fertility of the hybrids 
is not known, but from field observations I would suspect partial sterility. 
If the suspicion is proved to be a fact by experimental methods, then the 
parents, C. cuneatus and C. prostratus, belong to distinct species. 

X Ceanothus Bakeri is another named hybrid which appears to have re- 
sulted from a cross between C. prostratus var. laxus and either C. Greggii 
var. vest it us or one of the many varieties of C. cuneatus. The single plant 
known to me of this hybrid was collected by C. F. Baker from Kings Canyon, 
Ormsby County, Nevada, in 1902. The three fragmentary specimens resemble 
somewhat specimens of x C. flexilis and x C. connivens, both of which prob- 
ably have genes of C. prostratus in their makeup. This small bit of evidence 
would favor the assumption that C. prostratus and C. Greggii or C. cuneatus 
belong to distinct species. 

Specimens representing the named hybrid x C. connivens were collected 
from near Sheep Ranch, Calaveras County, California, by E. L. Greene and 
by T. S. Brandegee. Field studies in the Calaveras-Sheep Ranch area in 
1941 failed to disclose any specimens which quite resembled the specimens 
of the earlier collectors. However, in the Calaveras-Tuolumne area, where 
C. cuneatus, C. prostratus, and C. fresnensis grow, many h.ybrid jilants occur, 
some of which appear much like x C. connivens. It is probable that x C. con- 
nivens represents a cross between C. cuneatus and either C. prostratus or 
C. fresnensis. 

In concluding this paper mention should be made of the numerous other hy- 
brid plants which occur between C. cuneatus and C. jrrostratus wherever these 
two species occupy the same local area (Plates 35 and 86 and Histogram 3). 
Ceanothus cuneatus is primarily a component of hard chaparral in the Upper 



Vol. XXV] McMIXX: FIELD HYBRIDS IX CEAXOTHUS 



335 



Sonoran Life Zone, while C. prostratus occurs chiefly in the open pine forests 
of the Transition and Lower Canadian Life Zones. However, along the none- 
too-well-defined borders of these life zones these two entities overlap in their 
distribution and also in their characteristics. Some of the hybrids observed 
appear to have good fruits and seeds ; whether the seed produce strong and 



no. of 
plants 



Histogram 3 



40- 




' I  2  3  4 
C. prostratus Benth, 



6  7 8 

C. cuneatus Nutt. 



index 
value 



Hybrids 



Histogram 3 represents the distribution of plants into 9 groups based upon an index 
value determined by studying 8 characters of C. cuneatus. The plants were taken from a 
random sampling in an area where C. cuneatus and C. prostratus overlapped in their dis- 
tribution. Plants with 8 characters Avere classified as C. cuneatus; plants with none of these 
characters were classified as C. prostratus; and those with 1, 2, 3, 4, 5, 6, or 7 characters 
were classified as unnamed hybrids. Plants with a decreasing number of C. cuneatus charac- 
ters possessed an increasing number of C. prosti-atus ones. ^ 



fertile offspring is not known. Since the hybrid forms do not vary as much as 
F, and F3 offspring usually do from heterozygous parents, and since there 
are relatively few individuals compared to the number of individuals of both 
parents, I believe the hybrids are partially sterile and that their parents are 
distinct species. 



336 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th ser. i 

LITERATURE CITED 
Brandegee, K. 

1894, Studies in Ceanothus. Proceedings California Academy of Sciences (2), 4 : 173-222. 

Clausen, J. C, D. D. Keck, and W. M. Hiesey 

1939, The Concept of Species Based on Experiment. American Journal of Botany, 26 : 

] 03-106. 

HowEi.i.. J. T. 

1940, Studies in Ceanothus. Leaflets of "Western Botany, 2 :232. 
1940a, Studies in Ceanotlius. Leaflets of Western Botany, 2 : 259-262. 

McMiNN, H. E. 

1939, An Illustrated Manual of California Shrubs. 307-309. J. W. Stacey, Inc., San 
Francisco. 

Nobs, M. A. 

1941, Personal Communications. 

Stebbins, G. L. 

1942, The Genetic Approach to Problems of Eare and Endemic Species. Madrono, 6: 
241-258. 

1942a, The Eole of Isolation in the Differentiation of Plant Species. Biological Symposia, 
6:217-232. 

VAX Eensselaer, M. and H. E. McMinn 

1942, Ceanothus. Santa Barbara Botanic Garden, Santa Barbara. 



I wisli to express my sincere appreciation to Mr. M. A. Nobs and Mr. John Poindextei 
for f urjiishing the data and photographs for plates 29 to 36 and text figures 1 to 3. 



EXPLANATION OF PLATES 



PLATE 28 
Section Euceanotltus 

A. Branchlet with alternate leaves, thin, early falling stipules, and unihel-like clusters 
of flowers in racemes. 

B. Fruit without apical horns. 

C. Cross section of leaf showing stomata protected by guard cells on lower surface. 

Section Cerastes 

D. Branchlet with opposite leaves, thick, corky, persistent stipules, and umbel-like 
fruit -cluster. 

E. Fruit udth apical horns. 

F. Cross section of leaf showing stomata in sunken pits on the lower surface. 



[338 I 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV, NO. 14 [McMINN] PLATE 28 







[ 339 ] 



PLATE 29 

Section Euceanothus 
Fig. 1. C. arboreus. Fig. 10. C. Parri/i. 

Fig. 2. C. cordulatu.s. Fig. 11. C. Parryi. 

Fig. 3. C. cyaneus. Fig. 12. C. thijrsiflonis. 

Fig. 4. C. foliosus. Fig. 13. C. Ihymiflorus. 

Fig. 5. C incanu^. Fig. 14. C. tomentosus. 

Fig. 6. C integerrimus var. californicus. Fig. 15. C. velutiniis. 
Fig. 7. C. Lemmonii. Fig. 16. C. veluti)iu.s var. laevigatus. 

Fig. 8. C leucodermis. Fig. 17. C. spinosus. 

Fig. 9. C. papillosus. 



i 



[ 340 ] 



PROC CALIF, ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 14 



[MCMINN] PLATE 29 



^1 

Fig. 1 



Fig. ^ 






Fig. 12 



Fig. 2 







Fig. 7 



4^- 

Fi£^. 13 




Fig. 3 



Fig. 8 







Fig. 9 



Fig. 10 



Fig. 5 



Fig. 11 



Fig. 14 



1^ 

Fig. 15 






Fig. 16 



Fig, 17 



[ 341 ] 



PLATE 3U 

Section Cerasies 



Fig. 1. C. proNfratiis var. occidentalis. 

Fig. 2. C. cuneatus. 

Fig. 3. C. cnneatus. 

Fig. 4. C. cuneatus. 

Fig. 5. C. cuneatus. 

Fig. 6. C. Ferrisae. 

Fig. 7. C. gloriosus. 



Fig. 8. C prost rains var. laxus. 

Fig. 9. C. prostratus var. laxus. 

Fig. 1(1. C pro.strafus. 

Fig. 11. (7. ramulosus. 

Fig. 12. C. rigidus. 

Fig. 13. C. rigidus. 

Fig. 14. C. rerr«co.s«.s. 



342 



PROC. CALIF. ACAD. SCI . 4TH SERIES, VOL. XXV. NO. 14 



[MCMINN] PLATE 30 



4^ 



Fig. 2 



Fig. 6 






Fie. 11 



Fig. 7 



Fig. 12 



Fig. 3 



Fig. 8 



.0, 




Fig, 13 



Fig. 5 




Fig. 9 



^' 



Fig. 14 




Fig. 4 




Fig. 10 



[ 343 ] 



PLATE 31 



Diagrams of the growth forms of C. velutinus, C. cordulatus, 
and the hybrid x C. Lorensenii. 



[344] 



r 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV, NO. 14 



[McMINNJ PLATE 31 





X C.Lorenzenii 
( hybrid) 




£• c oraxxlatus 



[345] 



PLATE 32 



Photographs of specimens of C. velutinus, C. cordulatus, and a graded series of five 
of their hybrids taken from an area where the two species occur together. 



[ 346 ] 



J 



N 

<0 

UJ 
h 
< 

J 

a. 

i—i 

z 
z 

i 

u 

S 





z 
> 

X 
X 

J 

o 
> 

lli 
i 

I 
I- 



u 

d 
< 
u 
< 



_i 
< 
u 

u 


o: 







PLATE 33 

Chromosomes of C. cordidatus, C. velutinus, and of the hybrid x C. Loremenii. 
C. corduJatus, Nobs 76, Snow Flat, elevation 4500 feet, Nevada County, California ; 
12 regular pairs. C. veluti7ius, Nobs 79, Agate Bay, Lake Tahoc, Placer County, Cali- 
fornia; 12 regular pairs, x C. Lorenzenii, Nobs 117, Observation Point, Lake Tahoe, 
Eldorado County, California; 12 regular pairs, x C. Lorenzenii, Nobs 118, Observation 
Point, Lake Tahoe, Eldorado County, California ; 12 regular pairs, one very small 
pair, and 3 fragments. 



348 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 14 [McMINN] PLATE 33 



ui::ijfiut 



C, cordulatus 



No, 117 



uiaiiutii 



X C. Lorenzenii 



No. 118 



ittmiim-' 



C. velutinus 



««Ji:::i:i 



[349] 



PLATE 34 

Photographs of specimens of C. thyrsiflorus, C. papillosiis, and a graded series of five 
of their hybrids taken from an area where the two species occur togetlier. 



[ 350 ] 



Ui 

H 
< 

Q. 

I— 1 

Z 
Z 

i 

u 

2 



O 

z 
> 

X 
X 

J 

o 
> 

ui 

hi 

5 

LlI 
0) 

I 



U 

d 
< 

u 
< 



< 

u 

u 
o 

a. 




' / 



PLATE 35 

Photographs of specimens of C. prostratus, C. cuneatus, and a graded series of four of 
their hybrids taken from an area where the two species occur together. 



[352] 



10 

(n 

UJ 

H 
< 

_J 

0. 

I — 1 

z 
z 

s 

o 

5 



o 

z 

> 

X 
X 



o 
> 

en 

q: 

UJ 

tn 

I 
I- 



u 

d 
< 
u 
< 



< 
u 

u 
o 

q: 
a. 




PLATE 36 

Diagrams of the growth forms of C. prostratus, C. cuneatus, and 
one of their hybrids. 



[354] 



ID 

n 
u 

< 

J 

D. 

I — I 
Z 
Z 

u 

S 



o 

z 

> 

X 
X 

J 


> 

ui 

E 
111 

U) 

I 

H 



u 

d 
< 

u 
< 



_i 
< 
u 

d 



(T 
Q. 



-^^ 




O 

c 
u 






3 

a 

O 

a 

^1 



I 



« 



PROCEEDINGS ^^ 

OF THE V 

CALIFORNIA ACADEMY OF SCIENCES ^ 

Foil nil Series 
Vol. XXV, No. 15, pp. 357-376 December 27, 1944 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 15 
A REVISION OF PHACELIA SECTION MILTITZIA 

BY 

JOHN THOMAS HOWELL 

Assistant Curator, Department of Botany 
California Academy of Scie?ices 

INTRODUCTION 

THE SPECIES which are here presented as the section MUtitzia of the genus 
Phacelia have formerly been treated either as constituting the genus MU- 
titzia A.DC. or as forming the more numerous part of the genus Emmenanthe 
Benth. {Emmenanthe § MUtitzia A. Gray). I have recently expressed my 
belief that there is definitely a generic break between MUtitzia and Emmenan- 
the but that MUtitzia exhibits both morphologically and habitally a relation- 
ship so close to Phacelia sect. Microgenetes that it should be treated as a section 
in Phacelia (1944, pp. 12-16). The plump, corrugated seeds found in sect. 
MUtitzia are like those in sect. Microgenetes, while the compressed, reticulate 
seeds of Emmenanthe are different from any in Phacelia. Moreover the lateral 
attachment of the ovules to fleshy placentae in MUtitzia is characteristic of 
Phacelia, while in E^nmenanthe the pendent ovules are basally attached to the 
wing-like margins of membranous placentae. The marcescent persistent co- 
rolla, by which the species of sect. MUtitzia may be distinguished from those 
of sect. Microgenetes, was the character by which MUtitzia was separated 
from Phacelia although the character was not finally diagnostic since several 
species in Phacelia sect. Eutoca also have persistent corollas. Although it is 
this corolla character which MUtitzia and Emm.enanthe have in common, all 
other structural characters of MUtitzia are entirely Phacelia-like and indicate 
a relationship to Phacelia that must be interpreted as sectional rather than 
generic. 

The section MUtitzia consists of only nine species and these are confined to 
the desert areas of the western United States between the Rocky Mountains 

. [ 357 ] 



M.-si CALIFOIiXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

un the east and the Sierra Nevada and Cascade Range on the west, extending- 
northward to latitude 45° N. or thereabouts and southward to about 35° N. 
In this relatively limited area, the northern part of Nevada is not only the 
geographic center for the distribution of the species, but is also the center for 
species concentration. Here are found seven of the nine species, while in adja- 
cent southeastern Oregon are five and in eastern California four. To the 
eastward not more than two species are found in any one state. Three of the 
nine species are relatively common and widespread, P. adenophora, P. scopu- 
lina, and P. tetramera; two are more localized in distribution and less common, 
P. inundata and P. lutea; four are to be counted among- the rarest of western 
plants: P. glaherrima, knowai only from three collections; P. inyocnsis and 
P. suhmutica known from a few collections from restricted areas; and P. 
salina, known from five stations in three different states. 

Even to the present, there has been considerable confusion and not a little 
difficult}' in limiting species, part of this being due without doubt to the 
meagerness of collections available for study. A critical examination of numer- 
ous specimens assembled from various collections and institutions has revealed 
a series of mostly clear-cut and readily defined entities. Habit, foliage, and 
vesture have been found quite variable within species so that plants unlike in 
aspect are sometimes referable to the same species because of the critical but 
dependable characters in flowers and fruits. Because so little is known about 
the field occurrence of the subspeeific variants, only a single variety has been 
recognized taxonomically, but undoubtedly others will be recognized as the 
variants become better known. Phacelia adenophora, P. lutca, and P. scopulina 
are the only species presenting facies so polymorphic that they may be some- 
times confused with related species. The first, M'hich may be considered the 
most primitive species in the group because of its larger flower-parts and 
hairy corolla-tube and filaments, is sometimes difficult to distinguish from P. 
Jiitea aiid P. inundata. Phacelia scopidina, the most widely distributed species 
in the section, is also the most variable ; to the west there are intergrades wnth 
P. lutea, in the southeast P. suhmutica is very closely related, and P. salina 
seems to be merely an ecologically specialized variant with smaller parts. 

From a consideration of the environments in which these phacelias are 
found, it is very apparent that speeiation in the group has produced entities 
which occupy several of the most highly specialized situations in our Avestern 
deserts. Thus P. inundata is nearly confined to play as of evanescent ponds 
and shrunken lakes. Phacelia tetramera and P. salina are commonly restricted 
to highly saline flats. Although P. scopulina is generally rather common in 
sandy or gravelly soil of juniper slopes or sagebrush flats, it, too, sometimes 
occurs locally in more alkaline situations. Phacelia adenophora grows on sage- 
brush slopes, but in certain forms it is characteristic of clay flats of high 
plains. The closely related P. lutca seems to be characteristic of wash slopes 
of clay hills and valleys of eastern Oi-egon; and the highly localized P. inyo- 
ensis seems to be confined to boggy spots among granitic sands bordoi-ing the 



Vol. XXV] HOWELL: PEACELIA SECTION MILTITZIA 359 

western Sierra. In no other group of like size in the genus are there so many 
forms edaphically specialized and localized, rare creations to thrill the soul 
of the botanical traveler who might chance upon one flourishing amid the vast 
loneliness and seeming desolate sterility of a saline desert. 

ACKNOWLEDGMENTS 

This revision has been prepared in the Herbarium of the California Acad- 
emy of Sciences (CAS), and, although the collection there has furnished the 
basis for research, my study would have been far from complete if I had not 
been able to borrow abundant collections from other institutions. These insti- 
tutions are here listed, together with the symbol used in citing specimens : 

Chicago Natural History Museum (CM) 

Dudley Herbarium, Stanford University (DS) 

Gray Herbarium, Harvard University (G) 

Intermountain Herbarium, Utah State Agricultural College (lU) 

Missouri Botanical Garden (M) 

Greene Herbarium, University of Notre Dame (ND) 

New York Botanical Garden (NY) 

Pomona College (P) 

Eocky Mountain Herbarium, University of "Wyoming (EM) 

University of California (UC) 

United States National Herbarium (US) 

Willamette University (W) 

Washington State College (WS) 

For the privilege of borrowing specimens, I am very grateful; and again I 
wish to express my gratitude and appreciation to many friends who have 
furnished me with field data and specimens in this group over a period of 
years. Especially am I grateful to H. D. Kipley and R. C. Barneby for a com- 
plete set of their numerous excellent collections which they have obtained 
during the course of their critical field work in western American deserts 
from 1940 to 1944. The number of specimens cited from their collections in 
the present revision is indicative of the important contribution they have 
made to my study. 

Also it should be noted that because type specimens in the United States 
National Herbarium and in the Herbarium of the New York Botanical Garden 
are not available for study during wartime, some collections that are undoubt- 
edly in those institutions are not cited in this paper. 

TAXONOMY 

Phacelia sect. Miltitzia (A. DC.) J. T. Howell, Leafl. West. Bot. 4 : 15 (IdU) . Miltitzia 
A. DC. in DC. Prodr. 9: 296 (1845). Emmenanthe § Miltitzia (A. DC.) A. Gray, Proe. 
Amer. Acad. 10: 328 (1875). 

Flowers 5-merous or occasionally 4-merous; corolla mareescent, yellow or 
frequently tinged with lavender or violet ; corolla-scales present or lacking ; 
filaments glabrous except in one species; ovary generally more or less pubes- 



360 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

cent, the style pubescent at least at base except in one species, ovules 4-26, 
laterally attached to fleshy placentae ; capsule apiculate or muticous ; seeds 
transversely corrugated. — Annual herbs of saline or subsaline habitats in 
western North American deserts; herbage frequentlj^ fleshy, more or less 
pubescent and glandular, occasionally glabrous; stems branching and gen- 
erally widely spreading ; leaves entire to deeply pinnately divided. 

KEY TO THE SPECIES 

A. Corolla-tube hairy inside between stamens; filaments liairy 1. P. adenophora 

A. Corolla-tube glabrous inside; filaments glabrous. 

B. Seeds faintly but definitely transversely striate, the striations about 12; ovules 

20-26 3. P. inundata 

B. Seeds prominently transversely corrugated, the corrugations 6-12 ; ovules 4-20. 

C. Corolla 5-8 mm. long; corolla-scales generally conspicuous and united to base 

of filaments about 0.5 mm.; style and branches (1.75 or) 2-3.75 mm. 

long; hypogynous disk conspicuous Avith prominent rounded or deltoi<l 

lobes alternating with the stamens 2. P. lutea 

C. Corolla 1.3-4.5 mm. long; corolla-scales inconspicuous, obsolescent, or lacking, 
free from the filaments at base if present ; style and branches 0.25-2 mm. 
long. 
D. Plants glabrous or nearly so ; corolla subrotate in anthesis, the lobes often 
about as long as the tube; ovary and style glabrous or nearly so. 

4. P. glaberrima 

D. Plants more or less hirsutulous or villous and capitate-glandular or viscidu- 

lous; corolla eampanulate, the lobes shorter than the tube (or about 

equaling the tube in P. tetramera) ; ovary generally sparsely to 

densely hairy, the style hairy at least at base. 

E. Flowers mostly 5-mcrous; corolla 3-4.5 mm. long. 

F. Style and branches 1.25-2 mm. long; hypogynous disk incon- 
spicuous; ovules 10—15. 
G. Style hairy to above middle; capsule apiculate. 

5. P. scopulina 
G. Style glabrous except at very base; capsule nearly or quite 

without apiculation 6. P. submutica 

F. Style and branches 1 mm. long; hypogynous disk relatively con- 
sj)icuous. 
H. Ovules 18-20; seeds nearly 1 mm. long, the corrugations 

6 or 7 7. P. inyoensis 

H. Ovules 7-9; seeds 1.5-2 long, the corrugations about 9-11. 

8. P. salina 
E. Flowers commonly 4-merous; corolla 1.3-1.8 mm. long. 

9. P. tetramera 

1 . Phacelia adenophora J. T. Howell 

Leafl. West. Bot. 4:15 (1944) 

Emvienanthe gJandulifcra Torr. ex S. Wats., Bot. U.S. Geol. Explor. 40th Paral. (King's 

Exped.) 257 (1871). 
Miltitsia glandulifera (Torr.) Heller, Muhlenbergia 8:20 (1912). 
Miltitsia glanduUfera (Torr.) Heller var. calif ornim Brand, Univ. Calif. Publ. Bot. 4:224 

(1912). 
Not PlKicelia (jhnuhilifcm Piper, Contrib. U.S. Nat. Herb. 11 :472 (1906). 



Vol. XXV] HOWELL: PHACELIA SECTION MILTITZIA 361 

Stems few or several, spreading' from the top of the annual taproot, pros- 
trate or strongly ascending, slender or rather stout, mostly 1-3 dm. long, 
capitate-glandular and villous-puberulent or hirsutulous; basal leaves not 
very numerous and the upper leaves rather sparse, to 3 em. long and 1.7 cm. 
wide, oblongish to subovate, pinnately lobed or divided, the lobes oblongish, 
obtuse, entire or rarely dentate-lobed, the uppermost leaves sometimes entire, 
hirsutulous and little glandular, petioles hirsutulous and glandular, to 2.5 cm. 
long; racemes mostly exceeding the leafy part of the plant, to 1 dm. long, 
sessile or nearly so, flowers somewhat distant below, congested above or the 
flowers on spreading or recurved pedicels, pedicels stout or slender, 1-3 mm. 
long; flowers 5-merous; calyx-segments in flower 2—4.5 mm. long, 0.4—1 mm. 
wide, in fruit 5-6.5 mm. long, 0.5-1.25 mm. wide, linear to narrowly oblong- 
oblanceolate, sometimes a little spatulate, obtuse, hirsutulous and more or less 
glandular ; corolla marcescent and a little accrescent, tubular to campanulate, 
yellow or the lobes more or less tinged with lavender, 4r-l mm. long, lobes 1.25- 
2 mm. long, outside of corolla hairy, inside spareely to densely hairy in the 
tube between the stamens; corolla-scales 1-2 mm. long, linear to narrowly 
semilinear-lanceolate, attached at base to base of filaments about 0.5 mm.; 
stamens 3-5 mm. long, somewhat unequal, the filaments hairy below the mid- 
dle, anthers 0.5-0.75 mm. long (or rarely shorter) ; styles and branches 2-4 
mm. long, the style hairy to the middle, branches 0.5-0.75 mm. long ; ovary 1 
mm. long, ovules 8-14 ; hypogynous disk relatively prominent, the lobes be- 
tween the stamens broad and rounded ; capsule 4-6 mm. long, oblong to oblong- 
ovate, subacute to truncate below the terminal apiculation, longitudinally 
sulcate on backs of valves, sparsely puberulent, sometimes a little glandular 
above ; seeds oblongish, 1-1.5 mm. long, black-brown, prominently transversely 
corrugated, corrugations 8-12, rounded and mostly broad. 

Alkaline soil of high plains and sagebrush slopes, from middle western Ne- 
vada northward to northeastern California and southeastern Oregon. 

Eeprescntative collections. Nevada : Virginia Mts., 5000 ft., 8. Watson 
No. 885 (G, type collection of Emmenanthe gtandulifera) ; about 6 miles south 
of Gardnerville, Douglas Co., Cantetoiv in 1941 (CAS, CM, lU, US) ; Carson 
City, Ormsby Co., Anderson in 1865 (G, NY) ; Reno, Washoe Co., M. E. Jones 
No. 3986 (CAS, DS, G, lU, M, NY, P, RM, Vi^), Eastivood No. 14773 (CAS, 
CM, P) ; 5 miles east of Sparks, Washoe Co., Eastwood and Howell No. 30 
(CAS, US) ; Pyramid Lake, Washoe Co., Kennedy No. 1037 (CM, DS, G, M, 
NY, RM, UC, US) ; 12 miles north of Reno toward Pyramid Lake, Washoe 
Co., Ripley a7id Barne'by No. 4491 (CAS, G). In Graj'- Herbarium is a speci- 
men labeled "Winnemucca Valley" collected by Lemmon in 1875, but the cor- 
rectness of the locality is to be questioned. 

California: Near Viewland, Lassen Co., Ripley and Barnehy No. 5755 
(CAS); 12 miles north of Susanville, Lassen Co., J. T. Howell No. 11884 
(corolla-limb pale lavender fading to lavender-rose; CAS, US, W, WS) ; 
Horse Lake, Lassen Co., J. T. Howell No. 11892 (corolla bright yellow; CAS, 



362 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

CM, G, lU, M, ND), J. r. Howell No. 11898 (corolla-limb lavender; CAS, 
NY, P, RM) ; Madeline Plains, Lassen Co., Bnice No. 2135 (UC, type of 3///- 
titzia glandulifera var. calif oniica) , Peirson No. 6851 (CAS, P), J. T. Howell 
No. 11909 A (CAS, DS, W); canyon below Jess Valley, Modoc Co., Payne 
No. 549 (CAS). 

Oregon, Harney Co. : Catlow Valley, Henderson No. 8690 (CAS) ; Blitzen, 
Train No. 90 (US) ; base of Steens Mts., Train (Univ. Colo.). These collec- 
tions from Oregon are not typical and show characters that vary toward 
P. lutea. 

Phacelia adenophora is a rather variable entity, but it is usually well marked 
by the conspicuous pubescence on the filaments and in the corolla-tube. In 
typical form as it occurs in middle western Nevada, it is a slender and some- 
what elegantly branched plant with the upper parts rather finely hirsutulous 
and copiously capitate-glandular. In the vicinity of Reno, Nevada, and to the 
northward, this typical variant passes by numerous intergrades into a coarser, 
more densely pubescent plant with flowers a little larger, the form which 
Brand called Miltitzia glandulifera var. calif ornica. From southeastern Ore- 
gon relatively few collections of P. adenophora are known, but, because the 
plants that grow there frequently exhibit characters intermediate to P. lutea 
which also grows in the same region, it is suspected that the intermediates 
may be of hybrid origin. 

Although P. adenophora is closely related to P. lutea and perhaps should 
be considered only a variety of that species, I regard the differences between 
the two specific in value. It is true that Asa Gray remarked that P. adenophora 
{Emmenanthe glandidifera) was ''probably a mere form" of P. lutea (1878. 
p. 171) , but an examination of specimens in Gray Herbarium discloses that at 
that time he had no specimen of P. lutea, and that he mistook for that species 
the more robust form of P. adenophora which Brand has called Miltitzia 
glandulifera var. calif ornica. Quite recently Jepson has treated P. adenophora 
and P. lutea as the same species (1943, p. 281, as Miltitzia lutea), but each is 
characterized by a distinctive appearance and each by its own peculiar set of 
variations which do not approach each other except in instances of suspected 
hybridization. I do not believe that there should be a change in the status 
of P. adenophora until its relation to P. lutea is clearly understood through 
more detailed study and critical field work. 

Within P. adenophora no variant seems sufficiently distinguished to be 
recognized taxonomically. Although Brand's var. calif ornica of northeastern 
California has considerable right to varietal recognition, I have concluded 
that it is not clearly enough defined either geographically or morphologically 
to be accepted at this time. Also I have seen several collections of a peculiar 
glandular-villous form with large floAvers and slender fruits from southern 
Oregon which should perhaps be accorded taxonomic recognition, but. be- 
cause of a possible connection which this plant may have with the imperfectly 
understood relations between P. adenophora and P. lutea in tliat region. 



VoL.XXV] HOWELL: PHACELIA SECTION MILTITZIA 363 

nominal recognition of this entit}^ too, should await more numerous collections 
and further field work. 

A collection intermediate between P. adenophora and P. lutea is Coville 
and Leiherg No. 68 (US), from the road to Coleman Valley, Harney or Mal- 
heur Co., Oregon. In it the corolla-tube is pubescent only at the base of the 
stamens, or glabrous, and the filaments vary from subglabrous to distinctly 
hairy. Another intermediate collection from Oregon looking quite like P. 
adenophora, but with the technical floral characters of P. lutea, is Willits 
No. 169 (DS) from Hart Lake, Lake Co. A collection of P. adenophora which 
resembles P. lutea, but which has heavily hairy corolla-tube and filaments, 
has been collected near Canby, Modoc Co., California, /. T. Howell No. 

12215 (CAS). 

2. Phacelia lutea (H. and A.) J. T. Howell 

Leafl.West. Bot. 4:15 (1944) 

Stems one to several from the top of an annual taproot, spreading or 
ascending to substrictly erect, 2-8 cm. long, hirsutulous and scarcely gland- 
ular to hirsutulous and capitate-glandular or almost entirely glandular- 
villous ; leaves entire or coarsely serrate, rarely serrate-lobed, oblanceolate 
and oblong to elliptic and ovate, mostly 1-2.5 cm. long, 0.3-1 cm. wide, 
hirsutulous and a little glandular to sparsely glandulai'-villous, obtuse at 
apex, cuneate at base, petiole to 2.5 cm. long, usually equaling or shorter 
than the blade ; racemes to 5 cm. long, sometimes more or less exceeded by 
and concealed by the leaves but generally exceeding the leaves and showy, 
pedicels 1-2 mm. long or the loAvest to 5 mm. long, the lowest sometimes 
reflexed in fruit ; calyx-segments in flower 3.5-5 mm. long, in fruit 6.5-9 
mm. long, 1-1.75 mm. wide, linear-oblong to narrowly oblong-oblanceolate, 
subacute, hirsutulous and more or less glandular to rather densely capitate- 
glandular ; corolla yellow, sometimes becoming lavender-tinged in age, 5-8 
mm. long, campanulate or tubular, the lobes erect or spreading in anthesis, 
1-2 mm. long, denticulate, corolla sparsely hairy outside, glabrous inside ; 
corolla-scales 2-3 mm. long or sometimes obsolescent, semilinear-lanceolate, 
attached at base about 0.5 mm. to the filaments ; stamens 3-5.5 mm. long, 
filaments glabrous, anthers 0.4-0.75 mm. long; style and branches 1.75-3.75 
mm. long, branches 0.25-0.75 mm. long, style hairy only near the base or 
to above the middle; ovary 1.25-1.75 mm. long, ovules 10-17; hypogynous 
disk prominent with conspicuous rounded or deltoid lobes between the 
stamens ; capsule 5.5-7 mm. long, oblong to elliptic, subobtuse, apiculate, 
the beak to 1 mm. long, sparsely hairy and a little glandular ; seeds 1-1.5 
mm. long, elliptic-oblong, subobtuse to subacute, brown, prominently cor- 
rugated, the corrugations about 9 or 10, conspicuously reticulate. 

Clayey and sandy slopes and flats in southeastern Oregon, occurring 
locally in western Idaho and northwestern Nevada. 

The type of P. lutea was part of a collection of plants "gathered in the sum- 
mer of 1837 by a friend of Mr. Tolmie" at "Snake Fort, Snake Country." In 



364 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

the introduction to the Supplement of Botany of Captain Beeehey's Voyage 
(p. 315), it is stated that "Snake Fort is built at the junction of Reed's River 
with the Snake [River], the position of which is in N. lat. 44° 20', long. 116° 
W." On modern maps the name Reed's River does not appear and the lati- 
tude and longitude indicate a point considerably east of the Snake RiA'er. 
On the map in Hooker's Flora Boreali-Americana, vol. 1, however, Reed's 
River is given and from its position it is clearly the Boise River of the present 
day, the chief affluent of the Snake River from the east between the Payette 
River and the great bend of the Snake. Hence, collections that have been 
available from northwestern Owyhee Countj^ Idaho, may be regarded as topo- 
types of this interesting species that was first collected more than a hundred 
years ago. 

As has been pointed out in the discussion of P. adenopliora , Asa Gray had 
no specimens of P. lutea at the time he prepared his account of Emmenmifhe 
§ Miltitzia for the Synoptical Flora (1878) . As is indicated by his annotations 
on specimens in Gray Herbarium, he did receive collections of P. lutea from 
Cusick by the time the second edition was published (1886), but there is 
nothing in the supplement of the second edition concerning these specimens. 
Cusick's plants, collected from three different stations under varying environ- 
ments, were probably more confusing than helpful to Gray. One part of the 
collection. No. 1233a, from "sandy hillsides of Malheur and vicinity" is typical 
P. lutea, but No. 12331), "same locality but on heavy clay soil," and No. 1233c, 
"barren valley in rather heavy soil," are quite different in appearance and 
were originally noted as probably Emmenanthe parviflora (P. inundata) by 
Gray, and were more recently identified as Miltitzia gland^difcra (i.e., P. ade- 
nophora) by Macbride. Although these puzzling plants and others like them 
from Professor Peck's collections are definitely referable to P. lutea, they 
vary toward P. scopidina and present a problem of relationship that can only 
be indicated at this time. These variants are discussed further under P. .scopu- 
lina, and under P. adenopliora the problem of intermediates between that 
species and P. lutea is discussed. 

One variant in P. lutea, howeATr, seems quite distinct and well deserving 
taxonomic recognition, a small-flowered plant with glandular-villous pubes- 
cence. This variety may be distinguished as follows : 

Key to Varieties of Phacelia liutea 

Corolla 6-8 mm, long; style hairy to the middle or above var. typica 

Corolla 5-6 mm. long; style hairy only near the base var. purpurascens 

2a. Phacelia lutea (H. and A.) J. T. Howell 

var. typica, nom. nov. 

P. lutea (H. and A.) J. T. Howell, 1. c. 

Euioca lutea H. and A. Bot. Beech. Voy. 373 (1840) ; Hook. Icon. tab. 354 (1841). 

Miltitzia lutea (H. and A.) A. DC. in DC. Prodr. 9:296 (1845). 

Emmertanthe lutea (11. and A.) A. Gray, Proc. Amer. Acad. 10:328 (1875). 



Vol. XXV] HOWELL: PEACELI A SECTION MILT ITZI A 365 

Plants hirsntulous and scarcely glandular to moderately glandular- 
liirsutulous, generally yellowish-green ; racemes mostly exceeding the leaves ; 
corolla 6-8 mm. long, showy ; style sparsely hairy to the middle or above, 
2.5-3.75 mm. long. 

Representative collections. Idaho: Squaw Creek near Marsing, Owyhee 
Co., Tucker No. 1029 (CAS ex Univ. Ida. So. Br.). 

Oregon: Harper, Malheur Co., Peck No. 20553 (CAS, W), Henderson No. 
8689 (CAS) ; 5 miles south of Harper, Malheur Co., Peck No. 16104 (W, 
approaching var, purpurascens) ; Roekville, Malheur Co., Henderson No. 
8691 (CAS, depauperate) ; Sucker Creek, 30 miles north of Jordan Valley, 
Malheur Co., Holmgren No. 2206 (CAS, lU) ; Malheur Butte, Malheur Co., 
Leiierg No. 2043 (CM, G, NY, UC, US) ; 15 miles southeast of Blitzen, Harney 
Co., Applegate No. 5591 (DS, W) ; near Burns, Harney Co., Lilla Luch in 
1927 (W) ; base of SteensMts., Harney Co., HoweZZmiSSo (CM, G, NY, US) ; 
Unity, Baker Co., Peck- No. 16186 (W) ; Paulina Creek northeast of Lapine, 
Deschutes Co., WUted No. 1679 (ND). 

Nevada: Virgin Valley, Humboldt Co., L. Kellogg in 1909 (P). 

An unsual variant, entirely glabrous except for an occasional hair in the 
inflorescence and for a few scattered hairs and glands at the top of the ovary 
and on the lower part of the style, has been collected by Ripley and Barneby 
in Owyhee County, Idaho, where it grew on a "friable alkaline clay bank 24 
miles southwest of Marsing" (No. 6153, CAS, G, US). A few of the corollas 
were noted by the collectors as being tetramerous. 

2b. Phacelia lutea (H. and A.) J. T. Howell 
var. purpurascens J. T. Howell, var. nov. 

Glanduloso-villosa vix hirsutula, caulibus foliis rhachidibus et segmentis 
calycis valde purpurascenti-tinctis; foliis racemos plerunique superantibus ; 
corolla 5-6 mm. longa; stylo 1.75-2.5 mm. longo, pilifero solum prope basin. 

Type: Herb. Calif. Acad. Sci. No. 127399, collected in "chocolate soil" on 
high hills opposite Humphrey's, Grant County, Oregon, by L. F. Henderson, 
No. 5092, April 30, 1925. There is a duplicate in Gray Herbarium. Only one 
other collection entirely typical of var. purpurascens has been seen : Cusick 
No. 2634, which was obtained "in red clay soil of Crook Co." (CM, G, M, ND, 
NY, P, RM,UC,US). 

3. Phacelia inundata J. T. Howell 

Leafl. West. Bot. 4 : 15 (1944) 

Emmenanthe parviflora A. Gray, U.S. Pacif. ER. Reports 6:84, tab. 15 (1857). 
Miltitzia parviflora (A. Gray) Brand, Das Pflanzenr. IV. 251:131 (1913). 
Not Phacelia parviflora Pursli, Fl. Amer. Sept. 1 :140 (1814) ; nor P. parviflora Phil., Anal. 
Univ. Chile 90:226 (1895). 

Stems several, spreading or ascending from the top of the annual taproot, 
mostly 1-4 dm. long, but the plants sometimes depauperate and the stems only 
0.1-0.3 dm. long, finely glandular and somewhat hirsutulous; leaves rosu- 



366 CALIFOSNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

late and also rather numerous up to the ends of the branches, mostly 1-3 cm. 
long and 0.4r-1.5 em. wide, oblong to oblong-obovate, subentire to pinnately 
parted and divided, the divisions obtuse, mostly entire, oblong or deltoid- 
oblong, the terminal division largest and generally lobed, the basal divisions 
frequently small and discrete, petioles 0.5-2 cm. long, finely glandular- 
hirsutulous, the blades finely hirsutulous and a little glandular beneath ; 
racemes mostly exceeded by the leaves, to 12 cm. long, the lowest flowers rather 
distant, the uppermost somewhat crowded, pedicels 1-2 mm. long or the lowest 
to 4 mm. long, erect or somewhat spreading in fruit ; flowers 5-merous ; calyx- 
segments in flower narrowly oblongish, 4 mm. long, 0.6-1 mm. wide, in fruit 
6-10 mm. long, 1-1.5 mm. wide, linear-oblanceolate to linear-oblong, some- 
times a little spatulate, obtuse, hirsutulous and a little glandular; corolla 
yellow, 4—5 mm. long, the lobes rotately spreading in anthesis, becoming erect 
later, lobes about 2 mm. long, pilose-hairy outside, glabrous within; corolla- 
scales narrowly semilinear -lanceolate, 1.25 mm. long, at base free from fila- 
ments, or the scales obsolete ; stamens quite unequal, 2-3 mm. long, filaments 
glabrous, anthers roundish, 0.3 mm. long; style and branches 1-1.25 mm. long, 
style hirsutulous nearly to the branches, branches 0.25 mm. long; ovary 1.5 
mm. long, densely hirsutulous, ovules 20-26 ; hypogynous disk rather promi- 
nent with rounded lobes between the stamens ; capsule 5.5-7 mm. long, oblong, 
abruptly narrowed above into a prominent deltoid apiculation and persistent 
style, sparsely hairy below, sparsely hirsutulous above and on the beak, not 
glandular; seeds 1.5-1.75 mm. long, subdeltoid, irregularly angled, mostly 
acute at the ends, tending to be strongly flattened, reticulate-pitted, finely but 
definitely transversely striate, the striations about 12-14. 

Summer-dried playas of desert lakes and sinks from middle southern and 
southeastern Oregon south to northeastern California and middle western 
Nevada. 

Collections. Oregon: Klamath Lake, Klamath Co., Xewhcrry (G, type); 
Silver Lake, Harney Co., Cusick No. 2727 (CM, G, M, ND, NY, P, RM, UC, 
US; plant noted as having a "very disagreeable odor") ; Steens Mts. above 
Alberson, Harney Co., Teck No. 14217 (NY, W). 

California : east side of Clear Lake, Modoc Co., J. T. Howell No. 12357 
(CAS, US) ; between Malin and Canby, Modoc Co., M. S. Baker No. 8307 
(CAS) ; west side of Eagle Lake at Spaldings, Lassen Co., /. T. Howell No. 
12510 (CAS, CM, DS, P, W). 

Nevada : Swan Lake, Washoe Co., Train No. 3006 (lU, NY) ; Washoe Lake, 
Washoe Co., M. E. Jones in 1897 (CAS, P) ; Franktown, Washoe Co., Ripley 
and Barnehy No. 5947 (CAS) . 

Phacelia inundata is one of the more rarelj'- collected species in the section 
Miltitzia. Although three of the nine collections that have been examined have 
come from northea-stern California, before this it has been reported only 
from southeastern Oregon and Nevada (Tidestrom, 1925, p. 448; Peck, 1941, 
p. 593). Only a single specimen has been seen (Peck No. 14217) in which the 



VoL.XXV] HOWELL: PHACELIA SECTION MILTITZIA 367 

collection data indicate an environment away from a desert lake or sink, while 
the Californian collections which I made came from lake-bottom areas that 
had been covered with water in the earlier part of the season. Because of its 
preference for so peculiar a habitat, P. inimdata appears to be one of the most 
highly specialized and advanced species in the genus from a physiological 
point of view. Morphologically it is quite distinct, too, its nearest relative 
perhaps being P. lutea. 

4. Phacelia glaberrima (Torr.) J. T. Howell 

Leafl. West. Bot. 4:15 (1944) 

Emmenanthe glaberrima Torr. ex S. Wats., Bot. U.S. Geol. Explor. 40th Paral. (King's 

Exped.) 257 (1871). 
Miliiisia glaberrima (Torr.) Brand, Das Pfianzenr. IV. 251:131 (1913). 

Stems several, spreading from top of annual taproot, 3-20 cm. long and 
probably longer, first internodes above the base very long, entirely glabrous ; 
leaves 1-1.5 cm. long, 0.4-1 cm. wide, glabrous or with a few hirsutulous hairs 
on the margins and near the apex, broadly oblanceolate and ovate to broadly 
elliptic, entire or with 1 to few coarse serrations, obtuse at apex, narrowly 
cuneate at base and frequently a little decurrent along the petiole, petiole to 
2 cm. long below, very short above ; racemes to 8 cm. long, flowers approximate 
or rather distantly spaced, pedicels mostly 1-3 mm. long ; flowers 5-merous ; 
calyx-segments in flower very unequal, 2.5 mm. long, 0.25-1 mm. wide, in 
fruit 4—5 mm. long, 0.3-2 mm. wide, oblanceolate to obovate, fleshy and with 
a prominent midrib near the base, glabrous or with a few short scattered 
hirsutulous hairs near the apex ; corolla 3-4 mm. long, subrotate in anthesis, 
the lobes 1-3 mm. long, becoming erect later, entirely glabrous within and 
without; corolla-scales obsolete; stamens 2.5-3 mm. long, filaments glabrous, 
flattened at base, anthers 0.4 mm. long ; style and branches 1-2 mm. long, the 
style glabrous, deltoid-subulate, compressed, the branches 0.5 mm. long, 
ascending ; ovary glabrous or sometimes with a few scattered hairs near the 
top, 1 mm. long, ovules 4-10 ; hypogynous disk prominent about uniformly 
wide all around and not conspicuously lobed; capsule 4-5 mm. long, ovate to 
elliptic, glabrous or nearly so, somewhat rounded above, prominently apicu- 
late, valves coriaceous with conspicuous thickened margins, longitudinally 
sulcate; seeds 1.5-2 mm. long, oblong or ovatish, prominently corrugated, 
corrugations rounded, reticulate, about 10 or 11. 

Phacelia glaberrima is a remarkable and localized endemic in the saline 
desert of north-central Nevada and, until it was rediscovered in the Keese 
River Valley in June, 1944, by H. D. Ripley and R. C. Barneby, it was known 
only from the collections of Sereno Watson made in 1868 on King's Expedi- 
tion. The three presently known collections are: Humboldt Sink, 4300 ft., 
Watson in Maij, 1868 (G, UC frag.) ; Reese Valley, 5000 ft., Watson in July, 
1868 (G, UC frag.) ; Reese River Valley 32 miles north of Austin, Lander 
Co., Eipley and Barnehy No. 6186 (CAS, G, US). Concerning the highly 



368 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

specialized liabitat of P. glaherrima, Mr. Barneby writes as follows : "The 
Reese River Valley is not verj^ exciting for the most part, one of those broad 
sterile valley's covered with sagebrush and with long fans of gravel leading 
into the hills, but at one point about 60 miles south of Battle Mountain the 
mountains close in and the river has to negotiate a short canyon. Here the 
mountains are mostly volcanic, but there are localized occurrences of lime- 
stone, and these have been broken down by weathering into white bluffs of a 
stiff gravel and clay mixture impregnated with alkali. On the sides of these 
bluffs, on which nothing else can exist, Phacelia glaherrima is abundant, its 
yellowish herbage very succulent and fragile and its abundant racemes of 
golden-yellow visible from afar. Occasionally it seeds down into the washes 
and there was one spot where it was associated with P. scopulma and P. 
gymnoclada, the three species, so similar and yet so different, sometimes 
enlaced into small "trispecific" mats. The corolla of P. glaherrima is very 
open-campanulate, almost rotate when fully expanded, and the leaves are 
occasionally crenate." 

Although P. glaherrima might be related to P. scopulina because of its 
smaller corollas which are devoid of corolla-scales, this rare Nevada plant is 
probably much nearer P. lufea and, in its subrotate corollas, it is reminiscent 
of the equally highly differentiated P. imindata, a species which I have also 
related to P. lutea. Among the numerous and varied Miltitzia collections of 
Ripley and Barneby are remarkable glabrous forms of both P. lutea and P. 
scopulina but from these P. glaherrima may be readily distinguished by the 
rotately spreading and deeply lobed corolla, the glabrous style, and the thick- 
rimmed valves of the capsule. From Watson's collections I concluded that the 
ovary was glabrous as originally described but in the ample material of 
Ripley and Barneby it is apparent that the ovary is sometimes sparsely and 
minutely hairy near the top. 

Phacelia glaherrima has been reported from the Flax River (Little Colorado 
River) in northern Arizona, where it was supposed to have been collected by 
Newberry, but I have located no Arizona collection (cf. Kearney and Peebles, 

1942, p. 737). J 

5. Phacelia scopulina (A. Nels.) J. T. Howell | 

Leafl. West. Bot. 4:16 (1944) 

Emmenanthe scopulina A. Nels., Bull. Torr. Bot. Club 25 : 380 (1898). 

Miltitzia lutea (H. and A.) A. DC. var. scopulina (A. Nels.) Brand, Das Pflanzenr. IT. 

251:131 (1913). 
Miltitzia scopidina (A. Nels.) Eydb., Bull. Torr. Bot. Club 40 : 479 (1913). 
Emmenanthe dispar M, E. Jones ex Eydberg, Fl. Eky. Mts. and Adj. Plains 708 (1917), as 

synonym of Miltitzia salina. 

Stems several from the top of an annual taproot, 2-10 cm. long, spreading 
or ascending and tending to be erect, sparsely to rather densely hirsutulous, » 
more or less cinereous, eglandulose or minutely viscidulous above; leaves 
oblanceolate to oblong or oblong-obovate, 1-3 cm. long, 0.5-1.5 cm. wide, 
entire, serrate- or crenate-lobed, or rather deeply pinnately lobed, the lobes 



VoL.XXV] BOWELL: FRACELIA SECTION MILTITZIA 3(59 

oblong, deltoid, or elliptic, the blade obtuse at apex, cuneate to rounded at 
base, hii-sutulous, petioles to 4 cm. long, frequently longer than the blades ; 
racemes rather few-flowered and more or less concealed by the leaves, to 
4.5 cm. long, pedicels 1-6 mm. long, erect or sigmoid-spreading, or the lowest 
recurved in fruit ; floAvers 5-merous ; calyx-segments 2.5-4.5 mm. long, about 
0.5 mm. wide, linear-oblong, in fruit becoming 6-7 mm. long, 0.5-1 mm. wide, 
linear-oblong or linear-oblanceolate, obtuse, usually Avithout a prominent 
raised midvein except at the base, hirsutulous, rarely capitate-glandular 
and perhaps sometimes a little viseidulous; corolla tubular-campannlate, 
somewhat accrescent in age, 3-5 mm. long, the lobes 0.75-1 mm. long, yellow 
or strongly tinged with lavender or violet, the outside hairy, the inside 
glabrous; corolla-scales inconspicuous or obsolete, filiform-linear, about 
1 mm. long, free from the base of the filaments ; stamens 2.5-3 mm. long, the 
longest frequently equaling the lobes of the corolla, filaments glabrous, 
anthers 0.3-0.4 mm. long; style and branches 1.3-2 mm. long, the branches 
0.2-0.3 mm. long, the style hirsutulous-hairy to the middle or above ; ovary 
1-1.25 mm. long, ovules 11-15 ; hypogynous disk inconspicuous and scarcely 
lobed between the stamens ; capsule 5-6 mm. long, oblong, obtuse or sub- 
acute at top below the prominent apiculation, hirsutulous-puberulent espe- 
cially above; seeds oblong or oblong-ovate, 1.25-1.75 mm. long, brown, 
prominently corrugated, corrugations rounded, 9-11, reticulate. 

Loose sandy or gravelly soil of desert flats and slopes, from southwestern 
Montana and Wyoming southward to western Utah and northern Nevada, 
and westward to eastern Oregon. 

Representative collections. Montana : Melrose, Silver Bow Co., Rydherg 
No. 2771 (NY, US). 

Wyoming : Green River, Sweetwater Co., A. Nelson No. 3056 (RM, type of 
EmmenantJie scopulina; G, M, US), A. Nelson No. 4712 (CAS, G, M, NY, P, 
RM, US, WS), M. E. Jones in 1896 (CAS, lU, M, NY, P, RM, US) ; between 
Eden and Big Piney, Sublette Co., Pay son and Pay son No. 2576 (CM, G, M, 
NY, P, RM,UC,US). 

Utah : Dutch Mt., Tooele Co., M. E. Jones in 1891 (P). 

Nevada: Wells, Elko Co., 31. E. Jones in 1930 (P), Eastwood & Howell 
No. 330 (CAS, CM, DS, ND, US), Ripley and Barnehy No. 5524 (CAS) ; 25 
miles north of Wells, Elko Co., Holmgren No. 955 (CAS, lU, UC) ; foothills 
of Independence Mts. southeast of Tuscarora, Elko Co., Ripley and Barnehy 
No. 5551 (CAS) ; near Wendover, Elko Co., M. E. Jones in 1929 (CAS, P) ; 
Cobre, Elko Co., M. E. Jones in 1906 (CAS, P), in 1907 (P) ; foothills of 
Schell Creek Range west of Connors Pass, White Pine Co., Ripley and 
Barnehy No. 6305 (CAS) ; Monitor Valley south of Lone Mt., Eureka Co., 
Ripley and Barnehy No. 6203 (CAS) ; 11 miles west of Eureka, Eureka Co., 
Ripley and Barnehy No. 6235 (CAS) ; Reese River Valley 32 miles north of 
Austin, Lander Co., Ripley and Barnehy No. 6187 (CAS). 

Oregon : Harney Lake, Harney Co., Henderson No. 8688 (CAS) ; 24 miles 



370 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

north of Wagontire, Harney Co., Peck No. 19394 (W, simulating atypical 
P. lutea) ; 8 miles west of Lakeview, Lake Co., Peck No. 15243 (W, more glan- 
dular than typical) ; south of Narrows, Harney Co., Bipley and Barneby No. 
6088 (depauperate, CAS). 

Although Brand considered Emmenanthe scopulina a variety of Miltitzia 
lutea, I can see no reason for following so conservative a procedure. Phacelia 
scopulina has a very distinctive appearance and is marked by excellent char- 
acters over a wide range; and, in a much more restricted range, Phacelia lutea 
is usually quite as distinct. In eastern Oregon, however, where the ranges of 
the two species overlap, plants have been collected which are confusingly 
intermediate in appearance, although from the critical characters of the 
flower-parts, such specimens have been definitely referable to one species or 
the other. A pair of such specimens has been collected by Prof. M. E. Peck, 
P. lutea coming from 7 miles east of Buchanan, Harney County {No. 20536, 
W), P. scopulina from 25 miles north of McDermitts, Malheur County {No. 
20600, W) . A series of collections of P. lutea accompanied by field notes made 
by Cusick in eastern Oregon and preserved in the Gray Herbarium would 
seem to indicate that some of the variations noted may be due to environmental 
conditions, but it seems likely that much of the variability results from the 
genetic makeup of the group and may be due to hybridization between P. lutea 
and P. scopulina. Cusick's collections are more fully described under P. lutea. 

Another collection of P. scopulina that exhibits noteworthy variability was 
made by Ripley and Barneby in Humboldt County, Nevada {No. 4557, CAS). 
In this collection there are two kinds of plants, one that is normally hirsutu- 
lous-pubescent throughout, the other that is quite glabrous except for a few 
scattered hirsutulous hairs on some of the leaves and for a few minute capitate- 
glands on the stems, calyx-segments, and upper part of the ovary. Because 
the only known collections of P. glaherrima were made in the northwestern 
part of Nevada not very far from the station where Ripley and Barneby found 
this odd variant, their plants were compared with specimens of P. glaherrima. 
The floral morphology of the two was found to be so unlike that it would be 
quite wrong even to suggest that the subglabrous individuals are intermediate 
between the two species. 

M. E. Jones distributed his 1896 collection of P. scopulina made at Green 
River with the name Emmenanthe dispar, hut apparently he never described 
the plant. Although Rydberg published the name as a synonym of Miltitzia 
salina, as indicated above in the synonymy, he had correctly determined and 
annotated Jones' specimen in Herb. N.Y. Bot. Gard. as Miltitzia scopulina. 

6. Phacelia submutica J. T. Howell, spec. nov. 

Herba annua plus minusve glanduloso-hirsutula; caulibus paucis vel mul- 
tis ex ba.si, patentibus vel suberectis, 2-8 cm. longis, purpurascenti-tinctis, 
sparse vel subdeiise vestitis pilis et hirsutulis albis gracilibus et paucis capitato- 
glandulosis; foliis plerumque dispersis, 0.5-1.5 cm. longis, 0.2-0.8 cm. latis,. 



VoL.XXV] HOWELL: PHACELIA SECTION MILTITZIA 371 

elliptico-oblongis ovatis obovatisve, integris vel tenuiter creiiulatis vel promi- 
nenter serrato-lobatis, apice obtusis, cuneatis basi, plus minusve hirsutulis, 
petiolo ad 2 cm. longo, saepe quam lamina longiore; racemis circa 2 cm. longis, 
saepe f oliis brevioribus, floribiis subcongestis vel infimis distantibus, pedicellis 
plerumque erectis vel adscendentibus, ad 6 mm. longis ; floribus 5-meris ; seg- 
mentis calycis ad anthesin 3.5-5 mm. longis, 0.3--0.5 mm, latis, in fructu valde 
accrescentibus, 6-10 mm. longis, 0.3-1 mm. latis, hirsutulis et capitato-glandu- 
losis, basi calcarif ormi-angulata, costa prominenti ; corolla tubulata, 3.5-4.5 
mm. longa, dilute lutea, baud purpureo-tincta, pubescenti extus, intus glabra, 
lobis 1 mm. longis, crenulatis ; squamis ad lineas reductis vel nullis ; sta- 
minibus inclusis, 2.2-2.75 mm. longis, filamentis glabris, antlieris 0.3 mm. 
longis, latioribus quam longioribus explanatis; stylo et ramis 1.25-1.5 mm. 
longis, stylo paulum pilifero tantum prope basin, ramis 0.2-0.25 mm. longis; 
ovario 0.75-1 mm. longo, hirsutulo supra, ovulis 10 vel 11 ; eapsula 3-5 mm. 
longa, elliptica vel elliptico-ovata, apice rotundata, mutica vel brevissime 
apiculata, longitudinaliter dorso valvarum sulcata, sparse hirsutula superne; 
seminibus 1.5-2 mm. longis, subovatis vel ovato-oblongis, obtusis, nigro-b run- 
nels, paulo angulatis, prominenter transverse corrugatis, rugis circa 12, pro- 
funde alveolato-reticulatis. 

Type: No. 163032 in Rocky Mountain Herbarium, University of Wyoming, 
collected on May 19, 1911, at DeBeque, Mesa County, Colorado, by George E. 
Osterliout, No. 4458. There is a duplicate in Herb. N.Y. Bot. Gard. Phacelia 
suhmutica is known only from Osterhout's collections made at DeBeque, others 
being : No. 4726 (RM, frag. CAS) and No. 5978 (RM, frag. CAS) . 

Although P. suhmutica is obviously related to P. scopulina from which it 
was undoubtedly derived, it may be readily differentiated by a number of 
distinctive characters that indicate, each in a small way, a separate specific 
entity. Phacelia suhmutica has a rather distinctive appearance which comes 
from its less leafy and more fiorif erous habit and which is also due to the less 
cinereous character of the pubescence. In P. suhmutica the fruiting calj^x- 
segments are usually much more elongate ; and while the pedicels of the lower 
flowers are reflexed or sigmoid-spreading in fruit in P. scopulina, they are 
mostly strictly erect or ascending in P. suhmutica. There is a noteworthy 
difference in the coloration of the corolla in the two species, too, that of P. 
suhmutica being yellowish, that of P. scopulina being more or less strongly 
tinged in age with lavender or violet. The stamens in P. scopulina equal or 
exceed the corolla-throat, sometimes in anthesis even exceeding the corolla- 
lobes, while in P. suhmutica the stamens are shorter than the corolla-throat. 
Although all these characters indicate real differences between P. suhmutica 
and P. scopulina, they are not so important either specifically or diagnostically 
as the characters that are found in the style, capsule, and seed. In P. suhmutica 
the style is glabrous except for a few hairs near the base, but in P. scopulina 
the style is hirsutulous-hairy to the middle or above, generally densely so. 
The capsule of P. suhmutica differs from the capsules of other species in this 



■M2 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

section except P. tetramera, in that it is either entirely devoid of a terminal 
apiculation or has a very insignificant one. By contrast, the capsule of P. 
scopulina is prominently apiculate. The differences in the seeds, although 
small, are nevertheless appreciable: the seeds of P. suhmutica are usually 
larger and more finely and numerously corrugated, the corrugations are not so 
i-ounded and are more deeply alveolate-pitted, and the seed-cover is blackish- 
brown and tends to be iridescent. 

Hence, although P. suhmutica is undoubtedly closely related to P. scopulina, 
I believe that it is an entity sufficiently distinguished to be accorded specific 
recognition. 

7. Phacelia inyoensis (Macbr.) J. T. Howell 

Leali. West. Bot. 4:16 (1944) 
Miltitzia inyoensis Macbr., Contrib. Gray Herb., n. ser., 49:41 (1917). 

Stems erect or spreading, slender, one to several from the top of the taproot, 
2-10 cm. tall, hirsutulous and finely capitate-glandular ; leaves few and scat- 
tered, not concealing the flowers, 0.5-1.5 cm. long, 0.3-1 cm. wide, oblong to 
elliptic or obovate, entire to coarsely and pinnately few-lobed, subobtuse at 
apex, euneate at base, hirsutulous and more or less finely capitate-glandular, 
petioles of basal leaves to 2 cm. long, those of the uppermost leaves only about 
3 mm. long ; racemes lax, loosely few-flowered, to 7 em. long, the lowest flowers 
distant, the uppermost approximate, pedicels slender and spreading, 2—6 mm. 
long, hirsutulous and glandular; flowers 5-merous; calyx-segments in flower 
a little unequal, 2-3 mm. long, 0.4-0.75 mm. wide, linear to linear-oblanceolate, 
in fruit 3.75-6 mm. long, 0.5-1.3 mm. wide, linear-oblong to oblanceolate, 
somewhat spatulate, narrowed below to claw-like base, subacute, glandular 
and hirsutulous ; corolla pale yellow becoming tinged with violet, tubular- or 
cylindric-campanulate, 3-3.5 mm. long, lobes erect, 0.6-1 mm. long, corolla- 
tube hairy outside to the base, glabrous inside; corolla-scales obsolete or pres- 
ent as a mere line about 1 mm. long, free from the base of the filaments; 
stamens 1.5-2.5 mm. long, filaments glabrous, anthers about 0.2 mm. long, 
wider than long after dehiscence; style and branches 1 mm. long, branches 
0.2-0.25 mm. long, style glabrous except at the very base ; ovary 1 mm. long, 
hairy above, ovules 18-20; hypogynous disk relatively prominent with round- 
ish lobes between the stamens ; capsule 3.3-4 mm. long, elliptic, rounded- 
obtuse, shortly apiculate, sparsely hirsutulous, valves shallowly longitudinally 
sulcate on the back; seeds nearly 1 mm. long, ovatish or deltoid-ovate, acute 
at apex, rounded at base, brownish, corrugations rather i^rominent, reticulate, 
6 or 7. 

Known only from Owens Valley, Inyo Count3^ California. 

Collectio')Ks. Foothills west of Bislioi), Heller No. 8324 (G, type; CAS, DS, 
ITS) ; Big Pine, K. Brandegee in 1913 (P) ; Lone Pine, K. Brandegee in 1913 
(CAS, P). 

According to data accompanying Heller's collection, the habitat in which 



I 



» 



VOL.XXV] HOWELL: PHACELIA SECTION MILTITZIA 373 

this rare species grows is "a small dry peat bog among the granite sands." 
.Vlthoiigh it is quite a distinct species, it is probably most closely related to 
P. saliiia and through it to P. scojmlina. 

8. Phacelia salina (A. Nels.) J. T. Howell 

Leafl. West. Bot. 4:16 (1944) 

Emmenanthe foliosa M. E. Jones, Zoe 4:278 (1893). 

Emmenanthe salina A. Nels., Bull. Torr. Bot. Club 25:381 (1898). 

Miltitzia foliosa (M. E. Jones) Brand, Das Pflanzenr. IV. 251 :131 (1913). 

Miltitzia salina (M. E. Jones) Eydb., Bull. Torr. Bot. Club 40 :479 (1913). 

Xot Phacelia foliosa Phil., Anal. Mus. Nac. Chile 53 (1891). 

Xot Fliacelia salina M. E. Jones ex Brand, Das Pflanzenr. IV. 251 : 119 (1913), as synonym. 

Stems short and stout or slender and spreading, 2 to several from the top 
of a slender annual taproot, 0.5-3 cm. or even to 6 cm. long, sparsely hirsutu- 
lous, eglandulose or minutely and sparsely capitate-glandular above and in 
the inflorescence ; leaves entire, sinuate-wavy, serrate or pinnately f ew-lobed, 
0.5-1.5 cm. long, 0.3-0.9 cm. wide, oblanceolate to elliptical or obovate, 
sparsely hirsutulous, obtuse, narrowly to broadly cuneate at base, frequently 
a little asymmetrical at base, petioles 0.5-3 cm. long; racemes exceeded by 
the leaves, laxly few-flowered and only slightly scorpioid, the flowers distant 
or tending to be congested near the ground at the base of the stems, pedicels 
1-5 mm. long, straight or spreading; flowers 5-merous; calyx-segments in 
flower 2.5-4 mm. long, in fruit 4.5-6 mm. long, 0.5-0.75 mm. wide, linear to 
linear-oblong or linear-oblanceolate, sparsely hirsutulous and sometimes a 
little glandular ; corolla yellow or tinged with lavender, tubular, 3-4 mm. long, 
the lobes erect, 0.9-1.25 mm. long, the corolla hairy outside, glabrous within ; 
corolla-scales generally lacking, if present filiform and about 1.5 mm. long, 
at base free from the filaments ; stamens 1.5-2.5 mm. long, filaments glabrous, 
anthers about 0.25 mm. long, roundish ; style and branches 1 mm. long, the 
branches 0.2-0.25 mm. long, style hairy only near the base or to near the 
middle; ovary 0.75-1 mm. long, ovules 7-9 ; hypogynous disk with relatively 
prominent rounded-deltoid lobes, the lobes alternating with the stamens and 
about 0.2 mm. long ; capsule 3-4 mm. long, broadly elliptical or elliptic-ovate, 
obtuse and apiculate, sparsely hairy, the valves thin-chartaceous with thick 
conspicuous margins, back of the valves shallowly sulcate; seeds brown, 1.5--2 
mm. long, acute or subacute at both ends, oblongish to ovatish, prominently 
corrugated, corrugations 9-11, broad and rounded. 

Of rare and local occurrence in alkaline soil in southern Wyoming, western 
Utah, and northern Nevada. 

Collections. Bitter Creek, Sweetwater Co., Wyoming, A. Nelson No. 3105 
(RM, type of Emmenanthe salina; CAS, G, M, US), A. Nelson No. 4786 
(NY) ; Deep Creek Valley above Furber, Tooele Co., Utah, M. E. Jones in 
1891 (P, type of Emmenanthe foliosa; CAS, DS, G, M, UC, US) ; in loose 
red sand, west end of White Pine Mts. below Little Antelope Summit, White 
Pine Co., Nevada, Ripley and Barnehy No. 3592 (CAS) ; Reese River Valley 6 



374 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. > 

miles nortliAvest of Austin, Lander Co., Eipley and Barneby No. 5926 (CAS) ; 
ea!?t base of Railroad Pass, Lander Co., Ripley and Barnehy No. 5932 (CAS). 

There is considerable variation in the data accompanying the original col- 
lection of Emmenanthe foliosa. The specimen in the Jones Herbarium at 
Pomona College which I am accepting as the tj'^pe gives "Furber, Nevada," 
June 8, 1891, while a second sheet gives "Deep Creek, alkaline flats above Fur- 
ber, Nevada." Data on specimens in Herb. Calif. Acad. Sci. and U.S. Nat. 
Herb, are "On Deep Creek, above Furber," Utah, June 8, 1891 ; and in Gray 
Herbarium in Jones' handwriting are "Deep Creek, Utah," June 4, 1891. The 
data given in the original description are still different : "Deep Creek, Utah, 
June 6, 1891, altitude 5000 feet." After a careful study of the plants consti- 
tuting these specimens, I have concluded that all are parts of one collection ; 
and by reference to the "Botanical Exploration of Marcus E. Jones from 1875 
to 1919" (Ms.), I believe the correct data for the collection to be: Alkaline 
flats. Deep Creek Valley above (i.e. south of) Furber, Tooele Co., Utah, June 8, 
1891. According to his journal (I.e., p. 19), Jones was at Fish Springs on 
June 4, at Ibapah on June 6, and at Furber on June 8. 

Phacelia salina is intermediate between P. scopulina and P. tetramera, the 
three species apparently representing a distinct phylogenetic line. Morpho- 
logically P. salina seems more closely related to P. scopulina, differing chiefly 
in the smaller size of most parts; but ecologically it is much nearer P. tetra- 
mera, both species being usually restricted to highly alkaline or mineralized 
situations in desert regions. Phacelia salina, P. inyocnsis, P. glaherrima, and 
P. suhmutica are to be counted among the rarest of western plants. 

9. Phacelia tetramera J. T. Howell 

Leafl. West. Bot. 4:16 (1944) 

Emmenanthe pusilla A. Gray, Proc. Amer. Acad. 11:87 (1876). 
Miltitzin pusilla (A. Gray) Brand, Das Pflanzenr. IV. 251:132 (1913). 
Miltitzia pusilla (A. Gray) Brand var. flagellaris Brand, I.e. 

Not Phacelia pusilla Buckl., Amer. Jour. Sci. 45:172 (1843); nor P. pusilla Torr. ex S. 
Wats., Bot. U.S. Geo]. Explor. 40th Paral. (King's Exped.) 253 (1871). 

Stems several from the top of the taproot, spreading or ascending, 2-10 cm. 
long, slender, finely glandular- villous and near the base sparsely hirsutulous ; 
leaves narrowly elliptic-oblong to oblong-oblaneeolate, 0.4-1.5 em. long, 0.2- 
0.7 cm. wide, entire or saliently few-serrate-lobed, obtuse, narrowly cuneate at 
base, sparsely hirsutulous, petiole 0.4-1.5 cm. long, glandular-villous ; racemes 
to 8 cm. long, very laxly flowered, the lowest flowers distant, pedicels to 4 mm. 
long, slender, recurved or spreading in the lower flowers ; flowers usuallj^ 
4-merous, sometimes 5-merous ; calyx-segments in flower 2.5 mm. long, oblong 
or narrowly oblong-ovate, in fruit unequal in width, 4r-5 mm. long, 0.5-1 mm. 
wide, narrowly to broadly oblong, hirsutulous and sparsel.y and minutely 
glandular, obtuse, united at base sometimes as much as 2 mm., the base 
cuneate; corolla whitish, 1.3-1.8 mm. long, eampanulate, the lobes erect, 0.5-1 
mm. long, the lobes sparselj^ hairy outside, glabrous inside; corolla-scales 



Voi..XXV| nOWELL: PHACELIA SECTION MILTITZIA 375 

none ; stamens 1 mm. long-, filaments glabrous, anthers ronndish, nearly 0.2 
mm. across ; style 0.25-0.4 mm. long, the branches indicated only by an emar- 
gination at the top of the style; ovary roundish, 0.75 mm. long, hirsutulous at 
top, ovules 20; hypogynous disk narrow and inconspicuous; capsule oblong 
to oblong-obovate, 3-4 mm. long, very obtuse, shortly apiculate or muticous, 
sparsely hirsutulous, the verj^ base sometimes a little inferior ; seeds 1 mm. 
long, narrowly ovate to quadrate, round at base, acute at apex, light brown, 
finely and sometimes indistinctly corrugated, corrugations about 7-9. 

Alkaline flats and washes of high valleys, eastern Oregon south to central 
Nevada and eastern California. 

B.epreseyitative collections. Oregon: Union Co., Cnsick No. 758 (CM, G, 
US ; first-cited collection of Miltitzia pusilla var. flagellaris) ; Malheur River, 
Harney or Malheur Co., Cusick No. 1946 (CM, G, M, ND, RM, UC, US, WS) ; 
Wagontire, Harney Co., Peck No. 20844 (W) ; 8 miles west of Lakeview, Lake 
Co., Peck No. 15243 (CAS, NY, W). 

Nevada : 8 miles north of Twin Bridges, Elko Co., Holmgren No. 871 (lU, 
UC); Steamboat Springs, Washoe Co., 8. Watson (G, type of Emmenanthe 
pusilla; NY) ; Washoe Lake, Washoe Co., M. E. Jones in 1897 (CAS, CM, 
DS, P, US) ; Empire City, Ormsby Co., M. E. Jones No. 3987 (CAS, DS, 
lU, M, NY, P, RM, UC, US); foothills of Toquima Range near Belmont, 
Nye Co., Ripley and Barnehy No. 3696 (CAS) ; northwestern Nevada, Lem- 
mon in 1875 (G) ; Grass Valley, Lander Co., Ripley and Barnehy No. 5920 
(CAS). 

California : "Sierra Nevada Mts.," Lemmon in 1875 (US) ; Chat, Plumas 
Co., M. E. Jones in 1897 (CAS, P) ; Long Valley, Mono Co., J. T. Howell No. 
14382 (CAS, CM, G, lU, P, US) ; valley east of Cedarville, Modoc Co., Ripley 
and Barnehy No. 6005 (CAS) . 

The earliest collection of this species, Watson No. 878 from Steamboat 
Springs, was included by Torrey in hLs Phacelia pusilla; and, although Gray 
cited Watson's collection in liis original description of Emmenanthe pusilla, 
Brand annotated it in the U.S. National Herbarium as Phacelia pusilla Torr. 
and cited it as that species in his monograph (1913, p. 122) . 

The most remarkable feature of P. tetramera is the tendency for the flowers 
to be 4-merous instead of 5-merous as is usual in Phacelia. The reduction in 
the number of parts seems to result from the suppression of one part of the 
calyx, corolla, and androecium, and not to be due to the union of adjacent 
parts. Although most of the flowers that have been examined are truly 4- 
merous, occasionally they are either partly or complete! j^ 5-merous. In some 
5-merous flowers, the fifth calyx-segment is much smaller than are the other 
four ; in one case two calyx-segments were united nearly throughout, while 
the other three were deeply divided. It has been noted that 5-merous flowers 
iu-e more numerous in specimens from Oregon than in those from California 
(vr Nevada. Certainly the reduction of flower-parts exhibited by P. tetramera 
is a most unusual tendency in Hydrophyllaceae . 



;576 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4tii Skk. 

REFERENCES 

Brand, A. 

1913. Hjdrophyllaceae in A. Engler, Das Pflanzenreich IV. 251 (Heft 59). 210 pp. Wil- 
helm Engelmann, Leipzig. 
Gray, A. 

1878. Synoptical Flora of North America, vol. 2, pt. 1, 393 pp. Ivisoii, Blakeinan, Taylor, 
and Co., New York. 
Howell, J. T. 

1944. A reconsideration of the genus Miltitzia. Leafl. West. Bot. 4:12-16. 
Jepson, W. L. 

1943. Flora of California, vol. 3, pt. 2, pp. 129-464. Associated Students Store, University 
of California, Berkeley. 
Kearney, T. H., and E. H. Peebles 

1942. Flowering plants and ferns of Arizona. U.S. Dept. Agric. Miscell. Publ. No. 42:>. 
1069 pp. Government Printing Office, Washington, D.C. 
Peck, M. E. 

1941. Manual of the higher plants of Oregon. 866 pp. Binfords and Mort, Portland, 
Oregon. 

TiDESTROM, I. I 

1925. Flora of Utah and Nevada. Contrib. U.S. Nat. Herb. 25. 665 pp. Government Print- 
ing Office, Washington, D.C. ^ 



PROCEEDINGS 

OF THE 



Marine Biological LabordWiy | 

X* I 33 K. -A- R "ST 

JAN 3 1 1947 

WOODS HOLE, MASS. 



CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXV, No. i6, pp. 377-420, pis. 37-46 November 15, 1946 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS 

No. 16 

ENDEMISM IN THE HAWAIIAN FLORA, AND A 

REVISION OF THE HAWAIIAN SPECIES OF 

GUNNERA (HALORAGIDACEAE) 

HAWAIIAN PLANT STUDIES U' 

BY 

HAROLD ST. JOHN 

Professor of Botany, University of Hawaii 



E 



ENDEMISM IN THE HAWAIIAN FLORA 

NDEMisM in its flora has long made the plants of the Hawaiian Islands 



famous. Of the islands or floristic regions of the world having a flora of 
more than a few species, they are said to have the world's largest percentage 
of endemics. 

Prof. A. Guillaumin (Proc. 3rd. Pan. Pacif. Sci. Congr. Tokyo 1 : 930, 1926) 
in his review of the Pacific island floras calculates that endemism for the 
Hawaiian Phanerogams is 66 per cent. He does not state how he arrived at 
this figure but his totals of the species are evidently those of Wm. Hillebrand. 
This early botanist, Hillebrand (PI. Hawaiian Ids. p. XVII, 1888), calculated 
that of the native Phanerogams, 574 were endemic, while 246 only were in- 
digenous, and 24 of early aboriginal introduction, and the total of native 
Phanerogams and Vascular Cryptogams was 860. It is apparent that Guil- 
laumin used this total of 860 indigenous Vascular Plants and Hillebrand's 
figure of 574 indigenous Phanerogams in computing the percentage of en- 

^ This is the eleventh of a series of papers designed to present descriptions, revisions, and 
records of Hawaiian plants. The preceding papers have been published as Occasional Papers 
Bishop Museum 10(4), 1933; 10(12), 1934; 11(14), 1935; 12(8), 1936; 14(8), 1938; 
15(1), 1939; 15(2), 1939; 15(22), 1940; 15(28), 1940; 17(12), 1943; Lloydia 7:265-274, 
1944; Bull. Torrey Bot. Club 72:22-30, 1945. 

[377] 



378 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

demism, which he calculated as 66 per cent. Had he used 705 native species 
and 574 endemics, which are Ilillebrand's truly comparable figures for the 
Phanerogams, the endemism would figure out as 81 per cent, which was the 
estimate of Hillebrand, previous to his death in 1886. All species published 
since that date or unknown to Hillebrand, Guillaumin seems to reject by his 
phrase that the area presents an extraordinary number of varieties which are 
sometimes considered as species of narrow geographic localization ("presente 
un nombre extraordinaire de varietes, considerees parf ois des comme especes et 
etroitement localisees."). D. H. Campbell used the same figures from Hille- 
brand, but he correctly calculated the endemism of Phanerograms as 81.42 per 
cent (The Derivation of the Flora of Hawaii, Univ. Ser. Stanford Univ., Publ. 
15, 1919). Also R. Kanehira apparently used the same figures as he lists the 
Hawaiian endemism as 81 per cent (Proc. 6th Pacif. Sci. Congr. 4 :610, 1940). 
Guillaumin concludes (pp. 921, 931) that in the Hawaiian flora the Australian, 
New Zealand, and even Polynesian elements are almost completely lacking, 
that the flora has strong relationships with the American flora and should be 
considered a part of the IMexican floristic region. The present writer disputes 
all of these conclusions of Guillaumin. 

The interpretations of C. Skottsberg seem more truly in accord with the 
facts. He divided the Hawaiian species of Phanerogams (Bull. Bishop Mus. 
16 : 6, 1925) as 32.3 per cent Old-Pacific ; 29 per cent Australian-Polynesian ; 
26.5 per cent Indo-Malayan ; 7.5 per cent American; 3.3 per cent widespread ; 
1.2 per cent Subantarctic ; and 0.2 per cent Boreal. The widespread or Pan- 
tropic and the Subantarctic may well have immigrated by the same route as 
the other groups from Oceania. If these, then, are all added, the total is 92.3 
per cent for the plants with a probable southwestern origin that came or whose 
ancestors came as a stream of plant immigrants through Polynesia. Later 
Skottsberg discusses the genera (Proc. Linn. Soc. 151 : 182-186, 1939), their 
relationship, and probable route of immigration. Of more than 200 genera, 
18 are Pantropic, 15 Paleotropic, 49 Malesian, 12 littoral are widespread, 57 
Austral, 22 Boreal, 9 aquatics apparently Boreal, and 13 Neotropical. This 
review of the Hawaiian genera gives the same picture as with the species : a 
large majority Indo-Malayan or Polynesian, small minorities Boreal and 
American. 

The same early source, Hillebrand, was used by J. C. Willis (Proc. Linn. 
Soc. London 148:86-91, 1936) in his contemporary discussion of regional 
endemism. Hillebrand's figures should not be discounted. They were based 
on his excellent flora and were correct for his time, that is 58 years ago. Still, to 
discuss the endemism of the Hawaiian flora today and to use only Hillebrand's 
figures is not the modern scientific method, which should use all available 
evidence. Since 1886 in Hawaii there has been a great deal of arduous explora- 
tion of the rain forests and the precipitous mountains. Here and elsewhere 
a large amount of floristic and monographic work has been published. To over- 
look and reject all the work of C. B. Clarke, Heller, Rock, C. N. Forbes, 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNEBA 379 

Beccari, Leveille, Skottsberg, Cliristensen, Degener, St. John, Sherff , Yuncker, 
Christophersen, Caum, Hitchcock, Hochreutmer, Pilger, von Poelhiitz, Radl- 
kofer, Oliver, Danser, Heimerl, Keck, Krajina, Whitney, Hosaka, Fosberg, 
Egler, Lam, Croizat, Sleumer, and others, is not the way to arrive at a modern 
estimate of the known Hawaiian flora. Skottsberg used the more sound method 
of including subsequent publications (Bull. Bishop Mus. 16:5, 1925) when 
he gave a tentative estimate of the indigenous Hawaiian Phanerogams as about 
900 species. Later he revised this figure to between 1,000 and 1,100 (Proc. 6th 
Pacif. Sei. Congr. Berkeley 4 : 685, 1940) . The writer keeps a check list of the 
Hawaiian flora. There are quite a number of the recently proposed segregate 
species which he has not included, awaiting time to verify their status, but 
many will doubtless be accepted. When eventually added in, they will increase 
rather than diminish the percentage of endemism. At present the writer's 
figures for endemism of Hawaiian Phanerogams is 90 per cent, based on totals 
of 1795 indigenous, and 1614 endemic species and their subdivisions. Esti- 
mates by some others exclude all subdivisions of species, but here each is in- 
cluded as a unit. 

Some Hawaiian genera contain species that are homogeneous and wide- 
spread, occurring unmodified on all or nearly all of the large islands. Ex- 
amples are given in the two following lists. 

I. Homogeneous Widespread Species in Genera Monottpic in the Hawaiian Islands 

Freycinetia arhorea Gaud. 

Joinvillea GaudicJiaudiana Brogn. and Gris 

Dianella sandwicensis H. and A.^ 

AnoectocMlus sandwicensis Lindl. 

Liparis Jiawaiiensis Mann 

Hahenaria holochila Hbd. 

Pilea peploides H. and A. 

Argemone alba Lestib. var. glauca Prain 

Osteomeles anthyllidi folia Lindl. 

Erythrina sandwicensis Degener 

Strongylodon lucidus Seem. 

Plumiago seylanica L. 

Osmanthus sandwicensis (Gray) B. and H. 

Nertera depressa Banks and Soland. 

II. Homogeneous Widespread Species in Genera with Several Hawaiian Species 

Smilax sandwicensis Kunth 
Chenopodium oahuense (Meyen) Aellen^ 
Peperomia leptostachya H. and A. 

There are other Hawaiian species widespread among the larger islands, but 
which are not homogeneous, having a tendency to vary. These variations are in 
part recognized as described varieties but in large part are unrecognized, 

2 The recently described Dianella lavarum Degener and D. multipedicellata Degener are 
considered exact synonyms. 

3 Chenopodium sandwicheum Moq. forma macrospermum Aellen and forma microspermum 
Aellen are not separable, as the larger and smaller seeds occur on a single plant. 



380 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

forming a part of the heterogeneous population now included in the species. 

Such ones are listed below. 

TIT. Polymorphic Widespread Species 
Eumex giganteus Ait. 
Charpentiera oiovata Gaud. 
Charpentiera ovata Gaud. 
Broussaisia arguta Gaud. 
Acacia Koa Gray 
Ilex anomala H. and A. 
Cheirodendron Gaudichaudii (DC.) Seem. 
Vacci7iium calycinum Sm. 
Vacciniutn dentatum Sm. 
Styphelia Tameiameiae (Cham.) F. Muell. 
Myrsine Lessertiana A.DC. 
Gouldia terminalis (H. and A.) Hbd. 

Other genera, usually with many species, show a conspicuous segregation 

into species, usually each species being restricted to a single island or even to 

a single mountain range or valley on the particular island. Such genera are 

the following : 

IV. Genera with Numerous Species that are Narrow Endemics 

Astelia, Schiedea, Pelea, Fagara, HibisctirS, Viola, Labordia, Haplostachys, Phyllos- 
tegia, Stenogyne, Cyrtandra, Coprosma, Hedyotis, Clermontia, Cyanea, Delissea, 
Lobelia, Bidens, Bubauiia, and Lipochaeta. 

The Hawaiian plants here listed illustrate a progressive series : 1, genera 
monotypic in Hawaii occurring as invariable wides on all the principal 
islands ; 2, species that are invariable wides, belonging to genera with several 
Hawaiian species; 3, species that are wides, but showing variabilitj^ on the 
various islands ; and 4, general with many Hawaiian species, typically with 
different, distinct species on each island. These species represent degrees of 
increasing differentiation and presumably of decreasing age. Thej' show evi- 
dences of speciation. The results of this species formation can be seen in the 
percentages of endemism here presented and discussed. 

REVISION OF THE HAWAIIAN SPECIES OF GUNNERA 

One of the most famous of the genera occurring in the Hawaiian Islands is 
Gunnera, called by the natives "apeape." It is a giant herb with a stem 12 to 
15 cm. or more in diameter, fleshy, 2 to 6 meters tall, the base decumbent and 
prostrate. The paired stipules are large, pale, and conspicuous. From near the 
summit are produced several alternate leaves on fleshy petioles 6 to 13 dm. 
long. The blades are rounded, shallowly lobed, like great parasols 4 to 20 dm. 
in diameter. Axillary panicles 8 to 10 dm. long bear numerous minute green 
flowers and later tiny yellow, orange, red, or purplish drupes. The plant itself 
is so bizarre as to hold the attention and interest of all observers, whether 
scientific or not. Thus its occurrence is shown even on the tourist maps. 

The habitat of the "apeape" is also noteworthy. It occurs on precipitous 
slopes with the soil saturated from the run-off of the boggy mountain summits. 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUN NEB A 381 

It is usually on the face or the foot of a precipice ("pali" in Hawaiian) and 
is often difficult of access. It forms a definite vegetational belt between the 
Middle Forest and the Upper Forest, at anywhere from 2,500 to 5,000 feet 
altitude. Because of the distinctive, giant leaves, this belt of "apeape," though 
narrow, is often so conspicuous that it can be seen from a distance of two or 
three miles. 

The "apeape" was discovered in 1819 by Charles Gaudichaud, who published 
it in 1830 as Gunnera petalo'idea. Subsequent collections have revealed that 
Gunnera occurs on Kauai, Oahu, Molokai, Maui, and Hawaii. Hillebrand in- 
terpreted these as different locality records for the single species. This has 
been the generally accepted view, that the same species occurred on all of 
the islands. 

The taxonomic history since the original publication by Gaudichaud has 
been brief. Alphonse De CandoUe described a specimen collected on Kaui 
[= Kauai] by the United States Exploring Expedition as G. petaloidea Gaud. 
13 Kauensis A.DC. (De Candolle, Prodr. Reg. Veg. 16 (2) : 597, 1868) . His four 
word description included only two characters, almost glabrous, the bracts 
shorter. We now find no differences in the Kauai plants in pubescence, and 
the bracts appear to be alike. This plant was treated later as an unnamed 
variety, G. petaloidea Gaud. /? (Hillebrand, Fl. Haw. Ids. 124, 1888). In 1930 
it was again distinguished by two botanists in almost simultaneous publi- 
cations. Gunnera kauaie^isis Rock in Caum (Occ. Pap. B. P. Bishop Mus. 
9(5) : 3^, pi. I, 1930, Sept.). Rock described it from a new type specimen. 
Rock, 5,053 from Mt. Waialeale. The second author, Krajina, redescribed it 
as G. Dominii Krajina (Acta Bot. Bohemica 9 : 50, ill. p. 51, 1930, without 
precise date), citing as synonyms the varieties of De Candolle and of Hille- 
brand, but listing only his own collection from "montis Waialeale." It is 
apparent from the characters given below that this Kauai plant with peltate 
blades and red to purplish drupes is a distinct species. Several attempts to 
learn from Prague the exact month and day of publication of Krajina's paper 
have failed. A new search is being made for the date. In the meantime, the 
name given by Rock is accepted as it is dated to the month. 

Krajina also described and illustrated from his own collections G. peta- 
loidea. He published as two new varieties: var. kaalensis (p. 50) from his 
collection on Puu Kaala, Oahu ; and var. mauiensis from his collection by 
Waikamoi Stream, east Maui. Krajina apparently did not examine Gaudi- 
chaud's type specimen, nor did he indicate whether or not either of his two 
varieties was to be considered var. typica though with a different name. Dupli- 
cates of Krajina's collections have not yet reached the Bishop Museum, nor 
has the writer seen them in other herbaria. Still, there is no difficulty in match- 
ing them. Dr. V. Krajina was a fellow at the Bishop Museum in 1929-30, 
a keen, energetic botanist who soon became my good friend and I took him on 
many field trips. His t^'pe of var. kaalensis was collected on my class trip to 
Puu Kaala at the well-known locality above the fire-brick trail from Schofield 



382 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4™ Ser. 

Barracks. My collection, St. John 10,072, was made at the same spot and time 
as Krajina's and there are numerous other collections from this locality. His 
type of var. mauiensis was collected at another often visited spot where the 
Olinda Pipe Line Trail crosses Waikamoi gulch. There are several other col- 
lections from the same spot. 

That recounts the known taxonomic history of this genus in Hawaii. The 
plant is known to all collectors and there are several standard, fairly easily 
accessible localities. The writer has collected it on Kauai, Oahu, Maui, and 
Haw^aii, but any new localitj' is always noteworthy. On Oahu it has been re- 
peatedly collected on the wet, windward side of Mt. Kaala in the Waianae 
Mountains. There are two or three collections from the drier, leeward side of 
the same mountain. Also, Miss Marie Neal of the Bishop Museum tells the 
writer that she has observed Giomera in the same mountain range on the adja- 
cent peak, Puu Kalena, on its northeast or windward side. 

The preferred habitat is steep, water-soaked slopes on the windward side 
of a mountain range. It has long puzzled the writer that the "apeape" was 
abundant in the Waianae Mts. and not in the Koolau Range, the more easterly 
mountain chain on Oahu. The Waianae Mts. are rather arid, lying in the lee 
of the Koolau Range which gets the impact of the moist northeast trade winds 
and receives most of their moisture, while but little of it reaches the Waianae 
Mts. Only the highest peak, Mt. Kaala (called Puu Kaala on the earlier, 1917, 
official topographic map) gets a considerable rainfall. It is just far enough to 
the north to receive the impact of the northeast trade winds which sweep over 
or past the descending, low^ north end of the Koolau Range. This is only about 
1,000 feet high at this point, not high enough to cause the precipitation of all 
the moisture. Hence, Mt. Kaala with its large flat summit at 4,025 feet altitude 
and its upper ridges get enough rainfall to support a luxuriant rain forest. 
The summit has a swampy forest except for some boggy openings near its 
center. The average annual rainfall is now known to be 91 inches (Feldwisch, 
W. E., First Progress Rept., Hawaii Territorial Planning Board 121, 1939). 

The Koolau Range, Oahu, running from southeast to northwest, forms the 
backbone of the eastern side of the island. It rises abruptly close to the shore, 
and the section above 1,500 feet in height extends for about 25 miles, 
making a barrier almost exactly at right angles to the prevailing northeast 
trades. There are numerous sharp peaks that rise from the range, the highest 
being Puu Konahuanui with twin peaks 3,105 and 3,150 feet high. The several 
rain gauging stations show a high rainfall, as this mountain range catches 
much of the moisture from the clouds. The rainfall is more than 100 inches 
annually for most of this ridge and at the wettest station is 311 inches. There 
are miles of wind-swept precipices ("pali") and steep slopes with much water 
seepage from the higher peaks, just the kind of habitat that usually supports 
Gunnera. The writer had botanized these brinks and declivities at many locali- 
ties along the Koolau Range, and had never found the "apeape." Yet, this 
Gunnera is a plant that one cannot easily overlook. With its bizarre leaves 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNEBA 383 

like huge iTmbrellas, it is always the biggest and most conspicuous plant in its 
area. There were no specimens from the Koolau Range in the Bishop Museum, 
yet the plant has been reported from there. Prof. Vaughan MacCaughey listed 
it as rare in the rain forests of Manoa Valley (Am. Journ. Bot. 4 : 600, 1917). 
In the same year in a short article about Gunnera petaloidea (Am. Journ. 
Bot. 4 : 38, 1917, the reprints incorrectly printed vol. 1) he stated that among 
its typical habitats were "Ka-ala and Kona-hua-nui summit ridges on Oahu 
(2,500-4,000 ft.) ." The 4,000 foot record is certainly that for the often visited 
station of Mt. Kaala in the Waianae Mts. Hence, we infer that his station on 
the 3,150 foot peak Puu Konahuanui was at about 2,500 feet altitude. Mac- 
Caughey did exploring and original research in Manoa Valley which heads 
on the slope of Puu Konahuanui. The plant is unmistakable, so there would 
seem no reason to question this record, yet corroborating specimens were much 
to be desired. 

Some years ago the writer had an enterprising young man as a student, 
John R. Coleman. He was enlisted in the United States Marine Corps and was 
stationed in Honolulu with a company doing guard duty at the docks. At his 
request he was given continuous and regular night guard duty. This left his 
day time free and enabled him to attend the University of Hawaii. It was not 
obvious when he slept, but he did not sleep in class. He completed the course 
on elementary botany and on week ends he often joined advanced classes on 
collecting trips to the mountains. One Sunday he induced two other marines 
to join him, and from Nuuanu Pali they started to climb the knife-edged ridge 
that culminates in Puu Konahuanui. As he led the way and scrambled up one 
precipitous slope, the bushes, soil, and all caved away. He managed to make 
headway over the landslide and surmounted the ledge. All the footholds were 
destroyed, so his companions could not follow, and turned back. Coleman went 
on, climbed to the summit, and descended by a more southerly route. The next 
day he brought me a 10 cm. tip from a fruiting inflorescence with half ripe 
drupes, of a specimen that seemed to be Gunnera. He reported collecting it 
near the summit on Oct. 19, 1930. This specimen gave partial proof of the 
occurrence in the Koolau Range, but it was incomplete. At last, in June 1942 
the proof was furnished by two students, L. Eubank and A. D. Conger, who, on 
a wet day amid swirling clouds and rain, ascended Puu Konahuanui. They ob- 
served and collected abundant and complete material. They reported the three 
small colonies to be on the Manoa or leeward slope. A second visit on Sept. 
13, 1942 by A. D. Conger, H. St. John, and R. P. St. John w^as also in the clouds 
and intermittent rain, but thrice the clouds lifted to reveal the slopes, valleys 
and the shore line. All three patches proved to be on the windward or Kailua 
side. Any mountaineer will understand how in following- forking, intricate 
ridges in dense clouds, one can lose track of the directions. One locality was 
a single plant some 250 feet below the lower or south peak which is mapped 
and triangulated as Puu Konahuanui. The two others were on the east face of 
a northeast ridge of the north or higher peak. There were numerous plants in 



384 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

a steep, brushy, hanging valley at 2,950 feet altitude. Though both Miss 
Eubank and Mr. Conger previously slid down the cloud-blanketed slope to 
the plants, then scrambled up again, on the second trip no one wanted to try | 
it again. It was very dangerous, and when clear of clouds it was obvious that 
an uncontrolled slip on the steep, slippery clay would give one a flying start 
over the 1,000 foot precipice. Later a fourth colony was located. On May 9, 
1943 the writer conducted a student party of nine from Tantalus to the sum- 
mit of Puu Konahuanui. Clouds blanketed the precipices on the east slope 
hiding the single plant on the South Peak. From the North Peak a descent was 
made down the secondary ridge towards Kailua. The middle and inaccessible 
colony of Gunnera was again seen far below in the cloud. Neither the writer 
nor his son Robert could locate the third colony, though this time a rope was 
at hand for a descent to it. The clouds hid it completely. Climbing up again, 
the descent was made along the main divide to the road at Nuuanu Pali. On 
this divide a new colony was found, also on the east face, in a moist, precipi- 
tous, hanging valley at 2,300 feet altitude. By the use of a rope the colony was 
visited and specimens collected. There were more than 12 plants, up to 4 
meters tall, some exposed, others half hidden by the thicket of Pipturus alhidus 
and Cyrtandra paludosa. The specimens collected have been deposited in the 
Bishop Museum. These plants from the Koolau Range were critically com- 
pared with those from the east side of Mt. Kaala in the Waianae Mts. and 
were found to be identical in structure and quite inseparable, though the 
blades are smaller, the length along the midrib not exceeding 4 dm. 

While investigating this material the writer took occasion to assemble all 
available specimens and to evaluate the species and varieties described from 
the other Hawaiian Islands by Rock and by Krajina. Its massive fleshy stems, 
giant fruit clusters, and rounded leaves, 1 to 2 meters in diameter, cannot well 
be compressed upon a standard herbarium sheet, so that the collector is forced f 
to take sample fragments which are inevitably incomplete, and the leaves 
selected are often the smaller ones. Hence, all taxonomists revising the group 
and writing descriptions from the available herbarium material present meas- 
urements which are true as far as they go, but minimize the size of the giant 
stems, leaves, and inflorescences. Several of the writer's whole numbers in- 
cluding many unmounted duplicates are available, but still there are gaps. 
From east Maui there are no mature stipules, and from west Maui there are 
no petals, and mature drupes and stones are needed from Molokai and from 
Makaha Valley, Oahu. Still the collections are numerous and have proven 
sufficient. At a glance they all look alike, but on close study differences are^ 
noted in the floral and fruiting morphology. It is curious that most of the 
characters are minute, a difference in shape to be seen with a lens or in size 
measured in fractions of a millimeter. Seedlings are often collected. Since they i 
produce smaller stipules and blades that are much smaller and more renif orm,^ 
they have not been included in our descriptions. The only large parts furnish- 
ing characters are the stipule.s and the leaves as to lobing and toothing and! 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNEBA 385 

pubescence. When contrasted throughout, several correlated characters mostly 
of floral and fruiting anatomy are found. When separated on these characters, 
the segregates are found to agree in leaf and stipule characters and to have 
good geographic correlations. Each island has one local kind, except Maui and 
Oahu which have two each. When evaluated, the number and sort of char- 
acters would indicate that these segregates are properly considered species. 
The original species was Gunnera petalo'idea Gaudichaud (Voy. "Uranie," 
Bot. 512, 1830). The plant was not illustrated and the 16 word description 
includes only the major features, but none of the ones now found serviceable 
in distinguishing the local species. Neither Rock nor Krajina gave any dis- 
cussion as to the nature of G. petalo'idea Gaud, or its type locality. That pub- 
lished by Gaudichaud was "In insulis Sandwicensibus (Alt. 450-500 hex.)." 
No island is specified, and the statement of altitude in fathoms is unusual, but 
was perhaps natural after months at sea on a sailing vessel ! Search has been 
made for any detailed itinerary of Gaudichaud. He published none, and the 
narrative by the commander, Capt. Louis de Freycinet, and those by other 
members of the expeditions give much on the physique and social habits of the 
native inhabitants, but nothing on the route of the collecting botanist. On the 
"Uranie" voyage they spent 20 days in the Hawaiian Islands and Gaudichaud 
was on shore 7 days. The writer has long marveled at the large number of 
the endemic species and genera that he collected and described on this first 
voyage. Starting from the harbor, with no good roads, no cut mountain trails, 
to get up into the dense, native forests and through them to the cloud zone 
was a herculean task. Then, over half of his collections were lost when the 
homeward bound ship was w^recked on the Falkland Islands. All were soaked 
in sea water, but he rescued many by drying them before a bonfire on the 
beach. These experiences are mentioned in a few brief words, but what stark 
tragedy they describe ! What a grand naturalist he must have been, yet no 
portrait of him is known ! He did not specify where his shore trips were made, 
but the ship made landings on Aug. 8, 1819 at Kayakoua [Kailua, Hawaii] ; 
Aug. 12 at Bay of Kohaihai' [Kawaihae, Hawaii] ; Aug. 17-25 at Raheina 
[Lahaina, Maui] ; Aug. 26-30 at d'Onorourou [Honolulu, Oahu] . So the seven 
days on shore were divided among these localities, but just how is not known. 
In one day from the^arid regions near Kailua and Kawaihae, it would not have 
been possible to ascend to the habitat of Gunnera on the far side of the Kohala 
Mts. Hence, the island of Hawaii can be eliminated. Back of Lahaina, Maui, 
the mountains rise abruptly from near the shore, and on the steep upper slopes 
of the valleys Gunnera is abundant. From Honolulu it would have been possi- 
ble to find Gunnera in either mountain range. However, the east cliffs of Mt. 
Kaala were distant and difiicult of access. It is unlikely that he could have 
gotten there. Puu Konahuanui is only 6 miles in an air line from the harbor 
of Honolulu. It is known that he climbed to the cloud zone at the crest of the 
Koolau Range, because he collected Lobelia Gaudichaudii which is restricted 
to that area. Hence, it can be deduced that Gaudichaud probably collected the 



386 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Gunnera in the mountains of west Maui or in the Koolau Range near Hono- 
lulu, Oahu. In 1935 the writer photographed in Paris Gaudichaud's type speci- 
men of 0. petalo'idea, reproduced here as PI. 38. The data on the sheet read 
"90. Uranie, Ins. Sandwich. C. Gaudiehaud." Stipules are lacking and the 
pieces of the inflorescence do not permit one to detect the minute floral differ- 
ences from the photograph. However, the margin of the leaf is well shown. 
The distinct, rounded lobes and the low, rounded teeth or crenations of the 
leaf margin match exactly those of the species of west Maui, so that it is to 
be taken as the type locality. 

The Hawaiian species all belong to the subgenus Panke (Molina) Schindler 
(Engler, Pflanzenreich IV, fam. 225:117, 1905). This subgenus has a re- 
markable distribution, occurring also in the Cordilleras from Mexico to Chili, 
and on Juan Fernandez (Skottsberg, C, Proc. 6th Pacif. Sci. Congr. Berkeley 
4: 696, map 14, 1940). 

In the ensuing systematic treatment are given key, diagnoses, and descrip- 
tions of the species as at present understood. 

All the specimens examined and cited and the new types are in the B. P. 
Bishop Museum, Honolulu, except the Gaudiehaud type of G. petalo'idea 
which is in the Museum d'Histoire Naturelle, Paris. 

Key to the Hawaiian Species op Gunnera 
Blades peltate ; drupes red to purplish 7. G. Tcauaiensis Eock 

Blades basifixed; drupes yellow, orange (or reddish in G. mauiensis) 
Stipules scurfy on back or finally glabrate, 

Petals entire, obtuse, glabrous ; sepals 0.6—0.8 mm long, rounded, lacerate ; anthers 
1.2-1.3 mm. long ; peduncle bractless except at base ; blade margin lack- 
ing mucros 5.G. Tcaalaensis (Krajina) St. John. 

Petals lacerate-fimbriate, often acuminate; sepals 1-1.2 mm. long, broadly deltoid, 
deeply lacerate; anthers 1.6-2.2 mm. long; peduncle bracted; (stipules 
apparently scurfy) ; teeth of blade margin with revolute mucros 

6. G. maTcahaensis St. John. 
Stipules not scurfy on the back, 
Stipules glabrous, 

Sepals 1.5-2 mm. long, ovate, deeply lacerate, the lacerations fimbriate; petals 
entire 1. G. petaloidea Gaud. 

Sepals 0.6-0.8 mm. long, broadly deltoid to rounded, lacerate ; petals minutely 
lacerate 2. G. moloTcaiensis St. John. 

Stipules pubescent. 

Petals 1.5 mm. long, entire or somewhat erose; sepals 0.4—0.6 mm. long, semi- 
oval, the obtuse tip minutely ciliate; anthers 1.9-2 mm. long, elliptic 
oblong 3. G. mauiensis (Krajina) St. John. 

Petals 0.8-1 mm. long, sparsely lacerate; sepals 1-1.2 mm long, oblong lan- 
ceolate, the tip lacerate; anthers 1.5-1.7 mm. long, oval 

4. G. Eastwoodae St. John. 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNERA 387 

1. Gunnera petaloidea Gaudichaud 

(Plate37, fig. d; pis. 38, 39) 
Gunnera petaloidea Gaudichaud, Voy. "Uranie," Bot. 512, 1826 (=: 1830). 

Giant herb, 1-6 m. tall; stem 2-5 m. long, 15-22 cm. in diameter, fleshy; 
often long decumbent, the surface with prominent leaf scars and many oval 
pits from which prop roots develop on older, mostly decumbent parts of the 
stem ; roots reddish, slimy at tip ; stipides 5-9 cm. long, oblanceolate, pale, 
strongly nerved, glabrous; petioles 2-4 dm. long, hirsute, swollen at the base ; 
blades 27-53 cm. long, cordate-reniform, shallowly palmately 9- or 11-lobed, 
firm chartaceous, dark green, glabrous or nearly so, impressed reticulate 
nerved, the veins sparsely hirsutulous, below the surface paler green, the veins 
even to the tertiary ones densely hirsute, the lobes 1.5-6 cm. long, broadly 
deltoid, the midrib and prominent lateral veins each ending in a prominent 
deltoid dentation, the margin crenate or low rounded dentate, the sinus 
broadly V-shaped ; inflorescence probably axillary ; bracts at base of inflor- 
escence 3-7 cm. long, lanceolate to oblanceolate, like the stipules; peduncle 
5-16 cm. long, hinsute, bractless; inflorescence 3-5.5 dm. long, 12-25 cm. in 
diameter, lanceoloid, hirsute towards the base, more or less glabrous towards 
the tip, the principal lower bracts 20-28 mm. long, broadly linear, acute, the 
margin membranous, the lateral branches ascending or diverging, spiciform ; 
flowers perfect, sessile; ovary barrel-shaped; calyx lobes 1.5-2 mm. long, 
ovate, deeply lacerate, then the lacerations fimbriate; petals 1.5 mm. long, 
obovate, cucullate, entire, glabrous ; the 2 anthers opposite the petals, anthers 
1.6-1.9 mm. long, narrowly elliptic-oblong; the 2 stigmas 2 mm. long, terete, 
densely puberulent throughout; drupes 2.5-3 mm. in diameter, subglobose, 
greenish-yellow; stone 1.5-1.8 mm. long, 0.9-1.3 mm. wide, 0.9-1 mm. thick, 
oval in outline, 4-angled, 4-sided, two of the angles prominent, the third less 
so, and the fourth still weaker and unequal, the stone laterally compressed, 
the surface dull, pale yellowish browaiish. 

Type: "In insulis Sandwicensibus (alt. 450-500 hex.)." The type specimen 
in Paris is labeled "90, lies SandAvich, C. Gaudichaud." This is illustrated in 
our Plate 38. 

Collections studied, all from western Maui: Eeka [of Hillebrand = Puu Kukui], ex herb. 
Lydgate, W. Hillebrand and J. M. Lydgate; gulch sides by Mt. Kukui, Sept. 24, 1916, 
G. C. Munro 426 and 594; Hanakaoo, shingles the valley sides in places, May 1910, C. N. 
Forbes 69.M; Olowalu Valley, central ridge, far up on the peak, large patches, May 21, 
1920, C. N. Forbes 2,433.M; without locality, J. F. RocTc and fi'[osme)-] 8215. 

A discussion of the type locality is given in our introduction. Gaudichaud 
could have discovered Gunnera either on Oahu or on western Maui. Neither 
his publication nor the type specimen in Paris gives any hint as to the island 
concerned. Luckily there are some differences in leaf shape, and by careful 
examination of the lobing and dentation of the leaf margin it has been possible 
to match the type with recent collections from near Lahaina, Maui. Hence, 



388 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

it is deduced that Gaudichaud collected the type in the mountains back of 
Lahaina, Maui. 

Most previous authors have combined under G. petalo'idea all of the Ha- 
waiian specimens. This was due, doubtless in large part, to the fact that few 
if any of them had sufficient material to allow the recognition of any diag- 
nostic characters between the specimens from the different islands. Conse- 
quently, their descriptions are mostly composite ones, so that one cannot now 
rely upon them. This also applies to the monographic treatment by A. K. 
Schindler (Engler's Pflanzenreich IV, fam. 225:117, fig. 34, 1905). The de- 
tails that he shows in his figure 34 of the blade, petal, and anther agree with 
the characters of the common Oahu plant. The entire, broadly triangular 
calyx lobes are different from any known Hawaiian plant, and it seems that 
they are incorrectly delineated. He cited one by Gaudichaud and a total of 
six collections, originating from three islands, so it is quite possible that his 
drawing was a composite one made from several of these collections. Hence, 
this Schindler drawing is considered to be in large part one of G. kaalaensis, 
but not fully accurate. 

Gaudichaud in 1830, when he described G. petalo'idea, in a footnote depre- 
cated his choice of a specific name which alluded to the presence of petals. He 
had learned meanwhile that some other species of Gunnera had petals, but 
nevertheless he retained the name of G. petalo'idea since he had mentioned it 
already in his introduction (p. 98) printed in 1827. 

Most recent botanists have misspelled the named as G. petaloidea, but 
Gaudichaud clearly published it as G. petalo'idea, spelling it thus with a 
diaeresis over the i to indicate a separation of the syllables, and spelling it 
consistently on two remote pages. This was not an error, but rather his delib- 
erate choice, so we must follow his spelling despite its strange appearance in 
a latinized name. 



4 



4 



I 



2. Gunnera molokaiensis St. John, sp. no v. 

(Plate 38, fig. e) 

Herba gigantea, laminis 43 cm. longis cordato-reniformibus 9-lobatis chartaceis infra 
pallidioribus nervis reticulatis hirsutulisque, lobis 5-8 cm. longis sinuato-rotundatis denti- 
culatis, inflorescentiis 15-40 cm. longis laneeoloideis, ramulis spicatis, sepalis 0.6-0.8 mm. 
longis late deltoideis vel rotundatis laceratis, petalis 1.5 mm. longis cueullatis spatulatis 
minute laceratis vel subintegris, antheris 1.5-1.9 mm. longis ovalibus, fructibus ignotis. 

Giant herb ; stipules 2 cm. long, lanceolate, glabrous ( ?) ; petiole (one seen) 
47 cm. long, with scattered dark pustules especially at base, densely hirsute at 
base, sparsely so to glabrate elsewhere; blade 43 cm. long, apparently cordate- 
reniform, shallowly palmately 9-lobed, firm, chartaceous, above dark green, \ 
the surface and veins remotely hirsutulous to glabrate, below pale green, the 
prominent reticulate veins all scabrous hirsutulous, the lobes 5-8 cm. long 
sinuous rounded, the outline ovate, the sinus U-shaped, the margin irregularly 
rounded denticulate ; inflorescence probably axillary ; peduncle 7 cm. long 
(when young) , bractless, tomentose towards the base; inflorescence 15—40 cm. 



Vol. XXV] -ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNEBA 389 

long, 4-15 cm. in diameter, cylindric to lanceoloid, tomentose on the axis and 
towards the base, glabrate above, the principal lower bracts 8-12 mm. long, 
linear-spatulate, the margin membranous, the lateral branches ascending, 
spicif orm ; flowers perfect, sessile ; ovary barrel-shaped, glabrous ; calyx lobes 
0.6-0.8 mm. long, broadly deltoid to rounded, lacerate ; petals 1.5 mm, long, 
cucullate, spatulate, minutely lacerate or subentire ; the 2 anthers opposite the 
petals, anthers 1.5-1.9 mm. long, oval; stigmas immature; drupes immature. 

Tijpe: Molokai, Pelekunu Pali, July 12, 1912, C. N. Forhes 305. Mo. B. P. 
Bishop Museum, Honolulu. 

Forbes was the only one to bring back Gunnera from the remote precipi- 
tous, windw^ard brink of the Molokai mountains. His locality was on the old 
trail which started from Kamalo, ascended the broad ridge to the cabin near 
Puu Kolekole, then northeastward across the swampy forest and bogs to the 
divide near Kaunohua at about 4,535 feet altitude, then descended in switch- 
backs down the steep head wall on the north side. This deeply eroded Pele- 
kunu Valley is a spectacular bit of mountain scenery. The Gunnera was prob- 
ably found just below the windward or northern crest of the divide. 

The writer has a record in his field notebook of observing Gunnera in 
Waikolu Valley, the next one to the west. More exactly this was at Hanalilolilo, 
near the head of the deeply eroded part of the valley. He did not get close 
enough to collect the specimens. 

G. molohaiensis is most closely related to the species of western Maui, G. 
petalo'idea Gaudichaud, which has the blades shallowly lobed, with lobes 1.5-6 
cm. long, the margin crenate or low rounded dentate ; the main veins beneath 
densely hirsute ; the sepals 1.5-2 mm. long, ovate, deeply lacerate, the lacera- 
tions fimbriate; and the petals entire. G. molohaiensis has the blades with 
marked, well rounded lobes, the lobes 5-8 cm. long, the margin irregularly 
rounded denticulate; the main veins beneath sparsely tomentose to early 
glabrate; the sepals 0.6-0.8 mm. long, broadly deltoid to rounded, lacerate; 
and the petals minutely lacerate. 

3. Gunnera mauiensis (Krajina) St. John, comb. nov. 

(Plate37,fig. g;pl. 40.) 

Gunnera petaloidea Gaudichaud var. mauiensis Krajina, Acta Botanica Bohemica 9 : 50, 

figs. B2, F2, Ij, 1 pi. 1930. 
Illustrations : by Bonine, in Eock, J. P., Monograph of the Hawaiian Lobelioideae, Mem. 
Bishop Mus. 7(2) : pi. 34, 1919; by Krajina, see above; by Degener, O., PI. Hawaii Natl. 
Park, pi. 66, 1930. 

Giant herb about 2-3 m. tall ; stem about 1-2 m. tall, the tip assurgent, the 
stem up to 6 m. in length and 15-20 cm. in diameter; bark near apex hirsute, 
with numerous glabrous, oval pits, where prop roots may later develop ; 
terminal bud completely covered by the numerous imbricate stipules ; stipules 
of seedlings 1.5-2 cm. long, ovate, membranous, strongly nerved, slender 
hirsute on the back, those of mature leaves 7.5-10 cm. long, elliptic-lanceolate 
with a broad truncate base, pale greenish, strongly nerved, sparsely puber- 



390 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

ulous base and tip ; petioles 3-10 dm. long, pustulate, densely hirsute at base, 
sparsely so elsewhere; blades 30-56 cm. long, cordate-reniform, shallowly J 
palmately 7-11-lobed, firm chartaceous, above dark green, remotely hirsutu- 
lous, soon glabrate, below pale green, the prominent veins hirsute, the reticu- 
late veinlets hirsutulous, the lobes 2-A cm. long, low rounded sinuate, the 
midrib and one or more principal lateral veins ending in low, deltoid denta- 
tions, the sinus broad U-shaped, the margin with low broad denticulations ; 
inflorescence supra-axillarj' ; bracts at base of inflorescence 5 cm. long, ob- 
lanceolate, membranous, glabrous ; peduncle 15-30 cm. long, hirsute, bractless ; 
inflorescence 4.3-10.5 dm. long, 9-24 cm. in diameter, lanceoloid, the axis 
hirsute, the principal low^er bracts linear-spatulate, acute, with membranous[ 
margin, the lateral branches ascending, spiciform, hirsute at base, glabrous] 
towards the tip; flowers perfect, sessile ; ovary depressed globose ; calyx lobes 
0.4—0.6 mm. long, semioval, the obtuse tip minutely ciliate; petals 1.5 mm. long, 
broadly cucullate-spatulate, glabrous, entire or somewhat erose ; the 2 anthers 
1.9-2 mm. long, elliptic-oblong; the 2 stigmas 2-3 mm. long, densely puberu- 
lous ; drupes 2.5-3.5 mm. in diameter, globose, pale salmon-red ; stone 1.6-1.9 
mm. long, 1.4—1.5 mm. wide, 1-1.2 mm. thick, broadly obovate in outline with 
three prominent angles of which one is lower due to a slight compression of 
the stone, and a fourth angle more or less developed or wanting, the surface 
pale yellowish brown, dull. The fresh flowers are rose-magenta. 

Type: "Maui : Makawao District ad plurimos rivulos fluminis Waikamoi in 
declivibus septentrionali-orientalibus montis Haleakala, ca. 1275 m. s. m., legi 
15. IV. 1930." V. Krajina. Herb. Charles Univ., Prag-ue, not seen. 

Collections studied, all from eastern Maui: Olinda, along pipeline from Idlewild for 
about 7 miles in deep wet ravine, July 16, 1927, 0. Begener and H. Wiebhe 3,404; the same, 
July 30, 1927, D. L. Topping, herb. Begener 8,185 ; and June 19, 1927, 0. Begener and H. 
Wieilce 2,240, 2,240a, 2,240}); Kula Pipe Line, 4,000 ft. alt., bank of Waikamoi Stream, Feb. 
11, 1930, K. St. John 10,302; Waikamoi, rain forest, 4,250 ft., Aug. 14, 1933, M. Neal and 
C. Hartt; Haiku Uka, Waikamoi Gulch, steep side walls, 4,400 ft. alt., Aug. 4, 1943, E. St. 
John 20,358. The foregoing are all from the same station on the pipe line trail east of Olinda. 
Ukulele, Sept. 4. 1919, C. N. Forbes 1,255.M; Haipuaena Stream, sides of wet valley, el. 
4,400 ft., Nov. 23, 1927, G. C. Munro 740 ; below Koolau Gap, Haleakala, extremely rainy, 
wet region, June 17, 1927, 0. Begener 8,184 ; Nahiku, Kuhiwa trail, Koolau Forest Eeserve, 
6,000 ft. alt., wet woods, Jan. 2, 1937, E. St. John and E. J. Catto 17,935; Kipahulu Valley, 
northerly side gulch near head of Oheo Stream, wet thicket, 5,800 ft. alt., Aug. 22, 1945, 
E. St. John and A. L. Mitchell 21,104. 

Most of the localities are in one general region, within a few miles of 
each other. They are on the windward slope of Haleakala between 4,000 and 
6,000 feet altitude. Here in the very wet forests are numerous narrow gulches. 
On their steep, often dripping side walls, the Ounnera is abundant. 

On the drier southeast side of Haleakala, Gunnera was reported in Kipa- 
hulu Valley by Mr. David T. Fleming. In the summer of 1945 while exploring! 
Haleakala for the National Park Service, it was possible for the writer to enter 
this area. The upper section of Kipahulu Valley is well watered and densely 
forested; and the steep head walls are nearly always blanketed with clouds 



Vol. XXV] ST. JOHN : ENDEMISM IN HAWAIIAN FLORA. GUNNEEA 391 

and very wet. The stream heads have cut narrow gulches on the precipitous 
head walls and in these the Gunnera occurs from about 4,500 to 6,000 feet 
altitudes. It was found possible via Wai Anapanapa to approach one locality 
and get specimens. These plants prove to be conspecific with those of the wind- 
ward slope of Haleakala. 

4. Gunnera Eastwoodae St. John, sp. nov. 

(Plate 3 7, fig. f; plate 42) 

Herba gigantea 1.5 m. alta, stipulis hispidis, petiolis 45-60 cm. longis, laminis 38-53 em. 
longis cordato-reniformibus parve 9-lobatis chartaceis infra pallidioribus nervis reticulatis 
hirsutulis lobis 8-10 cm. longis rotundatis apice breve acuminate marginibus apiculato- 
denticulatis, pedunculis 15-20 cm. longia hirsutis, inflorescentiis 37-64 cm. longis lance- 
oloideis axi primario tomentoso ramulis spicatis, floribus rubris, sepalis 1-1.2 mm. longis 
oblongo-lanceolatis ad apicem laceratis, petalis 0.8-1 mm. longis obovatis cucullatis 
laceratis, antheris 1.5-1.7 mm. longis ovalibus, drupis 3.5-4 mm. diametro luteis, pyrena 
1.6-1.8 mm. longa 1.4-1.5 mm. lata 1.1-1.2 mm. crassa late ovali-obovoidea tetragona 
straminea. 

Giant herb, 1.5 m. tall; stem stout and fleshy; stipules of seedlings 1.5-2.5 
cm. long, oval, erose, membranous, strongly nerved, sparsely hispid on the 
back, those of mature leaves apparently similar but narrower; petioles 45-60 
cm. long, sparsely hirsute; blades 38-53 cm. long, cordate-reniform, shallowly 
palmately 9-lobed, firm chartaceous, above dark green, remotely hirsutulous, 
the lobes 8-10 cm. long, semicircular, the end of the midrib and one or more 
prominent lateral veins ending in short acuminations, the margin irregularly 
apiculate denticulate, the sinus broad V-shaped, below the surface pale green, 
prominently reticulate veined, subappressed hirsutulous on all the veins and 
veinlets ; inflorescence probably axillary ; bracts at base of inflorescence not 
seen ; peduncle 15-20 cm. long, sparsely hirsute; inflorescence 37-64 cm. long, 
10-15 cm. in diameter, lanceoloid, the axis ferrugineous tomentose, bractless, 
the lateral branches ascending, spiciform, tomentose towards base, glabrous 
towards tip ; the principal lower bracts 15-20 mm. long, linear oblanceolate, 
the margin membranous ; the flowers perfect, bright reddish, sessile ; ovary 
ovoid truncate; calyx lobes 1-1.2 mm. long, oblong-lanceolate, the tip lacerate ; 
petals 0.8-1 mm. long, obovate, cucullate, sparsely lacerate; the 2 anthers 
1.5-1.7 mm. long, oval; the 2 stigmas 3 mm. long, filiform, densely puberulous 
throughout; drupes 3.5-4 mm. in diameter, globose, yellow; seeds 1.6-1.8 mm. 
long, 1.4r-1.5 mm. wide, 1.1-1.2 mm. thick, broadly oval-obovoid, the apex 
emarginate, 4-angled, but somewhat compressed, two angles equal, the third 
well developed upwards, the fourth weak, the surface dull, stramineous. 

Type: Hawaii, Waipio, Upper Hamakua Ditch Trail, 3,500 ft. alt., on 
moist cliff, Jan. 1, 1932, H. St. John and E. Y. Hosaka 11,478. B. P. Bishop 
Museum, Honolulu. 

Collections studied, all from the island of Hawaii : Waipio Valley, Upper Hamakua Ditch 
Trail, face of pali at head of valley, alt. 1,300 m., Nov. 19, 1926, L. H. MacDaniels 286 ; 
Kohala Mts., Waimea, Sept. 7, 1911, C. N. Forbes 488.H; Kohala, Oct. 1931, 0. Degener 



392 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Her. 

S,183 ; Alakahi, Kawainui or Kawainui ditch or Kawaiiiui Valley, 3n\y 12, 1909, J. F. Eoclc, 
129, and 4,374 or 4,375 or 4,376 or 4,382; Waimea, Hawaii, June 1910. J. F. Bock 8,432. 

The specimens with definite localities all came from the windward slope of 
the Kohala Mountains, a region of very wet, swampy forests, at the brink of 
the precipitous, deepl^y entrenched valleys. Waimea is the nearest town, with 
hotel and supply base. It is in open grasslands, not the sort of place where 
Gunnera grows. Doubtless Waimea was the base camp of the collectors, not 
the habitat and locality of the plant. Forest ranger Kawai reports another 
locality at the head of Honopue Gulch. 

Rock's collection in 1909 bears confusing data on the four sheets examined. 
Alakahi and Kawainui are distinct valleys, both tributary to Waipio. They 
diverge sharph% so it is not evident how the locality could be in both valleys. 
Each of his sheets bears two numbers. His number 129 is perhaps a trip num- 
ber, while the others are apparently serial numbers given to each sheet col- 
lected. Though bearing different numbers they are doubtless duplicate sheets. 
Degener's locality Kohala is also open to question. Kohala town, post office, 
and mill are in the lowland, remote from the area where Gunnera is known 
to grow. Perhaps this locality is to be taken as a general one, referring to the 
Kohala District or Mountains. 

This new species is dedicated to Miss Alice Eastwood, Curator of Botany 
in the California Academy of Sciences. Her many decades of creative work 
have enriched the science of botany, and her friendliness and generous assist- 
ance have endeared her to all botanists who have consulted her. 

5. Gunnera kaalaensis (Krajina) St. John, comb. nov. 

(Plate 37, fig. c ; plates 43, 44, 45 ) 
Gunnera petaloidea Gaudichaud var. Icaalaensis Krajina, Acta Botanica Bohemica 9:50, 
figs. F„ B„ Flo, P„ Ao, 1930. 

Giant herb ; stem 1-3 m. tall, about 15 cm. in diameter, decumbent at base, 
the surface marked with raised, pale, corky leaf -scars, and above them numer- 
ous oval, raised craters ; stipules 4-8 cm. long, lanceolate, scurfy on the back, 
membranous, strongly nerved ; petioles 3-6 dm. long, at first densely hirsute, 
later sparsely so, with scattered dark tubercles; blades 20-50 cm. long, cordate 
reniform, shall owly palmately 7-9-lobed, firm chartaceous, above dark green, 
hirsute on primary and secondary veins or glabrate, below paler green, 
rough hirsute on the prominent, reticulate veins and veinlets, the lobes low 
undulate, shallow, rounded, 3-7 cm. long, the sinus broad U-shaped, the 
margin coarsely salient dentate; inflorescence axillary; bracts at base of 
peduncle reduced, about 2.5 cm. long, membranous ; peduncle 7-12 cm. long, 
bractless, hirsute but soon glabrate ; inflorescence 15-47 cm. long, 4-14 cm. in 
diameter, narrowly lanceoloid, the axis hirsute or glabrate, principal lower 
bracts 10-20 mm. long, linear-lanceolate, the margin membranous and not or 
but partly fimbriate ciliate, the lateral branches ascending spiciform, hirsute 
at base, glabrous towards tip; flowers perfect, sessile, greenish; ovary de- 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNERA 393 

pressed globose; calyx lobes 0.6-0.8 mm. long, rounded, lacerate; petals 1.3- 
1.8 mm. long, spatulate-obovate, eucullate, entire, obtuse, glabrous; the 2 
anthers 1.1-1.3 mm. long, ovoid; the 2 stigmas about 2 mm. long, densely 
puberulous ; drupes 2.8-3.1 mm. in diameter, globose, yellow ; stone 1.8-1.9 
mm. long, 1.4-1.6 mm. wide, 1.3-1.5 mm. thick, broadly oval-obovoid, the apex 
rounded or emarginate, with 4 prominent angles, the 2 lateral strongest and 
equal, the 3rd nearly as strong but diminishing at base, the 4th nearly as 
strong as the 3rd in the upper half, in the lower half nearly wanting, the 
surface yellowish brown, dull. 

Type: "Oahu : Montes Waianae, in declivibus orientalibus sub summo 
montis Puu Kaala, 1150 m.s. m., legi 24. XI. 1929" [F. Krajina in Herb. 
Charles Univ., Prague, not seen J . 

Collections studied, all from Oahu, Waianae Mts., Mt. Kaala: Kaala Mts., H. Mann and 
W. T. Brigham 626; Mt. Kaala, Jan. 1920, E. Bryan; Mt. Kaala, elev. 3,200 ft., Feb. 11, 
1928, H. F. Bergman; Puu Kaala, near summit, Waianaeuka, alt. 3,900 ft., wet stream 
bank, Nov. 24, 1929, H. St. John 10,072 ; Mt. Kaala, on moist upper slope by spring, elev. 
1,100 ± m., April 21, 1929, E. H. Brijan Jr. 707 ; Mt. Kaala, Waianaeuka, 4,000 ft. elev., 
Feb. 2, 1930, E. Y. Hosalca 139; Scliofield trail to Kaala, steep wet slope, elev. 3,300 ft., 
April 24, 1932, E. P. Hume 547 ; Puu Kaala, 3,600-3,800 ft., Waianaeuka, Jan. 8, 1933, 
V. H. Krauss; Kaala, eastern slope, at summit near spring, Sept. 25, 1938, 0. Degener, E. 
Ordonez and 0. Selliiig 12,251. 

Koolau Eauge: Konahuanui, near summit, Oct. 19, 1930, J. R. Coleman; Puu Kona- 
liuanui, east face, n.e. of north peak, head of South Fork of Kahanaiki Stream, Kailua, 
2,950 ft. alt., thicket on steep pali, June 21, 1942, L. Euhanic and A. D. Conger 131. Puu 
Konahuanui, precipitous, small, hanging valley, east side of Koolau Divide, head of North 
Fork of Kahanaiki Stream, 2300 ft. alt., Kailua, May 9, 1943, H. St. John 20,335. 

The localities for this species are discussed in the introduction. 

Gunnera kaalaensis forms a narrow belt on the eastern side of Mt. Kaala. 
It is conspicuous and is collected by most botanists. The older collections, as 
that by Mann and Brigham in 1864—65 were probably made on the old trail 
that ascended from Waialua. The numerous recent collections are apparently 
all from the station on the trail from Schofield Barracks, above the Firebreak 
Trail in Waianaeuka. The collectors have stated the elevations all the way 
from 3,200 to 4,000 feet. Doubtless all of the collections on this trail were 
made at the base of the "pali" (precipice), or by the "spring" or stream 
above it at elevations between 3,500 and 3,700 ft. The plant does not seem 
to occur at the actual summit. On the earlier U. S. Government topographic 
map of 1917, the mountain was given the Hawaiian name, Puu Kaala, and an 
elevation of 4,030 ft. The recent one of 1938 calls the peak Mt. Kaala, and 
measures it as 4,025 ft. high. 

The collection from Puu Kaala, St. John 10,072, shoAvs on one sheet a young 
inflorescence which has the petals either entire or sparsely fimbriate. This 
fringing of the petals is not constant and has not been observed in any of the 
other numerous collections. 

Krajina (p. 50) in separating this as var. kaalaensis included among the 



394 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

characters that it had the blade shining above, while the blades of his var. 
mauiensis were not shining. In this present revision, this character is not sub- 
stantiated, no diagnostic difference in the shiningness of the blade being noted 
between these two or any others of the Hawaiian species of Gimnera. 

6. Gunnera makahaensis St. John, sp. no v. 

(Plate37, fig. b.) 

Herba gigantea, caule 1.5 m. alta 15 cm. diametro, petiolis ad 7 dm. longis, laminis 20-37 
cm. longis cordato-reniformibus arete 7-9-lobatis chartaceis molle hirsutis in nerviis vel 
glabratis infra pallidioribus molle adpressi-hirsutis in nerviis reticulatis lobis sinuatis 
dentieulatis mucronatis, pedunculo bracteoso, bracteis 15-20 mm. longis 4-5 mm. latis 
oblongo-spatulatis marginibus membranaceis bifimbriatis, pedunculo 5-7 cm. longo, in- 
florescentiis 15-60 cm. longis anguste lanceoloideis, axi tomentoso, bracteis inferioribus 
15-35 mm. longis, ramulis spicatis, floribus viridescentibus sepalis 1-1.2 mm. longis late 
deltoideis profunde laceratis, petalis 1.9-2.2 mm. longis obovatis cucullatis marginibus 
erosis vel laceratis saepe acuminatis, antheris 1.6-2.2 mm. longis ovalibus, drupis ignotis. 

Giant herb, 2 m. tall ; stem about 1.5 m. tall and 15 cm. in diameter ; stipules 
not seen; petioles about 7 dm. long, somewhat hirsute, soon glabrate, sparsely 
tuberculate; blades 20-37 cm. long, cordate-reniform, shallowly palmately 
7-9-lobed, firm chartaceous, above dark green, sparsely soft hirsute on the 
veins or glabrate, below pale green, appressed soft hirsute on prominent veins 
and reticulate veinlets, the lobes 1.5-4 cm. long, shallowly sinuate semi- 
orbicular, the sinus broad and open U-shaped, the margin minutely salient 
denticulate and the principal dentations revolute mucronate ; inflorescence 
probably axillary; bracts at base of peduncle 5-6 cm. long, oblanceolate, 
scurfy towards the tip and margin ; peduncle when young clothed with numer- 
ous imbricate bracts which are 15-20 mm. long, 4-5 mm. wide, oblong spatu- 
late, firm, glabrous, with a conspicuous broad membranous margin lacerate 
and ciliate fimbriate, the peduncle 5-7 cm. long ; inflorescence 15-60 cm. long, 
6-10 cm. in diameter, narrowly lanceoloid, the axis brown tomentose, the prin- 
cipal lower bracts 15-35 mm. long, oblong-linear, acute, firm, the conspicuous 
membranous margin lacerate and ciliate fimbriate, the lateral branches ascend- 
ing, densely spicif orm, tomentose except towards the glabrate tip ; flowers 
perfect, sessile, greenish; ovary depressed globose; calj^x lobes 1-1.2 mm. 
long, broadly deltoid, deeply lacerate; petals 1.9-2.2 mm. long, obovate cucul- 
late, the margin erose to lacerate, often acuminate ; the 2 anthers 1.6-2.2 mm. 
long, oval; the 2 stigmas 1.5-2 mm. long, filiform, densely puberulous ; drupes 
unknown. 

Type: Oahu, Pun Kawiwi-Mt. Kaala ridge, Makaha-Waianae Kai, Wai- 
anae Mts., moist thicket in gulch, above pali [precipice], 3,500 ft. alt., 
March 31, 1935, H. St. John 17,581. 

Collections studied: Oahu: Kaala Eange, Makaha V[alley], Feb. 12-19, 1909, C. N. 
Forbes; Mt. Kaala, Waianae Mts., on wet wooded slope, el. 4,000 ft., (specimen brought 
by other students probably from the Makaha side), Feb. 2, 1930, E. Y. HosaTca. 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNEEA 395 

The specimen preserved by Hosaka was given him by other students who 
pushed ahead and crossed the summit to the Makaha side. Hence, his general- 
ized data and elevation of 4,000 feet are not to be considered accurate. 

It is remarkable that there are two species on Mt. Kaala, G. kaalaensis being 
abundant on the wet windward slope, 300 to 500 feet below the summit. G. 
makaJiaensis is infrequent at about the same altitude on the drier leeward 
slope. Actually, the broad, flat summit is well watered and supports a swampy 
forest with a few boggy openings. Gunnera has not been found on the summit. 
The locality of this new species is along one of the streams draining from the 
wet summit down towards the dry IMakaha Valley. 

The islands Kauai, Molokai, and Hawaii each produce a single species. Maui 
bears two species, one on each of the isolated mountain masses of east Maui and 
w^est Maui. Oahu has one species in both mountain ranges with a considerable 
gap between the localities on Puu Konahuanui in the Koolau Range and Mt. 
Kaala in the Waianae Mts. It is known only on the eastern slopes of Mt. Kaala. 
Yet, on the drier, westward slope we find a second species. It is amply distinct 
from its close neighbor in characters of sepals, petals, anthers, bracts, leaf 
blade size, lobing, and toothing. It is the only Hawaiian species with the mar- 
ginal teeth of the blade bearing definite revolute cartilaginous mucros. It has 
seldom been collected because of its scarcity and the long difficult ascent to 
the locality. 

7. Gunnera kauaiensis Rock 

(Plate 37, fig. a.) 
Gunnera Tcauaienis Eock, Oee. Pap. Bishop Mus. 9(5) :3-4, pi. 1, 1930, Sept. 
G. Dominii Krajina, Acta Botanica Bohemica 9 : 50, 52, figs. Fj, Bi, Ii, Pli, Pj, Ai, 1930. 

Giant herb, 1-3 m. tall ; stem 1-1.3 m. tall, 10-15 cm. or more in thickness, 
fleshy, decumbent at base ; stipules 4-5 cm. long, obovate, erose, pilose towards 
the margin, membranous, strongly nerved; petioles 3-6 dm. to "more than 1 
meter" in length, at first brownish tomentose but soon glabrate except at base, 
purplish, with scattered pustules; blades 0.8-2 m. in diameter, suborbicular 
peltate about % way from one side, firm chartaceous, above dark green, 
sparsely hirsute to subglabrate, below pale green, hirsute on the raised reticu- 
late veins and veinlets, the margin palmately 9-11-lobed, the lobes semior- 
bicular, acute, 5-10 cm. long, with a broad U-shaped sinus, the margin coarsely 
doubly dentate, the teeth 2-4 mm. long ; inflorescence probably' axillary ; basal 
bracts not seen ; peduncle 6-10 or more cm. in length, brownish hirsute ; in- 
florescence 3-6 dm. long, ("1 meter or more long," fide Rock), 8-17 cm. in 
diameter, dense, lanceoloid, the axis densely brownish hirsute, at length sub- 
glabrate, lower primary bracts 15-18 mm. long, oblong-lanceolate to spatulate, 
glabrous, the margin scarious, the lateral branches ascending, spiciform, 
hirsute at base, glabrate at tip; flowers perfect; ovary depressed globose; 
calyx lobes 0.3-1 mm. long, deltoid, the margin thin, lacerate; petals 1.1-1.3 
mm. long, obovate cucullate, the margin ciliate or lacerate; the 2 anthers 
1.9-2.1 mm. long, oval ; the 2 stigmas 1.5-2 mm. long, filiform, densely puber- 



396 CALIFOBNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

iilous ; drupes 3^ mm. in diameter, globose, red to purplish ; stone 1.9 mm. 
long:, 1.6 mm. wide, 1.4 mm. thick, oval, somewhat compressed, the apex emar- 
ginate, conspicuously 3-angled, the 2 lateral angles equal, acute, the 3rd angle 
prominent and acute in the upper half, wanting below, the 4th angle unde- 
veloped, the surface stramineous, dull. 

Type: "Kauai, along the stream beds of Kaluiti and Kailiili, just below the 
summit of Waialeale, elevation 4800 to 5000 feet, September 22, 1909 (fruit- 
ing), J. F. Kock no. 5053. Type B. P. Bisliop Mus." On one or more of the 
five isotypic sheets are found the numbers 5,050, 5,053, and 5,058. These are 
apparently individual numbers given by the collector to each sheet. Later he 
selected the number 5,053 to stand for the collection. They all bear also the 
number 129 which perhaps may be a trip number, though it duplicates the 
one for the collection from Hawaii in the same year. Rock's three series of num- 
bers on his collections are not well understood by the botanists of Honolulu. 

Collections studied, all from Kauai: Kahili summit, Aug. 1909, C. N. Forbes; Kilohana 
Outlook above Wainiha, Waimea Drainage Basin, West Side, July 3 to Aug. 18, 1917, 
C. N. Forbes 903.K; Kilohana lookout, N. W. end Alakai Swamp, Na Pali-Kona forest 
reserve, wet steep slope, el. 4,023 ft., Dec. 27, 1930, E. St. John et al, 10,788; Avithout locality, 
Oct. 1916, J. F. Rock. 

Rock (p. 3) described the "Panicle terminal." Neither his specimens nor 
any others of those available in Honolulu show an attached inflorescence. 
All of the other Haw^aiian species have their inflorescences developing near 
the stem tip, but truly axillary, and it seems probable that this one does too. 
The inflorescence is not a true panicle, rather it has an erect common axis 
bearing numerous ascending, unbranched spikes. 

This is the most distinct of the Hawaiian species, differing markedly from 
all the others. It alone has orbicular, peltate blades and drupes that are red to 
purplish. It also differs in the size and structure of the flowers and fruits. 

As indicated in the introduction, it is still an open question as to which of 
the two specific names of this species is the earlier. G. kauaiensis Rock was 
published in September 1930. G. Dominii Krajina was published in the same 
year, 1930 (Acta Botanica Bohemica 9 :50, 52, figs.. 1930). Previous requests 
for the exact date of publication, sent to Prague have been unanswered. Now 
we acknowledge with thanks the assistance of the librarians of the Gray Her- 
barium, the British Museum of Natural History, and the Director of the Kew 
Gardens. It is not known whether or not the several articles in volume 9, Acta 
Botanica Bohemica w^ere issued separately at various dates, or not. In any 
case, volume 9 was commonly distributed as a single bound volume, and the 
copies were received by the three institutions listed. Gray Herbarium on Oct. 
25, 1932, Kew on March 6, 1933, and British Museum on April 1, 1936, but 
these dates of receipt are so late as to be of no help. Krajina lists as G. peta- 
loidea var. mauiensis his own specimen collected on Maui on April 15, 1930. 
His manuscript must have been writteu later than April, and in all proba- 
bility after he returued to Prague. From this it can be deduced onlv that his 



Vol. XXV] ST. JOHN: ENDEMISM IN HAWAIIAN FLORA. GUNNERA 397 

paper was not published early in 1930. The first paper in vol. 9 was by the 
editor, Karel Domin, "A new species of Psitt acanthus from Dominica," which 
paper was not listed in the Botanisches Centralblatt until July 6, 1933. The 
succeeding volume 10 of Acta Botanica Bohemica contained a chronological 
bibliography of Dr. Karel Domin's publications. Two of them were published 
in volume 9, and they are listed in the bibliography as appearing in 1930. In 
one of these papers, page 256, Domin cites a plant specimen collected on Sept. 
20, 1930, which would imply that his paper was printed late in the year 1930. 
Volume 12 (1934) of the Prague journal Preslia is devoted almost entirely to 
bibliographic notes, and it lists Krajina's Gunyiera paper as published in 
1930, stating only the year. It is not known that this vol. 9 was issued only in 
complete volume form, and the exact date of Krajina's publication is still 
unknown. Since Rock's publication was dated Sept. 1930, it is tentatively 
accepted as the prior publication. 



PLATES 



PLATE 37 

Margin of terminal and one lateral lobe of blade of the Ha- 
waiian species of Gunner a. a, G. kauaiensis; b, G. malcahaensis ; 
c, G. Tcaalaensis; d, G. petaloidea ; e, G. moloTcaiensis ; f, G. East- 
woodae ; g, inauiensis. 



[ 4nn 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV NO. 16 [ST. JOHN] PLATE 37 




[ ^01 ] 



PLATE 38 

Gunnera petalo'idea Gaudichaud, type in Herb. Museum d'His- 
toire Xaturelle, Paris. 



[402] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 16 [ST. JOHN] PLATE 38 




Photo by H. St. John. 



[403] 



PLATE 39 

Gunnera petaloMea Gaud., at head of Kahana Valley, 3,800 ft. 
alt., west Maui, October, 1942. 



[404] 



01 
(0 
LiJ 
1- 
< 

a. 
I — I 
z 
I 




o 

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> 

X 
X 





> 

111 

tr 

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u 

u 
o 

0. 




PLATE 40 

Gunnera mauiensis (Krajina) St. John, Kula Pipe Line, bank 
of Waikamoi Stream, 4,250 ft. alt., east Maui, with figure of 

Koyal N. Chapman. 



[406 J 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 16 



[ST. JOHN] PLATE 40 




[407] 



PLATE 41 

Gunnera mauiensis (Krajina) St. John, Kula Pipe Line, bank of 
Waikamoi Stream, 4,250 ft. alt., east Maui, with figure of "Worth 

Aiken, 1925. 



[408 ] 



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h 
< 

_l 

Q. 

|—i 

Z 
I 

o 



I- 



o 

z 

> 

X 

J 


> 

hi 

ir 

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(fl 

I 

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0) 

d 

< 

< 



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u 
o 

(E 




PLATE 42 

Gunnera Eastwoodae St. John, tj-pe specimen. Upper Hamakua 
Ditch Trail, 3,500 ft. alt., Waipio, HaAvaii, January 1, 1932. 



[ 410 ] 



'J 
llJ 

H 
< 
J 
D. 
r— 1 
Z 
I 

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PLATE 43 

Gunnera kaalaensis (Krajina) St. John, Mt. Kaala, "Waiana- 

euka, 3,500 ft. alt., Oahu, with from left to right, figures (in 

upper right) of H. Pope and W. B. Storey. 



[412] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 16 



[ST. JOHN] PLATE 43 




Photo by Willis T. Pope. 



[413] 



PLATE 44 

Gunnera Tcaalaensis (Krajina) St. John, Mt. Kaala, Waiana- 

euka, 3,500 ft. alt., Oahu, with, from left to right, figures (in 

foreground) of H. Pope and W. B. Storey. 



[414] 



< 

J 
D- 
I — I 
Z 
I 

—> 

t- 



o 

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> 

X 
X 

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ui 

(E 
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u 

u 
o 

q: 
a. 




PLATE 45 

Gunnera Tcaalaensis (Krajina) St. John, Mt. Kaala, Waiana- 
euka, 3,500 ft. alt., Oahu, 



[416] 



PROC. CALIF. ACAD. SCI.. 4TH SERIES. VOL. XXV. NO. 16 



[ST. JOHN] PLATE 45 




Photo by Willis T. Pope. 



[417] 



f PLATE 46 

Distrilnition of Gunnera in the Hawaiian Islands. 
Square: Gunnera mololcaiensis ; Erect Triangle: G. petaloidea; 
Inverted Triangle: G. mauicnsis ; Shield: G. Eastivoodae ; Cir- 
cle: G. kauaiensis ; J: G. makaliaensis ; L: G. kaalacntiis. 



[418] 



10 

t 

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0. 

1— 1 

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Marine Biological Labordtory 

Xj X IB K. J^ T^ -X" 



JAN 3 1 1947 

WOODS HOLE, MASS. 



PROCEEDINGS 

OF THE 

CALIFORNIA ACADEMY OF SCIENCES 

Fourth Series 
Vol. XXV, No. 17, pp. 421-468, ig text figs. November 15, 1946 



ALICE EASTWOOD 
SEMI-CENTENNIAL PUBLICATIONS* 

No. 17 

A REVISION OF THE ARTEMISIA VULGARIS 

COMPLEX IN NORTH AMERICAf 

BY 

DAVID D. KECK 

Carnegie Institution of Washington 

Division oj Plant Biology 

Stanford University, California 

THE NORTH AMERICAN ALLIES of Artemisia vulgaris L, an Old AVorld species, 
form a polyploid complex of great diversity, extending from Alaska to 
Guatemala, and from Hudson Bay and the Mississippi Valley to the Pacific. 
The center of this diversity lies in the western United States. The extent of 
the problem posed to the systematist by this group of plants is at once appar- 
ent by referring to the last two complete treatments of these species. Hall and 
Clements (1923) included under Artemisia vulgaris L., with fifteen subspecies, 
all the American forms which, but fifteen years previously, had been treated 
as fifty -four distinct species by Rydberg (1916). 

These authors were alike in using comparative morphology as the principal 
tool in their taxonomic work, but their species concepts were at different 
poles from each other. Rydberg had come to believe that any striking morpho- 
logical form deserved a name, and that this should be a specific name, since he 
could see no advantage in using trinomials. The significance of geographical 
range was rarely considered. Rydberg's species often represent morpho- 
logically well-marked forms, but all too frequently they are found to be thor- 
oughly linked with others by intermediates, or to be unique variants, or to 
have no natural range of their own. Often his species are based on one morpho- 
logical character, which is soon found to occur sporadically here and there 

* Printed from the John W. Hendrie Publication Endowment, 
t Keceived for publication Feb. 6, 1943. 

[421 ] 



422 CALIFOENIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

over a large area in which other characters in the same plants show trends of 
variation in other directions. These limitations in Rydberg's species concept 
make his entities artificial rather than natnral. 

In their monumental monograph, Hall and Clements, at the other extreme, 
offered the most conservative species concept that could be defended on 
morphological grounds. The result was the telescoping of Rydberg's fift}"- 
four species of this complex into one species, as mentioned above. Later 
evaluation of the same materials by other workers has shown that the funda- 
mental units of Hall and Clements were carefully established. Other work- 
ers have upheld the majority of their basic units but have often treated 
their subspecies as species in order to conform with more widely accepted 
taxonomic practice. Hall and Clements' work is therefore sound from the 
morphological viewpoint, but the addition of one significant new criterion, 
chromosome number, has now made it possible to see a little more clearly the 
limits of the species in this group. 

The present paper takes the consequences of this discovery and presents a 
study of many new collections that were not available to Hall and Clements. 
It also utilizes genetical, garden, and field studies as background data for 
the herbarium studies. Several members of this complex were grown in the 
varied environment experiments conducted by the Carnegie Institution of 
Washington along a transect across central California, and some cytological 
and genetical work was done on them. The results of these experimental 
studies have been reported by Clausen, Keck, and Hiesey (1940a). Addi- 
tional forms have since become available for cytological study. The herbarium 
investigation was made at the time when the results from the transplant 
experiments were being prepared for publication, because the cytogenetic 
findings showed the need for a revised sj'Stematic treatment. 

The investigations of Clausen, Keek, and Hiesey (1940a and b) showed 
that several ecospecies must be recognized in this polyploid complex. On the 
other hand, from a limited amount of evidence these authors suggested the 
possibility that, to use the biosystematic term, only one cenospecies was in- 
volved in the whole subsection Vulgares of Artemisia, i.e., that at least a 
limited amount of interbreeding and gene exchange could still take place 
among its members. 

The morphological variation of the plants composing this complex is very 
great, making the delimitations of natural groups a difficult problem. The 
prominent characters of taxonomic use include size of parts such as leaves and 
heads, shape of leaves, and quality and quantity of the pubescence. IMost , 
of these are subject to environmental modification and must be interpreted 
with that in mind. Vegetative propagation by means of rhizomes is well de- 
veloped in most of the species, and notable variations may be perpetuated for 
long periods in this manner. Under adverse climatic conditions, as in the far 
north and at alpine heights, it may be only the exceptional year in which these 
plants are able to propagate themselves sexually, although they are strongly 



Vol. XXV] KECK: EEVISION OF ARTEMISIA VULGARIS COMPLEX 423 

established in these localities (Hulten, 1937, p. 327; Clausen, Keck, and 
Hiesey, 1940a, p. 336). Furthermore, single characters, which some authors 
have utilized in the past to differentiate "species" in this group, have been 
found very undependable for this purpose. It is only when a group of char- 
acters is found correlated to each other within a given territory that the 
existence of a natural S3'stematic unit can be surmised. Even then the sys- 
tematist must reconcile himself to having a residue of specimens that he must 
arbitrarily include or exclude from a given unit of lower order, because nat- 
ural crossings between such units produce recombinations that do not fit 
his key. 

When historical precedent has entrenched a name in the botanical manuals, 
only a thorough revision of the group can properly appraise it. For example, 
Thomas Nuttall in 1818 proposed four species in this group. One, based on a 
specimen from near St. Louis, with leaves lightly tomentose on the upper sur- 
face and densely tomentose beneath, he named A. ludoviciana. A second, based 
on specimens from the Wisconsin area, with leaves white-tomentose on both 
sides, he named A. gnaphalodes. These two species have been maintained in 
the majority of botanical works down to the present, yet this writer agrees 
with Asa Gray that there is no warrant for separating these variations even 
in a subspecific category. The only characters that have been proposed for dis- 
tinguishing them prove to be completely scattered in distribution, wdth every 
indication of complete interbreeding between the forms. 

This revision employs orthodox taxonomy rather than experimental tax- 
onomy (biosystematics), because the experimental evidence has been very 
incomplete. Consequently, it has not been possible to determine the exact 
genetic-ecologic boundaries of the various units, and thus to apply more than 
superficially the concept of species set down by Clausen, Keek, and Hiesey 
(1939a). Consequently, most reliance has necessarily been placed upon com- 
parative morphology coupled with distribution; and all of the available 
cytological information has been used. In order to recognize units as com- 
parable as possible with those determined by experiment, the geographic- 
ecologic principle has been followed in developing this species concept. That 
is, an attempt has been made to find key morphological distinctions that mark 
groups limited by natural environmental barriers, whether these barriers 
be geographical or ecological. From the kind and amount of intergradation 
observed between the natural units thus set off, the systematic category of 
the units has been determined. 

Under the biosystematic concept of species at least partial genetic barriers 
exist between species. A difference in chromosome number is a genetic barrier 
to free intercrossing, and so it in itself may be taken as evidence that distinct 
species are involved, provided that morphological differences have had time 
to follow the genetic break. In the present group of species the differences in 
chromosome number coincide with major morphological breaks in each in- 
stance examined. 



424 ' CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

In the vulgaris complex the morphological breaks between species are only 
moderately distinct. This prompted Hall and Clements to keep all its units in 
one species. As compared with almost all the other species of the section Ahro- 
tanum to which this group belongs, the specific differences in the subsection 
Vulgares are not morphologically large. A gauge for species is set, however, 
by the chromosome data where they are known. With these as a basis, the othet 
species have been delimited as regional groups of comparable order, showing 
relatively little intergradation. 

Regional groups that differ from each other in morphology and distribu- 
tion, but that intergrade completely where their ranges overlap, appear to 
lack genetic barriers. Such units are treated as subspecies, and all of these are 
geographical. Because of the extensive morphological variation in these plants, 
requiring broad subspecies, it is likely that the larger subspecies contain more 
than one ecotype. 

Morphological variants that from herbarium evidence do not appear to 
have either geographic-ecologic correlation or genetic barriers separating 
them are not named. These often striking variants are the formae of many 
taxonomists; but such uncorrelated forms cannot be considered of phylo- 
genetic significance. 

Cytological and Genetical Evidence. The chromosome numbers of 37 plants 
from eight American systematic units, including seven species, have thus far 
been determined in addition to the European A. vtdgaris. These chromosome 
numbers have been reported previously by Clausen, Keck, and Hiesey (1939b, 
1940a and b) , but several additional collections are reported in this paper for 
the first time. The enumeration of the collections that have been counted fol- 
lows the citation of specimens of the unit concerned. 

The European A. vulgaris is a diploid species, with n = 8 chromosomes. In 
contrast, all the American forms of the Vulgares that have thus far been in- 
vestigated follow a 9 series of chromosome numbers. This cj^tological difference 
alone is sufficient to keep all the American forms out of the Old World vidgaris. 
Artemisia Carruthii and A. Suksdorfii are diploid, with n = 9 ; A. serrata, A. 
longifolia, and A. ludoviciana sspp. typica and incompta are tetraploid, with 
w = 18 ; and A. Douglasiana and A. Tilesii ssp. unalaschcensis are hexaploid, 
with n = 27. Chromosomes of these forms are illustrated in figures 1-9. The 
two American diploid species thus far known, A. Carruthii and A. S^iksdorfii, 
have larger chromosomes than any of the other forms including the diploid 
A. vtdgaris, with the chromosomes of A. Suksdorfii the largest of all. 

The little breeding work that has been done with these plants has for the 
most part been reported (Clausen, Keck, and Hiesey, 1940a). This report 
covered selfings of A. Douglasiana from Santa Barbara, and of A. ludovi- 
ciana typica from Manitou, Colorado, and also two pentaploid hybrids that 
were obtained when these two races were crossed. An additional cross has 
been attempted between A. ludoviciana typica from Leevining, California, 
and A. Suksdorfii from Trinidad, California, in the hope that A. Douglasiana 



Vol. XXV] KECK: BEVISION OF ARTEMISIA VULGARIS COMPLEX 



425 




Figures 1-9. Heterotypic metaphases of meiosis in pollen mother cells and somatic meta- 
phases in roots. Fig. 1. Artemisia vulgaris, Skane, Sweden, Sn. Fig. 2. A. Carruiliii, Springer, 
New Mexico, 2n = 18. Fig. 3. A. Sulcsdorfii, White Horse, Washington, On. Fig. 4. A. serrata, 
St. Paul, Minnesota, 2n = 36. Fig. 5. A. longifolia, Billings County, North Dakota, 2w = 36. 
Fig. 6. A. Douglasiana, Medford, Oregon, 2n = 56. Fig. 7. A. ludoviciana typica, Manitou, 
Colorado, 16ii + liv Fig. 8. A. ludoviciana incompta, Yosemite Creek, California, 18„. Fig.9. 
A. Tilesii unalaschcensis, Unalaska, Aleutian Islands, 27„. Enlargement for all figures: ca. x 
2400. 



426 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

might be synthesized through amphidiploidy, but it did not have a fair chance, 
because the races tested differed too much in time of flowering. The supposed 
amphidiploid origin of A. Douglasiana is discussed in more detail on page 459. 

The Subsection in Perspective. The members of the vulgaris complex are 
widely distributed across the North American continent. There are two 
diploid species of radically different appearance. Along the Pacific shores of 
the United States from the Canadian boundary to northern California gro^vs 
Artemisia Suksdorfii, a stout, large-leaved herb in clumps often more than a 
meter high and half again as broad. On the more arid plains and plateaus from 
western Kansas and Colorado to Chihuahua is A. Carruthii, a low gray clump 
with tiny fascicled leaves, in marked contrast with the preceding diploid. 

Through the midlands of the continent there is a group of tetraploids per- 
plexing in their variation. On the periphery it is relatively simple to disjoin 
A. serrata and A. longifolia from the mass, the former in the upper Mississippi 
Valley, and the latter spreading somewhat westward from the prairies and 
badlands of the Dakotas. The remainder goes through an impressive amount 
of variation across the western half of the United States and down the length 
of Mexico, sometimes in accord with the climatic zone occupied, and again in 
seeming contradiction to the environment. From the lowlands and plains this 
distribution may be traced up through the timbered slopes to the alpine sum- 
mits, and from the regions of abundant rainfall out across the arid desert 
ranges. No lines of demarcation of even relative sharpness are at first apparent, 
for even the regional facies of the assemblage show rather frequent intergra- 
dation. As a consequence, this tetraploid diversity is gathered under one spe- 
cific name, A. ludoviciana. The regional races of individual caste within it 
are demarked as subspecies — seven in all. Where the sampling has been dense, 
as in Colorado, the abundance of examples makes the broad gamut of variation 
appear but the more confusing; however, from this it becomes apparent that 
the number of specific barriers to interbreeding in this assemblage is strictly 
limited — that the variation is intra-, rather than inter-specific. In Mexico, on 
the other hand, additional sampling may well uncover regional subspecies too 
poorly discerned for recognition at present. The potential variation is cer- 
tainly there. The increased sampling available to us now has resulted in the 
recognition of the validity of sulcata and albula as subspecies of ludoviciana 
from the arid Southwest, while the same abundance of material has failed to 
develop any substantial basis for separating gnaphalodes from ludoviciana, 
or Wright a from Carruthii, all of these results running at variance with 
rather recent conclusions drawn from less material. 

The two known hexaploids are A. Douglasiana, from a Pacific Coast region 
between A. Suksdorfii and A. ludoviciana, respectively its diploid and tetra- 
ploid presumed progenitors, its individuals becomiug the largest in all the 
complex; and A. Tilesii, a coarse herb with two subspecies distributed from 
the Pacific Northwest to the Arctic Ocean and to the western shore of Hudson 
Bay, but obviously coming to our continent from Asia, for the trail is clear 



Vol. XXV] KECK: EEVISION OF ARTEMISIA VULGARIS COMPLEX 427 

both across Bering" Strait and the Aleutians to Kamchatka, and westward to 
European Arctic Russia. Its near-relatives remained in the Old World. 

The chromosome numbers of A. Michauxiana and A. Lindleyana are not 
known as yet, and that information is desirable for the completion of the 
picture. The alpine A. Michauxiana is a quite well-marked entity with its 
small, much dissected and finely cut leaves, its large heads and low stature, 
and its near-restriction to the northern Rockies. Artemisia Lindleyana is a 
suffrutescent plant — -a distinction from the usual herbaceous forms — with a 
distinctive subracemose inflorescence but somewhat variable leaves. Its dis- 
tribution is a natural one along the low elevation watercourses of the Columbia 
River system. 

The least well-known member of the complex is A. Prescottiana, an anoma- 
lous species of airy aspect due to its long filiform divided leaves, sparse raceme, 
and very slender branching stem. The root is gone from the only specimen I 
have studied, but it might even have been annual. David Douglas collected it 
six score j^ears ago on the Quick Sand River, near The Dalles, but it seems to 
have escaped recollection, and it appears to be uncertain today even as to 
along which stream Douglas found it. 

Other conclusions on the derivations of the various systematic units that 
can be gathered from the several lines of evidence available are presented in 
the discussion following- the systematic treatment (pp. 465-467). 

Methods, Materials, and Acknowledgments 

From the early stages of the investigation it was apparent that a study of 
this complex in any one herbarium would be very inadequate because of the 
great variability found in nature and the broad geographic distribution, and 
that it would be essential to bring together an abundance of herbarium mate- 
rial for comparison. Consequently, the material from twelve herbaria was 
assembled for simultaneous comparison at our central laboratory at Stanford. 
In the case of the more widely distributed collections, this made possible the 
comparison of all the material from one locality at one time. On account of 
the variation within one habitat, this is a much safer method than comparing 
the materials in one herbarium at a time. 

So many herbarium specimens have been studied that their citation in 
entirety becomes impractical. Distribution maps therefore indicate wherever 
possible the approximate location of each collection, leaving for the citation 
lists only a limited number of collections. These have been chosen because they 
are reasonably representative and widely distributed to herbaria, or because 
they mark the range or otherwise have special interest. Citation is more com- 
plete for those groups that are rare, or that were previously understood 
inadequately. The complete documentation is to be found in those herbaria 
in which material was studied and annotated. In so far as possible citation is 
by counties in a north to south and east to west geographical sequence ; the 
county name is followed by a colon. 



428 • CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

The descriptions are cross-comparable and made a.s brief as permissible if 
they are to have a utilitarian value in a group marked by so great diversity. 
The distribution and ecological data are gathered from the herbarium speci- 
mens and from some personal field knowledge of the group. 

During the taxonomic investigation collections were studied and annotated 
in the following herbaria. The abbreviations are those emploj-ed in the cita- 
tion of specimens. In those herbaria marked with an asterisk, only selected 
groups were examined : 

C — University of California, Berkeley. 
CAS — California Academy of Sciences, San Francisco. 
CI — Carnegie Institution of Washington, Stanford University, Calif. 
*Minn — University of Minnesota, Minneapolis. 
NY — New York Botanical Garden, New York. 
Ph — Academy of Natural Sciences of Philadelphia. 
Po — Pomona College, Claremont, Calif. 

RM — Eocky Mountain Herbarium, University of Wyoming, Laramie. 
SU — Dudley Herbarium, Stanford University, Calif. 

US — United States National Herbarium, Smithsonian Institution, Washington, D. C. 
*UW — University of Washington, Seattle. 

Veg — Vegetation Type Map Herbarium of U. S. Forest Service, University of California, 
Berkeley. 

To the curators of these herbaria grateful acknowledment is made for the 
privilege of study generously afforded and for other help given. I am indebted 
to the late Sir Arthur W. Hill, Director of the Royal Botanic Gardens at Kew, 
for the loan of a few types and pertinent specimens from that institution. Par- 
ticular thanks are due my colleague, Dr. Jens Clausen, for critical aid in 
making taxonomic and cytologic judgments. To him also, together with my 
colleague. Dr. W. M. Hiesey, I am grateful for a critical reading of the 
manuscript. 

Systematic Treatment 

The great diversity of this group of plants has been mentioned. One of the 
most variable organs, which at the same time furnishes key characters of im- 
portance, is the leaf. In figure 10 some of this variation is shown in order to 
enable the reader to visualize the type of leaf-cut which words often inade- 
quately describe. One is not to assume that this illustration encompasses by 
any means the entire range of variation that is to be encountered in these 
systematic units, but these drawings are intended to supplement the key. 

Key to the Systematic Units 

A. Divisions of lower leaves again lobed or cleft, 5-15 mm. wide, directed 
forward, green and essentially glabrous above. Old World ; eastern North 

America 1. A. vulgaris 

AA. Divisions of lower leaves entire, or if lobed, the divisions less than 5 mm. 
A\dde, spreading, or leaves themselves entire. 
B. At least some of the leaves bipinnately parted with salient segments 

again toothed. Northern Rocky Mts 2. A. Michauxiana 



Vol. XXV] KECK: REVISION OF ARTEMISIA VULGARIS COMPLEX 429 




n = 9 

SUKSDORFII 



rigure 10. Typical leaf outlines in Artemisia subsection Vulgares. x %. 



430 CALIFORNIA ACADEMY OF SCIEXCES [Pko^^- 4th 8er. 

BB. Leaves entire to bipinnatifid with entire lobes. 

C. Leaves linear to linear-lanceolate, entire, or rarely divided into 
few usually caudate lobes, often caudate and revolute-niargined, 
the tomentum on the upper surface usually deciduous, 3-12 cm. 
long; involucre 4-5 mm. high. North Dakota to central Wy- 
oming 3. A. longifolia 

CC. Leaves not as above, or if slender and entire the involucre shorter. 

D. Leaves finely and evenly serrate, strongly discolored; plant 

1-2.5 m. tall; heads only 2 mm. wide. Upper Mississippi 

Valley 4. J. serrata 

DD. Leaves not finely and evenly serrate. 

E. Leaves fascicled, 1-3 cm. long only, divided into di- 
varicate linear-filiform lobes 0.5-1 mm. wide ; panicle 
narrow, spike-like; herbage mostly gray-tomentose 
(leaves sometimes green on upper surface), the heads 
tomentulose and ± shining. Kansas and Arizona to 

northern Mexico 5. A. Carruthii 

EE. Leaves not fascicled, or if fascicled their divisions more 
more than 2 mm. wide or l>right green. 
F. Involucre as high as or higher than wide (except in 
^ A. ludoviciana candicans, but in that the leaves not 

prominently discolored). 
G. Principal leaves narrow, 1 cm. or less wide 
exclusive of lobes when present, tomentose on 
both sides or green above; stems rarely more 
than 1 m. tall. 

H. Herbaceous to base 6. A. ludoviciana 

I. Principal leaves entire or merely 
lobed. 

J. Leaves 3-10 cm. long, the mar- 
gin not revolute. 
K. Panicle usually compact, 
elongated, not leafy ; leaves 
not thin, usually white- 
tomentose on both sides, 
rarely bright green on up- 
per surface; involucre 3-4 
mm. high. Ontario and Mis- 
sissippi Valley to Washing- 
ton and California. 

6a. .4. ludoviciana ssp. typica 
KK. Panicle open, leafy (some- 
times contracted) ; leaves 
thin, usually bright green 
above, white-tomentose be- 
neath ; involucre 2.5—3 mm. 
high. Arizona to Cliihua- 

hua 6b. A. ludoviciana ssp. sulcata 

JJ. Leaves mostly 1-2 cm. long, 

lanceolate, thickish, usually 

• white-tomentose on both sides, 

the margin often narrowly rev- 



Vol. XXV] KECK: REVISION OF ABTEMISIA VULGARIS COMPLEX 431 

olute; panicle open, the heads 
pendant in slender zt leafy ra- 
cemes. Southwestern U. S. and 
northern Mexico. 

6c. A. ludoviciana ssp. albula 
II. Principal leaves ± parted or di- 
vided. 
L. Involucre 4-5 mm. high, 4-7 mm. 
vsride; leaves white-tomentose 
on both sides, plane. Western 
Montana to northern Utah and 
California. .6d. A. ludoviciana ssp. candicans 
LL. Involucre 2.5-3.5 mm. high, 2- 
3.5 mm. wide; leaves usually 
green on upper surface (some- 
times tomentose on both sides 
in incompta). 
M. Panicle spike-like or race- 
mose; disk-florets 15-45. 
Montane ; Montana to Utah 
and California. 

6e. A. ludoviciana ssp. incompta 
MM. Panicle broader, branch- 
ing, leafy; disk-florets 
6-18. 
N. Lower leaves 5-10 cm. 
long, pinnately or ter- 
nately divided, the 
lobes entire, strongly 
discolored. Southwest- 
ern U. S. to southern 
Mexico. 

6f. A. ludoviciana ssp. mexicana 
NN. Lower leaves 2—5 cm. 
long, bipinnately dis- 
sected, green and ± 
glandular, or mod- 
erately tomentose on 
under surface. S.W. 
Texas to S.E. Arizona 
and Durango. 

6g. A. ludoviciana ssp. reddens 
HH. Suffrutescent with woody base; leaves 
narrow, entire to serrate-dentate apically, 
rarely more deeply lobed; panicle nar- 
row, spike-like or racemose. Banks of 

Columbia Eiver and tributaries 7. A. Lindleyana 

GG. Principal leaves broader, 1-5 cm. wide exclu- 
sive of lobes when present, frequently entire, 
» lobes if present few, short, entire, discolored ; 

K stems frequently more than 1 m. tall. 



432 CALIFOBNIA ACADEMY OF SCIEXCES [Proc. 4th Ser. 

O. Involucre terete or narrow-ovoid, less 
than 2 mm. wide, yellow-green and shin- 
ing; ray-florets 3-7; disk-florets 2-8; 
leaves green above, silvery tomentose 
beneath. Immediate Pacific Coast and 

Puget Sound region 8. ^. SuTcsdorfii 

00. Involucre campanulate, 2-3 mm. wide, ± 
tomentose; ray-florets 6-10; disk-florets 
10-25; leaves less strikingly discolored. 
Western Washington to northern Baja 

California 9. A. Douglasiana 

FF. Involucre wider than high. 

P. Stem branching at base ; herbage very spar- 
ingly arachnoid-pilose, somewhat more dense- 
ly so (but not tomentose) on under surface 
of leaves; leaves pinnatifid with 3-7 filiform 
divisions less than 1 mm. wide ; inflorescence 

racemose. Columbia Eiver 10. A. Prescotfiana 

PP. Stem unbranched at base ; leaves tomentose 
on under surface, strongly discolored, deeply 
incised, the divisions wider; inflorescence a 

spike-like panicle 11. A. Tilesii 

Q. Inflorescence overtopped by the leaves ; 
involucre 4-5 mm. high, 6-9 mm. wide ; 
heads 30-125-flowered. Siberia to Hud- 
son Bay 11a. A. Tilesii ssp. fi/pica 

QQ. Inflorescence well exceeding the leaves ; 
involucre 3-5 mm. high, 4-8 mm. wide; 
heads 30-55-flowered. Northern Japan, 
Kamchatka, Alaska to Pacific Northwest. 

lib. A. Tilesii ssp. unalascJicensis 

1. Artemisia vulgaris L., Sp. PI. 848, 1753. 

Figures 1 and 10 

Type statement : "Habitat in Buropae ciiltis, ruderatis." 
Stems 5 to 15 dm. tall, slender to moderately stout, herbaceous to base, 
commonly unbranched; lower leaves obovate to deltoid-ovate in outline, 
deeply incised into 3-7 lance-ovate to narro\vl.y lanceolate principal divisions, 
which are for the most part again irregularly and sharply toothed or lobed, 
the lobes and teeth all directed forward, 5-12 cm. long, often nearly as wide, 
green and glabrous or nearly so above, densely white-tomentose beneath, the 
margin obscurely revolute ; inflorescence an ample leafy panicle with ascend- 
ing branches crowded with subsessile heads, 3-15 cm. broad ; heads medium- 
sized, erect or nodding; involucre campanulate, 3-4 mm. high. 2.5-3.5 mm. 
wide, moderately tomentose, with 9-13 bracts ; ray-florets 6-9 ; disk-florets 
13-20. 

Common throughout Europe and eastward to Siberia ; probably adventive 
in North America, but in places appearing indigenous, occurring irregularlj'' 
from Newfoundland to Manitoba and Alabama. 



Vol. XXV] KECK: BEFISION OF ARTEMISIA VULGAEIS COMPLEX 433 

Material of this species was examined in only three herbaria. In addition 
to the localities cited bv Hall and Clements (1923), collections should be 
reported from the states of New Hampshire, Vermont, Massachusetts, and 
North Carolina. 

The chromosomes were counted in meiosis of plants grown from seed col- 
lected in Skane, southern Sweden, and kindly sent by Dr. Gote Turesson. 

2. Artemisia Michauxiana Bess, in Hook., Fl. Bor. Amer. 1 :324, 1833. 

Figures 10 and 11 

Type statement : "Rocky Mountains. Douglas. Columbia. Douglas, in Herh. 
Lindl." The type is in the Lindley Herbarium, according to a note attached 
to a sheet in the Bentham Herbarium at Kew, which is said to correspond 
exactly with the type (fide R. A. Rolfe) . This Kew sheet has been labeled in 
Douglas' hand, "In fissures of dry rocks near the Kettle Falls and sources of 
the Columbia. 1826." A piece of this sheet was removed and sent to New York, 
where it has been labeled as part of the type. The first collection cited doubt- 
less came from the Rocky Mountains of Canada; the second may have come 
from the state of Washington. 

Artemisia discolor Dougl. ex Bess., Bull. Soc. Nat. Mosc. 9:46, 1836. The type has been 
studied on loan from Kew. It was collected "on rocky situations near Spokane and 
the Kettle Falls," by David Douglas. It is a somewhat coarser plant than the type 
of A. Michauxiana, which came from a higher elevation, but the two can scarcely be 
separated morphologically. Its inflorescence is considerably more open-paniculate 
than that of Michauxiana, but this character proves to be variable and lacking in 
geographic significance in this region as is also the vesture of the herbage. 

A. graveolens Eydb., Bull. Torr. Club 24:296, 1897. Leaves greener than in the type of 
Michauxiana, but much dissected and to be classed here ratlier than in incompta. 
Type locality: Yogo Baldy, Little Belt Mts., Montana. 

A. subfflabra A. Nels., Bull. Torr. Club 27:36, Jan. 24, 1900. The same form as A. 
graveolens. Type locality : stony banks of Yellowstone Eiver, near Yancey's, Yellow- 
stone Park, "no. 5743." 

A. tenuis Eydb., Mem. N. Y. Bot. Gard. 1:431, Feb. 15, 1900. A slender, small-headed form 
of graveolens, i.e., the green form of the species. Type locality: Emigrant Gulch. 
Park Co., Montana. 

A. vulgaris ssp. discolor Hall et Clem., Carn. Inst. Wash. Publ. 326 : 74, 1923. 

A. vulgaris var. discolor Jeps., Man. Fl. PI. Calif. 1141, 1925. 

A. vulgaris ssp. Michauxiana St. John, Ees. Stud. St. Coll. Wash. 1:106, 1929. 

A. vulgaris ssp. Michauxiana var. typica St. John, ihid. 

A. discolor var. glandulifera Hend., Ehodora 32:27, 1930. Leaves not tomentose beneath 
but glandular. Type locality: eastern base of Steins Mts., Harney Co., Oregon. 

A. vulgaris var. glandulifera Peck, Man. Higher Pis. Ore. 766, 1941. 

Stems 2-4 (-7) dm. tall; herbage white-tomentose throughout when young 
to nearly glabrous and green ; leaves short and broad, rather crowded, com- 
monly bipinnately divided with the secondary segments again toothed, the 
linear lobes widely spreading, acute, rarely elongated, even the uppermost 
leaves rarely entire, 2-5 cm. long, typically bright green above and white- 



434 



CALIFORNIA ACADEMY OF SCIENCES 



[Proc. 4th Ser. 




Figure 11. Distribution of Artemisia Michauxlana. 



tomentose beneath, the margin plane or narrowly revolute; inflorescence a 
slender racemose or spike-like panicle, occasionally more open, not leafy; 
heads large, nodding at least at first ; involucre hemispheric, 3.5-4 mm. high, 
(2.5-) 3.5-5 mm. wide, glabrous or sparingly tomentose, with 9-16 bracts; 
ray-florets 9-12 ; disk-florets 15-35. 



Vol. XXV] KECK: EEVISION OF ARTEMISIA VULGARIS COMPLEX 435 

Rocky Mountains of northern Alberta and British Columbia, southward 
through the mountains to northern Wyoming, northern Idaho, and central 
Washington; local in Steins Mountains, Oregon, and in Washoe County, 
Nevada. Occurring at elevations of 500 to 2750 meters, and flowering from 
July to August. 

Alberta. Moberly's on Atliabaska, Scliaffer 1613 ; Cataract Creek, Brown 1471 ; Poboctan 
Pass; Lake Louise; Vermilion Mt. ; Highwood Eiver District, Brln'kDian 3209. British 
Columbia. Mt. Atlin, Lake Atlin region, Swarth 177 ; Telegrapli Creek, Walker 1208. Cari- 
boo: Mt. Sehvyn, Raup and Abbe 3946. Kootenay: Mt. Whaleback, Yolio Park, JJllce 334; 
Mt. Wapta; Kicking Horse Eiver; Carbonate Draw, Selkirk Mts., Shaw 335 ; Goldstream; 
Revelstoke. Yale: Eraser Valley; Lake House, Skagit River, Macoun 76927. New West- 
minster: Yale; Chilliwack Valley. Montana. Glacier: Midvale, Umbaoh 275 ; Many Glacier. 
Flathead: Avalanche Lake; Columbia Falls, Williams 709 ; Mission Creek. Lincoln: To- 
bacco Mts. Judith Basin: Yogo Baldy, Aug. 24, 1896, 7000 ft., J. E. Flodman 881 (NY, 
type of A. graveolens) ; Long Baldy, Little Belt Mts., Aug. 19, 1896, 7-8000 ft., Flodman 
881 (US, labeled isotype of A. graveolens). Deer Lodge: Eyans Lake; Mt. Haggin. Galla- 
tin: Mt. Bridger. Park: Emigrant Gulch, 6500 ft., Rydberg and Bessey 5201 (NY, type 
of A. tenuis; isotype, US) ; Electric Peak. Wyoming. Sheridan: Little Tongue Eiver Can- 
yon, Williams 3233. Johnson: headwaters of Clear Creek and Crazy Woman Eiver (NY). 
Big Horn: Doyle Creek, Goodding 385. Park: Meeteetse Creek, Griffiths. Yellowstone Park: 
Yellowstone Eiver near Junction Butte, Nelson 5743 (EM, type of A. sub glabra; isotype, 
NY). Teton: Leigh Lake, Williams 994. Utah. Summit: Blacks Fork, Uinta Mts.; Upper 
Henry's Fork Basin, Maguire et al. 14412, 14586. Idaho. Idaho : mouth of Salmon Eiver, 
Eastwood 13337. Washington. Okanogan: Oroville; Mt. Chapaca, Elmer 589; Tonasket; 
Hidden Lakes; Angels Pass, Thompson 7033. Whatcom: Newhalem, Muenscher 9919. 
Chelan: Nigger Creek; Three Brothers. Yakima: Mt. Adams, Suksdorf (C). Oregon. 
Harney: Steins Mts., Cusick 1991; alpine rivulet near eastern base of Steins Mts., beyond 
Alvord, Henderson 8489 (CAS, isotype of A. discolor var. glandulifera) . Nevada. Washoe: 
head of Galena Creek, 8500 ft., Heller 10222. 

The exact southern limits of this species are difficult to determine on mor- 
phological criteria alone. Plants from the alpine zone in parts of Nevada, as 
in the Ruby, White, and Charleston mountains, which appear very similar to 
the tenuis form of Michauxiana, are referred in this paper to A. ludoviciana 
ssp. incompta. This is because intergrades seem to connect these forms with 
very narrow, dark green leaf -segments with forms having the more usual type 
of incompta leaf, and these Michaiixiana-WkQ forms are fairly well isolated 
from the other members of Michmixiafia. Their disposition would be more 
satisfactorily made if cytological data were available. 



3. Artemisia longifolia Nutt., Gen. 2 : 142, 1818. 

Figures 5, 10, and 12 

Type statement : "In rocky situations, on the banks of the Missouri, from 
the confluence of White River to the Mountains?" A single leaf preserved 
at the Philadelphia Academy, labeled "Artemisia longifolia Missouri" in 
Nuttall's hand, is doubtless from the type. 



436 



CALIFORNIA ACADEMY OF SCIENCES 



[Proc. 4th Ser. 



A. ludovieiana var. integrifolia A. Nels., First Eep. Fl. Wyo. 138, 1896. Based upon A. 

integrifolia of Pursh, Fl. Amer. Sept. 2:520, 1814. Not A. integrifolia L., 1853. 

Pursh attempted to identify a specimen collected on bluffs of the Missouri Eiver, 

October 1, 1804, by Lewis, no. 53, witli the Linnaean species. Lewis' specimen at 

the Philadelphia Academy has been examined. 
A. naironensis A. Nels., Bull. Torr. Club 26:485, 1899. A wide-leaved form of the species. 

"Type no. is 568, Willow Creek [Wyoming], July 20, 1894, distributed as a variety 

of A. ludovieiana." 
A. falcata Eydb., N. Amer. Fl. 34:271, 1916. The form with salient linear lobes on the very 

narrow leaves, but in all other respects typical. Found on the southeastern border 

of the range of this species; the type locality is Fort Pierre, South Dakota. 




Figure 12. Distribution of Artemisia longifolia and A. serrata. 

A. vulgaris ssp. longifolia Hall et Clem., Carn. Inst. Wash. Publ. 326:78, 1923. 
A. vulgaris var. longifolia Peck, Man. Higher PI. Ore. 767, 1941. 

Stems 2-8 dm. tall, usualh- rather stout, erect, herbaceous down to the 
woody caudex, white-tomentose ; leaves linear or linear-lanceolate, merely 
acuminate or, more often, caudate-attenuate, usually entire, sometimes with 
a few slender teeth or lobes, occasionally the lobes caudate-elongated, 3-12 
cm. long, 2-5 (-6) mm. wide, those of the inflorescence very narrow and short, 
aJl loosely floccose and usually becoming glabrate or green above, densely 
tomentose beneath, often strongly revolute ; inflorescence a strict short panicle, 
sometimes elongated or rather open; heads mostly erect and peduncled; in- 
volucre campanulate or ± turbinate, 4-5 mm. high, 3-4 mm. wide, tomentu- 
lose to densely tomentose, with 9-17 bracts ; ray-florets 3-10; disk-florets 8-26. 
n = 18. 



Vol. XXV] KECK: REVISION OF ARTEMISIA VULGARIS COMPLEX 437 

Prairies and badlands from Saskatchewan and Alberta to central North 
and South Dakota, eastern Montana, and southern Wyoming, often in heavy 
alkaline clays. Flowering from July to September. 

Saskatchewan. 1858, Bourgeau. Alberta. W. of Lacombe, Moss; Eosedale, Moodie 
1097. North Dakota. McLean, Morton, McKenzie, Stark, Billings, and Golden Valley 
counties. South Dakota. Evarts, Missouri Eiver ; Grand River. Stanley : Fort Pierre, Sept. 
29, 1897, Griffiths 757 (US, type of A. falcata; isotype, US; phototypes, C, NY, US). 
Washaliaugh : Pine Ridge Reservation. Washington : Sheep Mt. Harding : Glaring Fork ; 
Short Pine Hills. Lawrence (?): Crow Buttes. Pennington: Big Bad Lands, Osterlwut 
7529. Montana. Milk River. Dawson: Colgate, Sandberg et al. 1011; Glendive. Toole: 
Shelby, Jones. (Po). Wyoming. Natrona: Willow Creek, Nelson 568 (RM, type of A. 
natronensis ; isotype, Minn). Johnson: Lake DeSmet, Nelson S545. Carbon: Como Ridge 
near Medicine Bow, Osterlwut 6290. Converse : Badlands, T. 38, R. 68. Fremont : Muskrat 
Creek, Goodding 517. Sweetwater: Table Rock; Point of Rocks, Nelson 8148. 

Chromosome counts were made in root-tips of four plants from Billings 
County, North Dakota. The living plants were supplied us through the kind- 
ness of Mr. E. C. Moran. 

4. Artemisia serrata Nutt., Gen. 2 : 142, 1818. 

Figures 4, 10, and 12 
Type statement : "Near the Prairie du Chien, on the banks of the Mississippi, 
also on the banks of the Missouri, in open alluvial soils." Upper Louisiana, 

Nuttall (Ph, type). 

Artemisia ludoviciana var. serrata ToRR. et Gray, F1. N. Anier. 2:420, 1843. 
A. vulgaris ssp. serrata Hall et Clem., Carn. Inst. Wash. Publ. 326:79, 1923. 

Stems 10-25 dm. tall, virgate, herbaceous; leaves lanceolate to linear- 
lanceolate, acuminate, sharply and rather finely serrate from apex well down 
toward base (rarely subentire), 8-12(-15) cm. long, 12-20 mm. wide, those 
of the inflorescence prominent but gradually reduced in size, all dark green 
and glabrous above, white-tomentose beneath; inflorescence a leafy panicle 
5-15 cm. broad; heads small, mostly erect, rather crowded; involucre cam- 
panulate, 2.3-3 mm. high, ca. 2 mm. wide, slightly floccose, with 8-12 bracts; 
ray-florets 6-11 ; disk-florets 3-16. n = 18. 

Upper Mississippi Valley ; Minnesota to Illinois and Iowa ; introduced in 
New York. Flowering from July to September. 

Illinois. Winnebago, Jo Daviess, Rock Island, and Henderson counties. Wisconsin. 
Dane: Wiibe (US). Minnesota. Chisago: Taylors Falls, B. C. Taylor. Isanti: Sandberg 
(C). Mille Lacs: Milaca, Sheldon 3278. Stearns: Grand Lake; Rockville, Campbell. Henne- 
pin : Fort Snelling ; Minnesota River near mouth of Nine Mile Creek. Wabasha : Lake City. 
Winona: Wiscoy, Holzinger ; Winona. Houston: Jefferson. Iowa. Fayette: Finlc 406. 
Poweshiek: Grinnell. Jasper: Baxter (US). 

Chromosome counts were made in root-tips of two plants from St. Paul, 
Minnesota. These plants were supplied through the kindness of Dr. C. O. 
Rosendahl. 



438 CALIFOBXIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

5. Artemisia Carruthii Wood ex Cairuth, 
Trans. Kans. Acad. Sci. 5 : 51, 1877. 
Figures 2, 10, and 13 
Type locality : Kansas. Type not seen. 

Artemisia Wrightii Gray, Proc. Amer. Acad. 19:48, 1883. The type collection, Wrlglit 
1279, made at Santa Rita del Cobre, Grant Co., New Mexico, is scarcely morphologi- 
cally distinguishable from material from the type region of Carruthii. The leaves 
are greener and glabrate on the upper surface in Wrightii, tomentose in Carruthii, 
but this feature does not appear in this case to distinguish ecotypes or geographical 
sulispecies. While the Wrightii form does not occur to the eastward, it grows together 
with the typical form at many localities in New Mexico. The very broad range of 
overlapping of the two forms largely obscures any difference in ecological preference 
that may exist betAveen them, although the incipient stages of ecotype formation 
may be involved. 

A. kansana Britt. in Britt. and Brown, 111. Fl. 3 :466, 1898. A form identical with Carruthii. 
Type locality: Lane County, Kansas. 

A. coloradensis Osterh., Bull. Torr. Club 27:506, 1900. Leaf-segments unusually wide, 
otherwise indistinguishable. In the type, from Dale Creek, Larimer Co., Colorado, 
the segments are less than 1 mm. wider than those of Carruthii from Kansas, but 
other collections from the same county are broader and intergrade toward the Brit- 
tonii form of ^. ludoviciana typicd. 

A. Balceri Greene, PI. Baker. 3:31, 1901. Allied to the Wrightii form in that the leaves 
are green on the upper surface, but these are atypically scattered and wide, giving 
some suggestion of an extremely narrow-leaved form of A. ludoviciana mexica.'na. 
This does not appear to warrant separation from the more usual forms of Carruthii, 
however, for plants of this appearance are found in the western portion of the range 
of the species in scattered stations between which more typical forms are found. In 
other words, the variant is of spasmodic occurrence and by no means of uniform 
appearance. The inflorescence is usually open-paniculate. Type locality: canyon of 
the Gunnison River, near Cimmaron, Colorado. 

A. Pringlci Greenm., Proc. Amer. Acad. 40:50, 1904. Heads larger than usual, containing 
tvdce the average number of florets for the species as a whole in the type collection 
(ca. 18 involucral bracts, 14-18 ray-florets, 20-34 disk-florets), but subject to con- 
siderable variation in the region from which the type came. Not readily separable by 
any character from Carruthii. Typo locality : plains near the city of Chihuahua. 

A. Wrightii coloradensis A. Nels. in Coult. and Nels., Man. Rocky Mt. 568, 1909. 

A. mexicana Balceri A. Nels., ihid. 569. 

A. vulgaris ssp. Wrightii Hall et Clem., Carn. Inst. "Wash. Publ. 326 : 80, 1923. 

A. vulgaris var. Wrightii Palmer et Steyerm., Ann. Mo. Bot. Gard. 22:677, 1935 ; Rhodora 
40:134,1938. 

A. vulgaris Carruthii F. C. Gates, Trans. Kansas Acad. Sci. 42 : 138, 1940. 

A. CarruthiiYar. Wrightii Blake, Jour. Wash. Acad. Sci. 30:472, 1940. 

Stems 3-6 dm. tall; herbage white-tomentose thronphont or the upper sur- 
face of the leaves preen ; leaves divided into divaricate linear-filiform lobes, 
these sometimes again toothed, 1-3 cm. long, those of sterile shoots and of the 
inflorescence often entire, reduced, the margin closely revolute; inflorescence 
a leafy narrow spike-like panicle, 1-3 cm. wide, occasionally rather expanded ; 



Vol. XXV] KECK: BKVISION OF ABTEMISIA VULGABIS COMPLEX 439 




Figure 13. Distribution of A. Carruthii. 

heads small, nodding when young; involucre eampanulate, 3-3.5 mm. high, 
1.5-3 mm. wide, tomentulose, often shining, with 8-14 bracts; ray-florets 
6-14 ; disk-florets 6-20. n = 9. 

Western Kansas and northern Texas to central Utah, Arizona, and northern 
Chihuahua, at elevations from 800 to 3000 m. Adventive in Rhode Island, 
Indiana, Missouri, and eastern Kansas. Flowering from late July through 
September. 



440 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

Kansas. Eooks: Rockport. Lane: plains, Aug. 15, 1895, HitchcocTc 302 (NY, type of A. 
Icansana; isotypes, RM, US). Greeley: Tribune. Kearney: Hitchcoclc. Hamilton: Syracuse, 
Bose and Fitch 17010. Clark: White. Meade: Meade. Seward: Liberal. Texas. Potter: Ama- 
rillo, Bailey 12S. Randall: Paloduro Canyon. Colorado. Larimer: Dale Creek, Sept. 7, 1899, 
Osterhout 2010 (RM, type of A. coloradensis ; isotypes, NY, Po, RM). Eagle: Red Cliff. 
Park: Como. Douglas: Pine Grove. El Paso: Palmer Lake; Colorado Springs. Teller: 
Florissant. Fremont: Canon City; Wet Mountain Valley. Saguache: Marshall Pass, 10,000 
ft., BaTcer 879 ; San Luis Valley. Montrose: Bedrock. Gunnison: Black Canyon, Baker 698 
(C, NY, Po, RM, SU, US, topotypes of A. Bakeri). Ouray: 5 mi. N. of Ouray. San Miquel: 
Norwood Hill. Huerfano : Cucharas Valley. Costilla : La Veta Pass. Mineral : Wagon Wheel 
Gap. Hinsdale: Lake City. Archuleta: Piedra. La Plata: W. of Hesperus. New Mexico. 
Union: Folsom. Colfax: Ute Park. Rio Arriba: Dulce. San Juan: Tunitcha Mts., Standley 
7622. Quay: Nara Visa; Llano Estacado. San Miguel; Las Vegas. Santa Fe: Canoncito. 
Sandoval: Jeniez Springs. Valencia: Grants. Lincoln: Ruidoso. Dona Ana: Organ Mts. 
Socorro : Negrito Ranger Station. Catron : Patterson. Grant : Santa Rita del Cobre, Wright 
1279 (Gray Herb., type of A. Wrightii; isotype, NY; phototypes, C, NY); N. of Cliff. 
Hidalgo: Cloverdale, Mearns 456. Utah. Sevier: Mt. Marvin, Rydberg and Carlton 7567. 
Piute : Circle Valley. Garfield : Panguitch Lake. San Juan : Montezuma Canyon, E. of 
Monticello, Rydberg and Garrett 9687 ; Elk Mts.; Bluff. Arizona. Apache: Carrizo Mts., 
Standley 7510 ; Greer. Navajo: Oraibi, Voth 76. Coconino: Grand Canyon; Mt. Humphreys; 
7 mi. E. of Flagstaff; Bellemont, Jones 4074 ; Williams. Mohave: Hualpai Mts., Kearney 
and Peebles 12727. Yavapai: Prescott. Gila: Black River Bridge. Greenlee: Metcalf. Gra- 
ham: Mt. Graham. Cochise: Chiricahua Mine, Blumer 1794. Chihuahua. Lake Santa Maria; 
plains near Chihuahua, Pringle 625 (Gray Herb., type of A. Pringlei; isotj'pes, Ph, SU, 
US) ; Santa Eulalia Plains, Wilkinson. Chihuahua or Coahuila. Charco de Grado, Oct. 
1852, Thurber 791. Sonora. San Luis Mts., Mearns 2461; Badehuachi, Lloyd 404. 

Chromosome counts were made in root-tips of three plants grown from seed 
from Springer, Colfax County, New Mexico, and kindly supplied by Mrs. 
Gladys Nisbet. 

In Colorado, on the one hand, occasional collections appear to be inter- 
mediate with the Brittonii form of A. lucloviciana typica, while in New 
Mexico, on the other, certain collections are found that are intermediate with 
A. ludoviciana mexicana. More chromosome counts will help clarify this 
situation and show whether, as is now suspected, all forms of liidovwiana are 
tetraploid, and of Carruthii diploid. 

6. Artemisia ludoviciana Nutt., Gen. 2 : 143, 1818. 

6a. Artemisia ludoviciana Nutt. ssp. typica Keck, Cam. Inst. 
Wash.Publ. 520:330, 1940. 

Figures 7, 10, and 14 
Type statement: "On the banks of the Missisippi. near St. Louis; also on 
the alluvial plains of the Missouri." A specimen at Philadelphia from St. 
Louis, collected and starred by Nuttall, is the type. It consists of only four 
leaves, lightly tomentose above and densely tomentose beneath. Another 
Nuttall specimen at Philadelphia, labeled by Nuttall "Artemisia Ludoviciana. 
Platte plains," is not starred, but is a large branch in flower with all leaves 
entire and tomentose on both sides. 



Vol. XXV] KECK: EEVISION OF ABTEMISIA FULGAEIS COMPLEX 



441 



Arternisia ludoviciana Nutt., loc. cit. 

A. gnaphalodes Nutt., ihid. Type statement: "On dry savannahs about Green Bay, Lake 
Michigan, and on the banks of Fox river, and the Missouri." A starred specimen at 
the Philadelphia Academy, collected by Nuttall at St. Louis, was considered by Hall 
and Clements to be the type, but its location is inconsistent with the type statement. 
It consists merely of the top of a flowering stem, in which the leaves are white- 
tonientose on both sides. See discussion of this synonym on p. 423. 

A. Pursliiana Bess, in Hook., Fl. Bor. Amer. 1:323, 1833. Type locality: "On the plains of 
the Saskatchewan. Dr. RicJmrdson. Drummond." See the following. 




Figure 14. Distribution of A. ludoviciana ssp. typica. 



A. Pursliiana a latifolia Bess., ibid. Specimen at Kew (Herb. Hooker), which has been 
studied, taken in the Eocky Mts. by Drummond, is doubtless of the type collection. 
By his var. latifolia, Besser meant the typical form of Purshianu (as both species 
and variety seem to rest on the same type) as contrasted with his var. y3 angustifolia, 
also described here and based on a plant from "Red Eiver. Douglas, in Herb. Lindl.," 
which is said to have lanceolate leaves 1% inches long and 3 lines wide. From this 
description it appears probable that this variety, too, is referable here. 

A. ludoviciana var. gnaphalodes ToRR. et Gray, Fl. N. Amer. 2 :420, 1843. 

A. ludoviciana var. latifolia ToRR. et Gray, ibid. 

A. vulgaris var. gnaphalodes O. Kuntze, Rev. 1 :309, 1891. 

A. vulgaris var. ludoviciana O. Kuntze, ibid. 

A. rhizomata A. Nels., Bull. Torr. Club 27:34, 1900. A narrow-leaved form. Type locality: 
Sweetwater River, Wyoming. 



442 CALIFOBMA ACADEMY OF SCIENCES IProc. 4th Ser. 

A. rhizomaia var. pahularis A. Nels., ihid. A small form of the preceding. Type locality: 
Red Desert, near Creston, Wyoming. 

A. diversifolia Rydb., Bull. Torr. Club 28:20, 1901. A form with the principal leaves merely 
pinnately cleft, but the few lobes very narrow and acuminate. It closely resembles 
forms of ssp. candicans from the general region of northern Idaho, and should per- 
haps be interpreted as a transitional form. Type locality: Priest River, Idaho. 

A. silvicola Osterh., Bull. Torr. Club 28:645, 1901. A form with lanceolate, nearly entire, 
strongly discolored leaves and large heads. Type locality : Maclntyre Creek, Larimer 
Co., Colorado. 

A. Briitonii Rydb., Bull. Torr. Club 32:129, 1905. Leaves short, with 3 to 5 short rather 
divergent lobes, heavily tomentose on both sides. Type locality: Golden, Colorado. 

A. pudica Rydb., ibid. 130. Inflorescence loosely branched, the large heads nodding, leaves 
rather thin and elongated. A spasmodic variant from widely scattered stations in the 
Colorado Rockies. Perhaps usually a shade modification. Type locality: Gunnison. 

A. pabulari,s Rydb., op. clt. 33 : 157, 190(). 

A. gnaphalodes diversifolia A. Nels. in Coult and Nels., Man. Rocky Mt. 569, 1909. 

A. mexicana silvicola A. Nels., ibid. 

A. H err iota Rydb., Bull. Torr. Club 37:455, 1910. A large-leaved form from Edmonton, 
Alberta. 

A. cuneata Rydb., N. Amer. Fl. 34:269, 1916. Leaves cuneate obovate, with a few salient 
teeth around the wide apex. Type locality: Idaho Springs, Colorado. 

A. argophylla Rydb., ibid. 274. A robust form with exceptionally large heads. The type, 
from Longs Peak, Colorado, is well east of the range of ssp. candicans, which form 
it resembles. 

A. vulgaris ssp. ludoviciana Hall et Clem., Carn. Inst. Wash. Publ. 326 : 76, 1923. 

A. vulgaris ssp. gnaphalodes Hall et Clem., ibid. 77. 

A. ludoviciana var. pabularis Fern., Rhodora 47:248, 1945. 

A. ludoviciana var. Briitonii Fern., ibid. 

A. ludoviciana var. cuneata Fern., ibid. 

Stems 3-10 dm. tall, slender to moderately stout, herbaceous to base; leaves 
many, linear to lanceolate, oblanceolate or elliptic, sometimes cuneate, entire or 
few-toothed or -lobed especially apically, the lobes regularly entire, 3-10 cm. 
long, permanently and densely white-tomentose on both sides or loosely floc- 
cose to green and glabrate above, plane ; inflorescence an elongated usuallj' 
compact panicle, 1.5-5 em. broad ; heads medium-sized, erect or nodding, often 
in glomerules; involucre ovoid to campanulate, 3^ mm. high, 2—3 mm. wide, 
usually densely tomentose, occasionally glabrate, with 7-13 bracts ; ray-florets 
5-12 ; di.sk-florets 6-21. n = 18. 

Lowlands and mountains up to middle elevations (2200 m., or occasionally 
up to 2900 m. in the Colorado Rockies) from Ontario to Alberta, Washington, 
California, New Mexico, and Arkansas ; sporadically introduced from Indiana 
to Virginia, New England, and Quebec. Flowering from July through Sep- 
tember. 

Ontario. Thunder Bay: near Fort William. Saskatchewan. Assiniboia, Indian Head, 
Macoiin 10084 ; Ribstone Creek ; Moose Jaw, Johnson 069. ? province, plains of the Saskatche- 
wan, Dnimmond (Kew, type(?) of A. rurshinna var. hitifolia). Alberta. Stettler District, 
Donalda to Bashaw; Peace River; Grand Prairie; Edmonton, Macovn and Ilcrriot 72825 
(NY, type of A. Ilerriotii) ; 'RoseAa.\e,Moodic 1183 ; Banff; Lethbridge. Bkitlsh Columbl^. 
Kootenay: Beavennonth, Shaxv 1153; Natural Bridge, TJlke. Indiana. Marshall, Fulton, 



Vol. XXV] KECK: BEFISIOX OF AETEMISIA VULGARIS COMPLEX 443 

Newton, and Sullivan counties. Illinois. Dupage, Ogle, and Eock Island counties. WiscoN- 
.siN. La Crosse, Sauk, Dane, and Rock counties. Minnesota. St. Louis, Norman, Traverse, 
Winona, Pope, and Hennepin counties. Iowa. Emmet, Fayette, Linn, Benton, Poweschiek, 
Boone, and Decatur counties. Missouri. Clark : Mead. Jackson : Sibley. Arkansas. Craig- 
head: S. of Jonesboro, Bemaree 7179. North Dakota. Cass, Eanson, Stutsman, Benson, 
Burleigh, McLean, and Golden Valley counties. South Dakota. Grant, Spink, Kingsbury, 
Harding, Butte, Meade, Lawrence, Custer, and Fall River counties. Nebraska. Douglas, 
Cass, Lancaster, Kearney, Brown, and Cherry counties. Kansas. Riley, Saline, McPherson, 
SedgA\dek, and Pratt counties. Oklahoma. Woods, Cimmaron, Kingfisher, and Harmon 
counties. Texas. Wood: Eeverchon 3961. Montana. Richland, Glacier, (Glacier National 
Park), Flathead, Sanders, Lewis and Clark, Yellowstone, Cascade, Carbon, Sweetgrass, 
Park, Gallatin, Jefferson, Deer Lodge, Silver Bow, Ravalli, and Beaverhead counties. 
Wyoming. Crook, Albany, Sheridan, Johnson, Natrona, Park (Yellowstone National Park), 
Teton, Lincoln, Carbon, Sweetwater, and Uinta counties. Colorado. Sedwick : Julesburg. 
Weld: New Windsor. Larimer: Mclntyre Creek, Aug. 24, 1900, Osterhout 2242 (NY, type 
of A. silvicola; isotypes, C, RM, US) ; Meadows Inn, Longs Peak, Aug. 14, 1907, Clements 
(NY, type of A. argyrophylla). Boulder: Foot of Flagstaff Mt., Ewan in PI. Exsic. Gray. 
897. Routt: S. of Steamboat Springs. Jefferson: Golden, Oct. 8, 1882, N. L. Britton (NY, 
type of A. Brittonii). Grande: Hot Sulphur Springs. Clear Creek: Idaho Springs, Shear 
4617 (NY, type of A. cuneata). Rio Blanco: Buford. Garfield: Glenwood Springs, Eggle- 
ston 14982. Otero: La Junta. Fremont: Canon City, Brandegee 501. Huerfano: Cucharas 
Valley. Alamosa: Alamosa. Gunnison: Gunnison, Baker 573 (NY, type of A. pudica; 
isotypes, C, CAS, Po, RM, US). Mineral: Pagosa Peak, Balcer 634. San Miquel: Norwood 
Hill. Montezuma: Mesa Verde National Park. New Mexico. Colfax: Raton. Rio Arriba: 
Brazos Canyon ; Dulce. San Juan : Tunitcha Mts. San Miguel : Las Vegas. Sandoval : Sandia 
Mts., EUis 125. Grant: mts. near Condes Camp, Wrigld 1275. Utah. Rich: Bear Lake, 
Hammond 3879. Box Elder: Corinne. Weber: Ogden Canyon. Salt Lake: Parleys Park. 
Piute: Marysvale, Jones 5846. San Juan: Dry Wash, Abajo Mts., Ryherg and Garrett 
9615. Iron: Parowan. Arizona. Coconino: Grand Canyon, Eggleston 15665. Idaho. Bonner: 
Slough beach, forest between Priest Lake and East Fork, 630 m., MacDougal 190 (NY, 
type of A. diversifolia ; Clarks Fork Valley near Horse Plains, Leiberg 1582. Kootenai: 
Coeur d'Alene, Leiberg 1553. Lemhi: Gilmore. Fremont: Henry's Lake, Nelson 6796. 
Bannock: Lava. Bingham: Blackfoot. Blaine: Ketchum. Twin Falls: Shoshone Falls, 
Nelson and Macbride 1366. Gem: Sweet, Machride 1626. Nevada. Elko, Eureka, Lander, 
Humboldt, White Pine, Nye, Pershing, Churchill, Washoe, Ormsl^y, Storey, and Lyon 
counties. Washington. Pend Oreille: Kalispell Lake. Chelan: Lake Chelan. Klickitat: 
Columbia River, Suksdorf 1610. Oregon. Wallowa, Union, Baker, Grant, Deschutes, Crook, 
Malheur, Harney, Lake, and Klamath counties. California. Modoc, Nevada, Mono, Inyo, 
San Bernardino, and Riverside counties. 

Chromosome counts were made both in root-tips and in meiosis of one plant 
from Manitou and of two plants from Engelmann Canyon, all from El Paso 
County, Colorado, and in root-tips only of one plant from Leevining, Mono 
County, California, and of two plants from Upper Fern Valley, San Jacinto 
Mts., Riverside County, California. 

An additional very interesting tetraploid, whose chromosomes were counted 
in root -tips, is now referred here. It was collected in Poopenaut Valley, Tu- 
olumne River, Yosemite National Park, at 1065 m. elevation, Nohs 1 (CI, 
RM). This plant, from the midst of the area occupied by the hexaploid A. 
Douglas'iana, near the western base of the central Sierra Nevada, was pre- 
sumed to be a peculiar form of that species when first studied. It had dis- 



444 CALIFORNIA ACADEMY OF SCIENCES [Proc. 4th Ser. 

colored leaves like Douglasiana, but these were extremely narrow ; those on 
the upper half of the stem were several centimeters long but less than 5 mm. 
wide. The lower leaves were oblanceolate and wider (up to 20 mm. including 
lobes) , with rather short forward-projecting lobes in the outer half, precluding 
the possibility that this could be a form of A . ludoviciana ssp. incompta carried 
down from higher elevations. This plant is best explained as a relict of ludo- 
viciana ssp. typica stranded on the wester