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The pride of Guadalupe Island, the endemic Cistanthe guadalupensis, flowering on a small islet

off the southwest coast, with cliffs of the main island as a background; 19 April 1957
his plant is rare on the main island, surviving only on cliffs out of reach of goats,

but common here on goatless Islote Negro

THE FLORA OF GUADALUPE ISLAND, MEXICO

THE FLORA OF GUADALUPE ISLAND,
MEXICO

Reid Moran

Published by the California Academy of Sciences
San Francisco, California

Memoirs of the California Academy of Sciences, Number 19

San Francisco
July 26, 1996

SCIENTIFIC PUBLICATIONS COMMITTEE:

Alan E. Leviton, Editor

Katie Martin, Managing Editor
Thomas F. Daniel

Michael Ghiselin

Robert C. Drewes

Wojciech J. Pulawski

Adam Schiff

Gary C. Williams

All rights reserved. No part of this publication may be reproduced or transmitted in any
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any information storage or retrieval system, without permission in writing from the
publisher.

Library of Congress Catalog Card Number: 96-084362
ISBN 0-940228-40-8

TABLE OF CONTENTS

PADS tACh inne cyauee raychaty Mecrorne ann rs See Sta eres eae Raye aacte wine ereeteuatn ys nthevs dies Vil
FE SUITI CIRM tnt nies, case ton ain care toNelg in enag ee A Sie Rated ae OMA oN Ones Seer e vill
Mera LUT UY OTe ogee eee eens otra eas steve ay eueete eats eer neTsueesie Saeetene Geensyeneegee eters eaee l
Guadalupe Island
| DS Yos ao} 100) Dene eneaye onemeriecterinie trae eee crcecniean rere Caoeaecia Piece ecient cae ne rena aoe |
PAGO UMAMIES sy cearacie ae, | oe cotyetceee terescesteee ercverane + cede tear gem ave nah ucemayeertosy tu stotenete 9
Climates aro Sacanne = ote noone oon na See eatma noemen es wanna aes iN}
FELIS COG arctemenycceaeig see a tin eons tans ste Spy once ened esoars acne deh eievetexePe there 15
Other Blotais ss 44264628 seonsae nee Sesh eee nen eee on a OE Hae Ree Ee on 1S
The Vascular Plants
INALIVeOnMhOLe] SNe z en remem tare earn cto ot eet ee rece mitaiegs anc aretans spete as enresels 19
Goatsvand Plants) a. 42: eto wen ee ere Sere eterds sain ere erat ug Pa eet ass 23
Rasteandabicsente Vegetal online rere sersie ot ein cieeonnse sets. ShOPR dee have as 28
Eextincetbiantssand! Nearly; Extinct.a. wets coties tas athena awe atesedatenns 39
Composition and Relationships of the Flora.............000..00 0000022. 41
BlanteGollectors’ ccpaewrecancausttees castiatucis senate see ete tea rea Gre es S05, ee. omens: ator fensesas cot cute chan’ 44
Catalogue of the Plants
ST Ss tae ee MR sect anenraath tee Merah coh sey ee gee ated ay agen avis yA ce ea 53
GY TMMNOS METIS er yan-ceh er ivaes aims scsenere, Guess Maye e haar aneyareiay dis sn neath tes wach ne 56
DDG OLS OR ie ND 5 ay Sas bs ey sodestin eran tS aehieetiond Mita SOR MPNG, in ariarnas. Bela Rola ad (ela, she air ts 66
MOnOCOtSaagas catact a2 Se to that aia e Gap omatanat amend de albane toe 158
Doubtiuland*Exclided Species: 6 ics cece goes odes ee vs gel sae geen 168
NCKNOWIECEMENUS,, 5 ssn ccnn cis a mtyes Sua eis 6 leche as Lae S oat Reendeeantete Dares ays 170
[eiteraturey Gited ha. ear. ourcr se aayeaeee oS adie eon Sees Ae Sart ae eee yaa ene emai 170

Indexeto- Plant Nam 6S oc. 2 fa sosssd secenats- on cots Oe eusvay ataucha lat aiead wink doves See teteen arated & 185

THE FLORA OF GUADALUPE ISLAND, MEXICO

Reid Moran

Department of Botany, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118

Guadalupe Island is a volcanic oceanic island 260 km west of the peninsula of Baja
California, Mexico, and separated by ocean depths of 3600 m. Its area is 250 km™
and its height 1295 m. Rainfall averages 133 mm near sea level at the south end but
evidently is much more at the high north end, where there is no record. For at least
a century and a half, feral goats have degraded the island by destroying the vegetation,
thus (1) extinguishing native species, (2) fostering soil erosion, and (3) drying up
springs by killing trees whose fog drip had added greatly to rainfall.

Once-extensive forests of pine, oak, and cypress on the high northern third of the
island, and woods of juniper and palms (Brahea), have shrunk drastically and are
still shrinking, the remaining old trees unable to reproduce because goats eat the
seedlings. Sage scrub (Senecio etc.) in the northern half of the island has been replaced
by weedy grasslands, with shrubs surviving only on cliffs and some kinds gradually
dying out. Some scrub (Ambrosia, Atriplex, Lycium, Hemizonia) remains at low levels
in the drier southern third of the island, where fewer goats range for lack of water.
Offshore islets are refugia for rare plants, including five never found on the main
island.

The total known vascular flora is 216 species, but not all there at one time: over 30
of them are probably now extinct on the island, and new weeds keep coming as if
to take their place. Whatever we may mean by native on a volcanic oceanic island,
I count 45 species, mainly European weeds, as relative newcomers, thus leaving at
most 171 to consider as possibly native. However, a dozen of these were first found
in parts of the island where earlier collectors had failed to find them. The cumulative
evidence is that some must be recent arrivals; but not knowing which are, I keep
15 west American plants in the category of possibly or probably introduced, leaving
156 as less doubtfully native. Of these natives, 34 species and subspecies (21.8 percent)
are endemic, including the monotypic genera Baeriopsis J. T. Howell and Hesperelaea
A. Gray, though five or more endemics are extinct. Of the native flora, 114 species
(73 percent) grow also in Alta California and only eight (5 percent) in Baja California
but not northward. In particular, 101 (65 percent) grow on California islands; and
19 of these (12.2 percent) are insular endemics. Of the Guadalupe endemics, six have
closest relatives on other islands. Thus the relationships of the Guadalupe flora are
mainly to the north, with the islands of southern California: Guadalupe Island is an
outlier of the California Floristic Province.

It is most important before more plants are lost, to remove all goats from the island,
reversing the process of degradation and encouraging in every way the renewal of
the natural vegetation. Even at best, some rare plants may die out unless propagated
and replanted. Guadalupe Island with its unique flora is a Mexican national treasure
in dire need of protection.

Received March 3, 1995. Accepted June 6, 1995.

Vil

Resumen

La Isla de Guadalupe is una isla oceanica voleanica situada a 260 km al oeste de la
peninsula de Baja California, Mexico, y separada por profundidades oceanicas de
3600 m. Su area es de 250 km cuadrados y su altitud alcanza 1295 m. La precipitacion
pluvial promedia 133 mm cerca del nivel del mar en el extremo sur pero evidente-
mente is mucho mas en el elevado extremo norte, donde no existen registros. Por
cuando menos un siglo y medio, las cabras ferales had degradado la isla destruyendo
la vegetacion, de esta manera (1) extinguiendo especies nativas, (2) propiciando
erosion del suelo, y (3) secando manantiales al matar arboles cuya condensacion de
niebla complementaba grandemente la Iluvia.

Los en un tiempo extensos bosques de pinos, encinos, y cipreses, en el elevado tercio
norte de la isla, y montes de enebros y palmeras (Brahea) se han reducido drasti-
camente y continuan reduciendose, los arboles viejos remanentes imposibilitados para
reproducirse por que las cabras devoran los renuevos. El matorral de salvia (Senecio
etc.) en la mitad norte de la isla, ha sido reemplazado por maleza de pastizales, con
arbustos sobreviviendo unicamente en acantilados y algunos gradualmente muriendo.
Algun matorral (Ambrosia, Atriplex, Lycitum, Hemizonia) permanece a bajos niveles
en el mas seco tercio del sur de la isla, a donde van menos cabras por falta de agua.
Los islotes litorales son refugios para plantas raras, incluyendo cinco que nunca se
han encontrado en la isla principal.

La flora vascular total conocida es de 216 especies, pero no todas al mismo tiempo:
mas de 30 de ellas probablemente se han extinguido ya de la isla, y nuevas malezas
siguen Ilegando como para tomar su lugar. De lo que de alguna manera consideremos
nativo en una isla voleanica oceanica, yo recuento 45 especies, principalmente malezas
europeas, como arribos relativamente recientes, asi dejando cuando mucho 171 que
se consideran posiblemente nativas. Sin embargo, una docena de estas fueron por
primera vez encontradas en este siglo, en partes de la isla donde no las habian hallado
recolectores anteriormente. La evidencia acumulativa es de que algunos son arribos
recientes; pero no sabiendo cuales son, yo mantengo 15 plantas oeste-americanas en
la categoria de posible 0 probablemente introducidas, dejando 156 como menos
dudosamente nativas. De estas especies nativas, 34 especies y subespecies (21.8 por-
ciento) son endemicas, incluyendo los generos monotipicos Baeriopsis J. T. Howell y
Hesperelaea A. Gray, aunque cinco 0 mas endeémicas estan extinguidas.

De la flora nativa, 114 especies (73 porciento) crecen tambien en Alta California y
unicamente ocho (5 porciento) crecen en Baja California pero no mas al norte. En
particular 101 (65 porciento) crecen en las islas de California; y 19 de estas (12.2
porciento) son endémicas insulares. De las endemicas de Guadalupe, seis tienen sus
mas cercanas especies emparentadas en otras islas. Asi, las relaciones de la flora de
Guadalupe se encuentran principalmente al norte, con las islas del sur de California;
la isla de Guadalupe es un desplazamiento de la Provincia Floristica de California.

Es de la mayor importancia antes de que se pierdan mas plantas, remover todas las
cabras de la isla, revirtiendo el proceso de degradacion y fomentando en todas las
formas posibles la recuperacion de la vegetacion natural. En el mejor de los casos,
algunas plantas raras no podran sobrevivir a menos que puedan ser propagadas de
semillas nativas y replantadas.

La isla de Guadalupe con su singular flora, es un tesoro nacional mexicano en de-
sesperada necesidad de proteccion.

Vill

GUADALUPE ISLAND FLORA
Introduction

Guadalupe Island, westernmost outpost of Mexico, 1s a volcanic oceanic island some 260
km (162 statute miles) off the peninsula of Baja California, 337 km (210 miles) southwest
of Ensenada, and 400 km (250 miles) south-southwest of San Diego, California (Fig. 1).
It is separated from the continent by ocean depths of some 3600 m (2000 fathoms) and
evidently has never been connected.

Like other high oceanic islands, Guadalupe is remarkable for endemism in the flora and
fauna; but it is a naturalists’ paradise despoiled by feral goats, housecats, and mice. These
pests need to be exterminated before they destroy more of the unique endemics of this
remarkable island; and it 1s important to reverse the desertification, which in the long run
threatens even the goats themselves. Pending removal of all goats, fenced exclosures could
help the recovery of rare plants now close to extinction; but even so, some rare ones
might not survive unless they could be propagated, of course from native seed, and re-
planted. As further insurance, botanic gardens on the mainland can help preserve geneti-
cally adequate samples and seeds of Guadalupe plants. Perhaps the Center for Plant Con-
servation could slightly extend its area of concern to include this southernmost outher of
the California Floristic Province and could coordinate the effort. Meanwhile the Rancho
Santa Ana Botanic Garden is starting to help with this work.

Indeed, relationships of the vascular flora are mainly to the north, with the Channel Islands
of southern California, and to some extent with the islands off northwest Baja California
(Fig. 1). Hence the flora is of particular interest to Californian botanists like me. Thorne
(1969) described these islands as to area, height, distance from the mainland and from
each other, and geological history; and for these islands he discussed disharmonic biotas,
immigrant patterns of distribution, relict and autochthonous endemism, gigantism, con-
tinuous flowering, hybridization, and vulnerability. Wallace (1985) tabulated the known
floras of the eight Channel Islands plus Guadalupe, but there has been no full account of
the Guadalupe flora. So at last, here it is.

Guadalupe Island

DESCRIPTION

Guadalupe Island (Fig. 2) is about 37 km (23 miles) long north and south, 12 km (8 miles)
wide, and 1295 m (4257 feet) high, with an area of about 250 km~ (98 square miles). It
is rather uniformly high in the northern third, the highest point, Mt. Augusta, scarcely
rising above the general level. In the southern half is one rather sharp 975-meter
(3200-foot) peak, El Picacho. From there, the surface drops away to rather low elevations
at the south end, with scattered cones and craters (Fig. 16). Two of these features form
islets offshore: Inner and Outer Islets (Fig. 17).

The island is a seamount built on the axis of a fossil ridge crest or spreading center,
starting at most 10 million years after the abandonment of the ridge and perhaps much
earlier (Batiza 1977). The island consists of two partly overlapping shield volcanoes, both
collapsed calderas, vestiges of the southern one at El Picacho, the younger northern one
remaining as a half-crater at the north end (Figs. 3, 4). These are largely overlain by a
thick series of flank and fissure eruptions, from a set of north-trending fissures marked
by lines of cinder cones. Volcanic rock types form a remarkably complete series from
alkali olivine basalt to trachyte. The oldest exposed rocks are subaerial flows of alkali

REID MORAN

120° 118° 116° 114

_ Santa Barbara
Point Conception

Santa Cruz I.
San Miguel Ls

_ Los Angeles

=34
= Santa Rosa | cia
San Pedro Hill
Santa Barbara I. « California
col
pan Nicolas Tics Santa Catalina I.
= San Clemente \ be
_ San Diego
Los Coronados Is. ,
= 32°
= nee ; _ Ensenada
: Os Santos Is, . .
Pacific Ocean Baja Y
California
_ San Quintin
- — 30°
San Geronimo |. .
Guadalupe I \) ~
Sebastian
San Benito Is. ., Vizcaino
- Cedros | Bay — 28°
Nauvidad Ly

100 km

————_— aaa

100 miles

| | | | | | | |

FIGURE 1. The California Islands, off southern California and northwest Baja California. Based
on Hydrographic Office Chart 5760, 1942,

olivine basalt radiometrically dated at 7+2 million years BP (Engel and Engel 1964,
1971).

The northern upland is what remains of a caldera about 10 km (6 miles) wide, whose
collapsed eastern side is eroded away by streams forming two large canyons. The western
rim of the caldera is the main ridge of the island, which from the high point at Mt.
Augusta curves around to the north and northeast, to end in North Point, the northern tip

GUADALUPE ISLAND FLORA

118° 25' 118° 20' 118° 15' 118° 10'
zi} 1 1 1 \ 1 | 1 | | | | | E
= North Point =

Pilot Rock
- Pilot Rock Beach ~
Ties The Glacier

29° 10' - 4,9 Old Lookout e

Elephant Rock ae WS se = pe Northeast Anchorage
- feels me ae Barracks Canyon — (Campo Norte) 2

Wr a Oak Canyon
: * (i ae Hh mn Lower Circus
- tie 7 oad aa) ~ Lower Valley :
Elephant Seal Beach i
7 ephant Seal Beac a eas Spring e
1231 m & DiSgeos Esparsa Canyon
= Mt. Augusta aaa 1298 ~ ; <a South Esparsa Canyon z
29° 05' — Steamer Point S =
Dike Point

7 El Comal 7
= Long Canyon -

: Juniper Canyon

C Pist
= SEE Red Cinder Cone -
' Lobster Cam
29°.00' - Pp a
El Picacho
West Anchorage
S (Campo Oeste) Pillar Point =
: Double Point i
28055" — Yellow Point _
Islote Negro ————_QQ) \ ; > {f Plaza de Toros
Arroyo Melpomene yA vi if :
Ce. Oyae South Bluff
- Campo Sur = 7
(Weather ee Melpomene Cove
~ Inner Islet — Middle Rock Z
: lote Te
— — eae Over .
3.1 miles (Islote Zapato)

28° 50' 5 1 ' 1 1 I! ' ' 1 1 ' ' 1 ' 1 ! \ 1 F

FIGURE 2. Guadalupe Island, with place names used in this flora, Based on USDMA Chart 21661,

12th edition, 1982.

of the island (Figs. 3, 4). Below the rim in the caldera is an irregular shelf or upper valley
(Fig. 3). Here is the only permanent spring left on the island (Fig. 5), evidently fed partly
by fog drip from the cypress grove west of the ridge; in August 1981 it was flowing at

a constant 0.275 liters/second (Gonzalez 1981).

Early visitors described other springs

nearby, and according to Greene (1885), the springs fed plenty of shallow and tepid pools.

REID MORAN

FIGURES 3 and 4. Northeast upland, north from Mt. Augusta, the remnant of a caldera collapsed
and eroded away to the east (right); 2 March 1965. The far rim is the northeast ridge of the island,
ending in North Point to the right. FIGURE 3. Upper valley, draining south and then east, to Esparsa
Canyon (see Figs. 7, 8). Near the middle of the picture is the principal spring of the island (see
Fig. 5). FIGURE 4 (view to the right of Fig. 3). Lower Circus or inner are of cliffs, and below it
the lower valley, the near end draining into Esparsa Canyon, the far end into Barracks Canyon (see
Fig. 9).

GUADALUPE ISLAND FLORA

These springs have since dried up, doubtless from the loss of much of the cypress grove.
Stewart (1965) mentioned three sizable springs still there in 1899; and springy places still
remained north of the principal spring into the late 1950s. Here there was once a ranch,
or probably a series of ranches, mentioned but not described by early visitors; adobe ruins
seen near the spring as late as 1958 are now gone, but others to the northeast have not
yet entirely melted away.

Just to the east in the caldera is a lower, inner are of cliffs, which Franceschi (1893c)
called the Lower Circus, and below this a lower valley (Figs. 4, 6). The caldera drains
to the east, mainly by the two largest canyons of the island, Esparsa Canyon (Figs. 7, 8)
on the south and Barracks Canyon (Fig. 9) on the north.

At the mouth of Barracks Canyon is Northeast Anchorage, for many years the main
landing on the island (Fig. 10). It has been the site of various settlements — of fishermen,
goathunters, and soldiers. Just above the beach are two conspicuous old white buildings
known as the barracks, probably dating from World War II. At my first visit in 1948 the
remains of a goatmeat cannery were still to be seen on the beach. A broad trail climbed
by zigzags out of the canyon, then up the east slope and through the lower valley to the
spring and the cypress grove. The landing is now called Campo Norte. It is connected
by road, by way of the spring, with the airfield (Campo Pista) near the middle of the
island, and that with Campo Oeste at West Anchorage and with Campo Sur at the southwest
corner of the island.

Seacliffs are high, especially around the northern end. North Point, actually the northeast
point, stands 400 m above the sea. The north end has seacliffs 500 or 600 m high and a
steep slope above (Fig. 11), and at the northwest corner of the island, exposed to the full
force of the Pacific swell, slightly more rounded cliffs are some 700 m high. Two-thirds
of the way down the west coast, west of and below El Picacho, is a break in the cliffs
— West Anchorage, an indifferent anchorage but sheltered from the north (Figs. 12, 13).
Over the years this was first a camp for sealers near a major rookery of the fur seals
(Anthony 1925, Hanna 1925), then more recently a seasonal fish camp, sometimes with
lobster or abalone fishermen, but now is part of the naval base and known as Campo
Oeste. Rather high cliffs again, continue to the south end (Fig. 13).

Another old rookery and sealing camp was on the low-lying southwest point of the island.
A marine base and weather station (Fig. 14) founded there in December 1946 was for
many years the only permanent settlement on the island; it is now called Campo Sur.
Just to the east is the south-end mesa (Figs. 31, 32), about 100 m high, with cliffs along
the south side facing Melpomene Cove. At the southeast corner of the island is South
Bluff, a rounded butte 310m high (Fig. 32). Just west of South Bluff is the mouth of
Arroyo Melpomene, which rises on the slopes of El Picacho, some 9 km to the north.

Close off the southwest coast is a small islet about 30 m (100 feet) high called Islote
Negro (Fig. 15). Prominent off the south end are two larger islets, Inner Islet and Outer
Islet (Figs. 16, 17). Inner Islet is a rounded dome 227 m (744 feet) high, with vegetation
on top and sheer cliffs below. Outer Islet is a seabound volcanic crater 206 m (677 feet)
high, with sheer cliffs on all sides but the northwest. There the sea has eaten farther into
the side of the crater, leaving only a low rim (Fig. 48), with a rocky slope down to the
water, where on a calm day it is possible to jump ashore onto the rocks and scramble up
into the crater. The floor of the crater is flat, and its sides slope up uniformly at about
45°, on most sides to a jagged rim topping the sheer outer cliffs (Fig. 34). But above on
the east side is a shallow hanging valley, a subsidiary crater, at 190 m (Fig. 35).

REID MORAN

ESS re wT oars Muse oa oh
x Wb

=a.

FIGURE 5. The spring, in upper valley, now the only permanent spring on the island, evidently
fed by fog drip on west side of ridge; 26 April 1958. With the loss of many trees, other springs
of this valley have dried up

FIGURE 6. Lower valley and cliffs of the Lower Circus, looking south to Mt. Augusta, with eastern
fringe of cypress grove beyond; 25 April 1958

GUADALUPE ISLAND FLORA

FIGURE 7. Southern rim of the collapsed caldera (right): looking east down Esparsa Canyon trom
Mt. Augusta, 26 April 1958.

FiGuRE 8. Mouth of Esparsa Canyon, on east side of the island, with Mt. Augusta and eastern
fringe of cypress grove on the skyline; 24 April 1958.

REID MORAN

FIGURE 9. Barracks Canyon, with Northeast Anchorage at the mouth, the skyline with Mt. Augusta
in the middle; 22 April 1958.

FIGURE 10. Northeast Anchorage, with the old barracks, 10 April 1948. The two pines on the
skyline are at the old lookout, at 800 m (see Fig. 42). The shrubs in the canyon mouth are the
recently arrived tree tobacco (Nicotiana glauca, M2838), shunned by the goats and so spreading
and now threatening to take the island.

GUADALUPE ISLAND FLORA

FIGURE 11. North end of the island, with 500-meter cliffs, 17 April 1957. On the skyline are
pines (Pinus radiata var. binata), on the slopes many palms (Brahea edulis) and a few scattered
pines.

Around the corner of the main island, on the east and leeward side, the seacliffs are lower
than on the west. Near the south end are the ruins of another old sealing camp, used
supposedly in the first part of the last century to house Aleutian islanders brought by
Yankee fur sealers to do the dirty work (Hanna 1925).

Farther north are several canyons or arroyos much shallower than Esparsa Canyon and
Barracks Canyon in the northeast.

PLACE NAMES

The map (Fig. 2) shows place names used in this flora. On U. S. Hydrographic Office
Chart 1688, from which I made my early labels for Guadalupe plants, the few long-es-
tablished place names are mostly English, though on Mexican maps many are now trans-
lated or replaced. The newer Mexican names include “Campo Norte” for Northeast An-
chorage, in the old reports simply called “the landing”; “Campo Oeste” for West Anchorage
or Jacks Bay or Campo Tepeyac; “Campo Sur” for the weather station or marine base at
the southwest corner of the island; “Campo Pista” at the air field; and “Campo de Lima”
for what was called the Lobster Camp. Dr. Carl Hubbs (Peterson et al. 1968, fig. 1) im-
provised such names as he needed to record his observations along the coast, such as
Islote Negro, Double Point, Dike Point, Yellow Point. Likewise, I have informally coined
a few names for plant labels, like “Oak Canyon,” “South Oak Canyon,” and “Hemizonia
Cliff.” “Esparsa Canyon,” from Hanna (1925), replaces “North Twin Canyon” of my early
labels; and “South Twin Canyon” becomes “South Esparsa Canyon.” Possibly this name
is a variant of Espaza, used by Palmer as Espaza Bay, apparently for an indentation on

REID MORAN

FIGURES 12 and 13. West Anchorage, a break in the cliffs two-thirds of the way down the west
coast; 17 April 1957. FIGURE 12. View from the west. FIGURE 13. The anchorage from the north,
with Scripps vessel Paolina T, the high cliffs continuing on southward towards Inner Islet.

GUADALUPE ISLAND FLORA

FIGURE 14. The village and weather station at the southwest corner of the island, 14 April 1970
Founded in 1946, it has now grown into Campo Sur.

FiGURE 15. Islote Negro, off the southwest coast, a goatless refugium for south-end plants; 29
April 1958. (See FRONTISPIECE.)

REID MORAN

FIGURE 16. South end of the island from the summit of El Picacho, showing Inner Islet and Outer
Islet off the south coast; 16 April 1970.

FiGureE 17. Outer Islet (left), a seabound volcanic crater, and Inner Islet, south of the main island;
22 August 1981. Outer Islet is a museum of plants rare or extinct on the main island; Inner Islet
is out of reach and tantalizing.

GUADALUPE ISLAND FLORA

the north coast (Watson 1876:119). Esparsa Canyon probably is what Palmer once referred
to as Landrum’s Canon (Watson 1876:119) and Brandegee (1900) called Sparmann’s
Canfion. I have adopted Franceschi’s (1893c) term “Lower Circus” for the lower, inner
are of cliffs remaining from the huge caldera that formed the northeast part of the island,
i.e. the cliffs below and east of the spring where the earliest collectors camped. The Old
Lookout is the site of the two ruined stone huts (Fig. 42) at 800m on the main north
ridge above and west of Northeast Anchorage. Melpomene Cove at the south end has
also been known as Whalers Bay or South Bay. The main arroyo at the south end I have
called Arroyo Melpomene. Inner Islet or Isla de Adentro is also known as Islote Toro,
and Outer Islet or Isla de Afuera was called Zapato as early as Palmer’s day (Safford
MS). However, these names have sometimes been reversed (e.g. by Berzunza 1950:28);
and the present names are unambiguous. Since my place names are a mixture of English
and Spanish, I am taking the easiest course in an English-language account and calling
the island Guadalupe Island rather than Isla Guadalupe or Isla de Guadalupe.

CLIMATE

The climate of Guadalupe Island is largely controlled by weather systems from the north.
In an occasional year a tropical storm from the south, most likely in August through
October, may bring heavy rainfall, strongly skewing the means from usual rainfall figures.

The low south end of the island, with south exposure, is comparatively desertic, but with
frequent fogs as the abundance of lichens suggests. At Campo Sur (6.4 m elevation) near
the southwest corner of the island is a Mexican government weather station, the only
source of weather data for the island. Hastings and Humphrey (1969) gave incomplete
monthly rainfall and temperature records for nearly ten years between 1951 and 1963
from this station. They showed rainfall ranging from 14 mm in 1951 to 88 mm in 1952
and to 693 mm in the exceptional year of 1962 (including 383 mm for March). The mean
annual rainfall was 129.3 mm, distributed as follows: 60.4 mm (44.7 percent) in winter,
61.4 mm (45.5 percent) in spring, 2.7 mm (2 percent) in summer, and 10.5 mm (7.8 percent)
in fall. For 1975 to 1981 Gonzalez (1981) reported a mean annual rainfall of 110.7 mm,
mostly in December, January, and April. Figure 18 shows mean monthly rainfall and
temperatures for 1987 to 1993. For this period rainfall ranged from 11 mm in 1988 to
283 mm in 1991, with a mean of 162.0 mm, distributed as follows: 96.0 mm (59.3 percent)
in winter, 9.2 mm (5.7 percent) in spring, 14.2 mm (8.8 percent) in summer, and 42.6
mm (26.3 percent) in fall. Comparison of the figures for these three data periods shows
the inadequacy of such short records in this arid region intruded by rare tropical storms.
The average for these 20 years of record is an annual rainfall of about 133 mm.

The much higher northern part of the island can be expected to have more rainfall, and
from the vegetation (including Pinus radiata) and from experience (in fact, from several
wet experiences) I am sure it does; but there is no record. As a first estimate for the high
north end, we might take the rainfall for other places where Pinus radiata is native and
where it is successfully grown. Dr. William Libby, a student of this pine, told me it grows
best with about 750 mm of rainfall but under some conditions can survive with as little
as half that much. The effective moisture, of course, is much greater because of fog drip.

Bryant (1887) wrote that fogs were dense and, driven by high winds, swept across the
island and saturated it like rain. He wrote that though rains were not heavy, the runoff
sent small torrents rushing down the arroyos. But in fact, rains can sometimes be very
heavy: I have seen mudflows, rolling rocks, a muddy torrent flowing out of Esparsa
Canyon, and waterfalls over the cliffs. Vegetation is now so sparse that runoff and erosion

REID MORAN

Isla Guadalupe, B.C., México (6 m)
Climate Diagram (Mean Data 1987-1993)

Precipitation (mm) Temperature ( C) Be

=
M A mM JS J A S OF ND
Month

—— Precipitation —— Temperature

Source: Based on data from Comisié6n
Naolonal del Agua, 8ARH, Méxioo (1993)

FIGURE 18. Mean monthly rainfall and temperature for 1987 to 1993 at the south end of Guadalupe
Island. Diagram by Jose Delgadillo.

are rapid. Between 1957 and 1988 I noted the loss of 8 out of 9 oaks in Oak Canyon
through undermining by the torrent that flows after rains, and the last oak is now reported

gone.

Before the prevailing northwest winds, clouds often bank up against the north end of the
island (Fig. 21), and Palmer wrote that fogs prevalent in winter are driven by strong, cold
northwest winds over the crest of the island. At other times clouds tend to dissipate
southward on the usually warmer south slopes. Moisture from the clouds, or fog (depending
on the point of view), condenses on the pine needles and cypress branchlets and drips to
the ground in big drops like rain, making the ground wetter there than elsewhere, enabling
these trees to grow there. The pines grow on the north slope (Figs. 19-21), right up to
the crest of the ridge where the clouds commonly dissipate, but no farther (Fig. 22). Fog
drip from pines and cypress evidently helps feed the springs at the north end. With the
loss of the great groves of the past, other springs of earlier days have dried up, and
continued loss of trees portends further drying and desertification of the island.

Hastings and Humphrey (1969) showed an annual mean temperature at the south end of
17.2°C, the means for different years ranging from 16.6°C to 17.6°C. Mean temperatures
by season were: 15.4°C for winter, 16.1°C for spring, 18.4°C for summer, and 18.8°C
for fall. For 1975 to 1981 Gonzalez (1981) reported a monthly mean temperature of
17.9°C, varying from 16.2°C for April to 20.4°C for October. For 1987 to 1993 the mean
annual temperature was 18.2°C.

GUADALUPE ISLAND FLORA

At the high north end not only is rainfall greater but temperatures clearly are lower,
though again there is no record. Lower temperatures result partly from the elevation but
especially from the frequent fog and cloud cover. Several authors report ice and snow in
winter. Palmer (Watson 1876) wrote that in December 1874 ice an inch thick formed in
the middle of the island, accompanied by two inches of snow and followed by hail and
five days of rain. Bryant (1887) wrote that the nights at that season (Dec—Mar.) were so
cold that ice occasionally formed and frost was of common occurrence. A heavy snowfall
was reported for the winter of 1948-49 (Moran and Lindsay 1950).

A suggestion of a formerly warmer climate comes from Pleistocene fossils interpreted as
of Sangamon Interglacial age (Hubbs and Jehl 1976).

HISTORY

Berzunza (1950) told something of the history of the island and listed scientific expeditions
and conservation efforts for marine mammals. Nelson (1921) summarized scientific work
until 1911 in Baja California, including Guadalupe. For an account of the botanical work,
see Plant Collectors, p. 44.

In October 1602, according to Berzunza, Sebastian Vizcaino sighted a large island which
from its position must have been Guadalupe. First to survey the island and chart it accu-
rately was French Admiral Abel du Petit-Thouars, on the Venus in November 1837 (du
Petit-Thouars 1956). He noted that it ordinarily served as landfall for the galleons returning
from the Philippines, which then went on to sight Cape San Lucas and continued to
Acapulco.

President Lopez Portillo of Mexico visited Guadalupe Island September 28, 1980, and
reportedly ordered studies to find how the island could best be used for the good of the
Mexican people. The Secretaria de Agricultura y Recursos Hidraulicos (SARH), contracted
with Ecoterra of Chihuahua, headed by biologist Dr. Martin H. Gonzalez, for a preliminary
inventory of the natural resources of the island (Gonzalez 1981).

OTHER BIOTA

The high oceanic island of Guadalupe is noted not only for its vascular flora but also for
endemism in birds and land snails. Readers may therefore wonder what is known of these
and other biota. Often that is little or nothing, from a few chance collections or none.

Crum and Miller (1956) summarized collections of bryophytes, at 26 mosses and 8
hepatics, calling the flora distinctly Californian. Crum (1972) added two more mosses.
Bonar (1939) reported three fungi, incidentally collected.

First to make thorough collections of lichens was William A. Weber in 1963, who reached
the eastern side of the island and the south end but not the higher north end; he accounted
for 104 species (Weber 1994). He found the lichen flora, like the vascular flora, more
closely related to that of southern California and its islands than to that of Baja California.
Because the lichens are a pristine remnant of a rich flora now largely gone from coastal
southern California, he proposed that the southern part of the island be designated a lichen
reserve.

No land mammals, reptiles, or amphibians have been found native on the island, but in
addition to the goats, house mice arrived by 1875 and house cats by 1885; and now there
are wild dogs on the northwest slope. The Guadalupe fur seal and the elephant seal, of

Ree
nN

REID MORAN

FIGURE 19. Pine grove (Pinus radiata var. binata) on the west slope of the northeast ridge, with
pines more scattered above; from near North Point, 22 April 1958.

FiGuRE 20. Palm (Brahea edulis) and pines (Pinus radiata var. binata) on the northeast ridge,
seen from the west, 13 April 1948.

GUADALUPE ISLAND FLORA

FiGuRES 21 and 22. Northeast ridge with the pine grove (Pinus radiata var. binata). FIGURE
Clouds banked up against the ridge, 3 May 1967. Moisture from the clouds condenses on the pine
needles and falls like rain, watering the pines, enabling them to grow there. (Oaks in foreground.)
FIGURE 22. Ridge from the south side, 13 April 1948. The pines come just to the crest but no
farther because the clouds commonly dissipate as they blow across to the hotter, drier south side.

21.

REID MORAN

course, are associated with Guadalupe not as endemics but because the island was the
sanctuary where a few somehow escaped the hunters in the last century and later, under
protection of the Mexican government, could recover to spread again; see Anthony (1925)
and Peterson et al. (1968).

From Palmer’s notes and small collection, Ridgway (1876) listed eight species and races
of landbirds, naming all eight as new — though one is now assigned to a mainland form.
Bryant (1887) found only one more to name, the Guadalupe petrel. As Howell and Cade
(1954) put it, the history of the avifauna is a sad one of reduction and extermination
through destruction of habitat by goats, predation by house cats, and, regrettably, some
excess of zeal by collectors. Apparently the Guadalupe rock wren, Guadalupe house finch,
and Guadalupe junco are still doing well. For the latest update on the birds, see Jehl and
Everett (1985). Hubbs and Jehl (1976) reported fossils of two birds from a Pleistocene
deposit interpreted as of Sangamon Interglacial age.

Pilsbry (1927) recognized 14 species and subspecies of ground-living land snails for
Guadalupe Island, eleven of them (79 percent) endemic. He found the fauna most closely
related to that of the southern group of Channel Islands, with 8 species or subspecies
occurring there also or having their closest relatives endemic there. Three others had
nearest relatives in mainland California and two in northern peninsular Baja California.
Most interesting is the genus Binneva, with two species, one endemic to Guadalupe and
the other to Santa Barbara Island and northwest Baja California. As late as 1922 Hanna
(1925) still found many land snails, but feral house mice have preyed on them until
apparently, as he predicted, few are left.

Palmer and later naturalists collected various insects and spiders, some of which were
reported by specialists: Arnaud (1959, 1963), Blaisdell (1925), Horn (1876), Scudder
(1876). Snow (1899) listed collections from the fur seal trip of 1897. With the decline
of the vegetation, presumably much of the original fauna is extinct. There is no compre-
hensive review evaluating the arthropod fauna, but Gonzalez (1981) included a list of 90
species, compiled by Dr. Scott Miller, various of them still unidentified. About 18 are
tentatively considered endemic.

Hubbs and Rechnitzer (1958) suggested that Guadalupe Island was proving to have a
high degree of endemism in its littoral marine as well as its terrestrial biota, and they
mentioned especially fishes. Briggs (1974) wrote that only about four shore fishes could
still be considered well-defined endemics, others named from Guadalupe having since
been found elsewhere. Gonzalez (1981) included a list compiled by Dr. Richard Rosenblatt,
of 125 shore fish species, including five endemics.

Strong and Hanna (1930) listed 87 marine molluscs from Guadalupe Island, mostly dredged
from one area off the south end. They found this fauna very similar to the faunas of
Catalina Island and the Coronado Islands and suggested that many had arrived on drifting
kelp. Chace (1958) expanded the list to 193, still based on rather few collections. He
found 149 (77.2 percent) belonging to the southern California fauna, 34 (17.6 percent)
belonging to the Panamic fauna, and 10 (5.2 percent) endemic. Lindberg et al. (1980)
reported a Pleistocene marine invertebrate fauna of 86 species, mainly molluscs, of San-
gamon interglacial age.

Concerning the marine algae, Dawson (1960) remarked that isolated Guadalupe Island,
unaffected by coastal upwelling, had an extremely interesting flora, with some Californian
species, some tropical Mexican coastal species, some Indo-Pacific insular species, and a
conspicuous group of endemics evidently derived from the Californian flora and showing

GUADALUPE ISLAND FLORA

the effectiveness of isolation and divergent ecology on speciation. Stewart and Stewart
(1984) summarized all records, listing 211 algal species, but mentioned other collections
still unstudied.

The Vascular Plants

NATIVE OR FOREIGN?

In the beginning, of course, Guadalupe Island was a steaming pile of bare volcanic rock,
with no native plants. All plants that grow there must be descendants of immigrants. So
how early must one have come, and by what means, to qualify now as “native”? Perhaps
seeming most native are those that have settled down and become endemic or perhaps
survived there but died out elsewhere. In any case, the distinction is not so clear and
obvious as it might at first appear. “Native” seems best defined a little vaguely as having
arrived long ago without human help.

The most foreign Guadalupe plants, and in some ways the easiest to deal with in the flora
if not in real life, are the 40 or so weeds that have come in the last two centuries, mostly
from Europe. Table | lists them with sourceland and with yeardate of first record on the
island; and in the Catalogue their names are starred. The table includes and notes those
few seen only once or twice and apparently never well established.

So far so good, and this might be the best place to stop. A few floras treat all plants as
equals, not saying whether they are native or foreign; and from some viewpoints, once
the plant is there, it may not matter. But most floras tell what plants are supposed to be
introduced, implying that the rest are native (and seeming to make the distinction with
confidence}—but really leaving most in the category of not supposed to be introduced.
In this flora I have tried to distinguish but have sometimes found it impossible. The
endemic plants and some others are easily called native and the European weeds intro-
duced, but other plants remain doubtful and debatable.

For such plants as Oligomeris linifolia and Plantago ovata, which grow also on other
continents, there is some question whether they are even native to North America, let
alone to the island. For North America, “native” generally means here before the Euro-
peans. This makes a convenient if indefinite timeline, after which European plants have
come in numbers but before which all is murky. But this vague definition may be hard
to apply, as with Oligomeris linifolia. That grows also in the Old World and its relatives
are all there, so clearly it came from there at some time: if that time was before the
arrival of Europeans, we can call it “native” and if after, “introduced”; but who knows
when it came? Furthermore, the Indians clearly moved some plants around before that
arbitrary timeline, and of course some plants undoubtedly have since been moving about
on their own, as they always have. So precisely where an American plant is “native” may
not bear close scrutiny.

Those Guadalupe Island plants clearly native to western North America we are likely to
take also as native to the island, but are they all? In fact, some authors have called
certain ones introduced. In her “List of species widely distributed, probably introduced”
on Guadalupe Island, Eastwood (1929:398) included, with no explanation, not only Euro-
pean weeds but also Aristida adscensionis, Daucus pusillus, Juncus bufonius, Linaria
canadensis, Muhlenbergia microsperma, Myosurus minimus, Parietaria “floridana” (= P.
hespera), Phalaris “intermedia” (= P. caroliniana), and Silene antirrhina. (She probably
intended two separate lists.) Greene (1885:221, 223) thought Eschscholzia californica

REID MORAN

TABLE 1.

Anagallis arvensis

Avena fatua

Avena sativa

Brassica nigra

Bromus tectorum

Erodium cicutarium
Erodium moschatum
Galium aparine

Hordeum murinum ssp. glaucum
Malva parviflora

Silene gallica

Solanum americanum
Sonchus oleraceus
Brassica rapa

Bromus berteroanus
Bromus diandrus
Chenopodium murale
Mesembryanthemum crystallinum
Polypogon monspeliensis
Avena barbata

Centaurea melitensis
Melilotus indica

Sonchus tenerrimus

Vulpia bromoides

Vulpia myuros var. hirsuta
Capsella bursa-pastoris
Lasthenia coronaria
Hordeum murinum ssp. leporinum
Hypochoeris glabra
Mesembryanthemum nodiflorum
Bromus rubens

Medicago polymorpha
Poa annua

Sisymbrium irio

Bromus hordeaceus
Nicotiana glauca

Ruta chalepensis
Cerastium glomeratum
Herniaria cinerea
Phalaris minor

Erodium brachycarpum
Lamarckia aurea
Raphanus sativus

Triticum aestivum
Sisymbrium orientale

Introduced Plants of Guadalupe Island.

Source
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
S. America?
Europe
Europe
S. America
Europe
Europe
Africa
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Western N. America
Europe
Europe
Africa
Europe
Europe
Europe
Europe
Europe
S. America
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe
Europe

Found
1875
1875
1875
1875
1875
1875
1875
1875
1875
1875
1875
1875
1875
1885
1885
1885
1885
1885
1885
1889
1889
1889
1889
1889
1889
1897
1897
1925?
1925
1931
1932
1932
1932
1932
1948
1948
1948
1958
1958
1958
1978
1983
1983
1983
1988

Notes

Not seen since 1885
Not seen since 1885
Not seen again

Not seen again

Reintroduced ca. 1988?

Doubtless planted

Probably not persisting

possibly and Mentzelia micrantha probably “of recent introduction.” Vasey and Rose
(1990:22) included Suaeda “fruticosa” (= S. taxifolia) among plants that “have very re-
cently gained a foothold here.” And Moran (1951:160) thought “Oenothera micrantha”
(= Camissonia robusta) was likely a new arrival.

GUADALUPE ISLAND FLORA

Without explanation, Brandegee (1900:22) called four west American plants introduced:
“Baeria coronaria” (= Lasthenia coronaria), “B. gracilis” (= L. californica), “Orthocarpus
purpurascens” (= Castilleja exserta), and Platystemon californicus. Palmer had found L.
californica abundant and widespread 22 years before, and there is little reason to doubt
that it is native in the present sense. The other three no one has found before or since,
suggesting they might indeed have been waifs that never got well started; but without
knowing Brandegee’s thinking, it is hard to judge. Three more plants likewise have been
found only once, and after the time of the first collectors: Layia platyglossa in 1897,
Stephanomeria diegensis in 1906, and Plagiobothrys acanthocarpus in 1958.

Islands (and especially oceanic islands) are simpler to think about than mainland areas.
But with some Old World islands, inhabited for centuries, distinctions between native and
introduced must be greatly blurred, especially for weedy and useful plants, so that only
the very latest arrivals can be recognized with confidence as foreign. Guadalupe, apparently
never visited by Indians, might at first glance seem clear and easy by comparison. Even
by 1875, however, when Palmer took the first plant records, it not only was losing natives
to the goats but also had gained a dozen European weeds—noted for their almost magical
abilities to arrive but usually needing some help. What else may have had help in arriving?

We may expect those west American plants that have achieved the high but nebulous
status of nativehood on Guadalupe to have come only slowly, over the millenia, by various
improbable means. Though a plant is unlikely to make its first natural arrival within any
short time span, however, conceivably one can come at any time, like this week; on
establishment, it will then automatically seem native so long as no one suspects it has
just come. Of course a new arrival may get a slow start and may remain rare and local
enough for several years to be overlooked by any collectors, so its arrival may not make
the evening news. And what of unnatural arrivals, meaning introductions with human
help? Although sealers and goat hunters of earlier years may have brought little but
weeds, we cannot know what they did bring. But what is more significant, the 1460
Angora goats reportedly brought from California in 1873 and 1874 (Esperanza 1874)
could surely have carried in seeds of west American plants, and probably they did.

There is, in fact, good reason to question whether certain west American plants found on
the island are indeed native. Some plants not found by early botanists undoubtedly were
overlooked because they were rare or were local in remote parts of the island. But some
that were not found then grow now where those early botanists camped and collected.
Can these be long-established plants that have reached new areas on the island with the
drastic changes wrought by the goats or for some other and unknown reason? Can some
be annuals that appear only sporadically and thus happen to evade the unlikely visits of
botanists to their home spot? Or can some perhaps be new arrivals?

For example, Greene found nine western plants in 1885 that Palmer had not noted in
1875, four of them near the springs where Palmer camped for some 15 weeks! Greene,
and later Franceschi, found Eschscholzia californica in that general area but in one place
only and away from the springs, where possibly Palmer had reason to miss it, unlikely
though that may seem. Near the springs, however, Greene found a fine growth of Filago
californica and fine large specimens of Agoseris heterophylla; so how could Palmer have
missed these? Were they just not flowering that year? Or had the goats been hungrier?
Greene raised no question about these two; but he was clearly perplexed that Palmer had
failed to note Dichelostemma pulchellum, which was now exceedingly common all about
the spring and on the slopes below — and large, with leaves three feet long and an inch
wide. Could all four of these plants have failed to flower there the year of Palmer’s visit?

REID MORAN

On the other hand, remember, Greene did say that the eschscholzia might possibly be a
recent introduction. Far from suggesting the same for the dichelostemma, however, he
named it the next year as an insular new species (though that is not recognized today).
So could all four of these plants have failed to flower there for Palmer? Perhaps they did
not flower there that year because they just hadn’t reached the island yet! The four give
cumulative support to this idea.

How is it possible to tell whether western plants recently found on Guadalupe are native?
Some clue might come from their distribution elsewhere, for example whether they grow
on others of the islands and thus show an ability to reach islands. However, their means
of reaching the other islands are as uncertain as their means of reaching Guadalupe: the
islands of southern California have a long history of ranching, with goats, sheep, and
cattle, which could have helped bring in foreign plants! And on the other hand, of course,
they certainly helped destroy natives.

All together, a dozen or more plants remain doubtful and debatable. Very likely several
of them are introduced, but which ones? Taking them as a group, the probability seems
high that at least a few are recent arrivals; but taking them one at a time and without
further evidence, the probability for any one is well below certainty. Having to start
somewhere, and grasping for some solid basis, | assume that very few west American
plants were brought in before the great influx of goats of 1873-74. I therefore tend to
assume that west American plants collected by Palmer in 1875 are native —even though
knowing some may not be. But what of those first found by Greene and later collectors?
If they came from parts of the island where earlier botanists collected without finding
them, then the later the date and thus the more botanists who had nor found them, the
greater the likelihood that they were recent arrivals. Surely some of those must be intro-
duced, and | think probably most; but still, some may be native. In the Catalogue I discuss
the status of each of these species, but always inconclusively. Since they can’t either
definitely be called native or definitely be called introduced, they must go into a separate
category, of Possibly or Probably Introduced (Table 2). This is a satisfying solution to,
or evasion of, a bothersome problem, though raising only slightly lesser questions of
exactly which plants to include in this category and whether to call them possibly or
probably introduced. In the discussion of different plants | use sometimes one word and
sometimes the other.

Table 2 lists Greene’s four additions plus eleven other plants found afterwards where
earlier collectors had failed to find them. A clear example is Camissonia robusta, now
common at Northeast Anchorage, where every early botanist went ashore without seeing
it, but apparently first found in 1948. The table also includes two plants collected only
by Brandegee in 1897, which he called introduced, though without saying why he thought
so or where on the island he found them. In the Catalogue each of these 15 names is
marked with a special symbol (#) to show that the plant is in this uncertain category.

Not included in Table 2 are several other west American plants first found in recent years
but seeming less likely new arrivals. For example, Rhus integrifolia is found only on
Outer Islet, not visited by early collectors. Antirrhinum watsonii is widespread but not
common at the south end, where either Palmer or Howell might have found it; but it
could not have come with goats from California so must have found some other way,
leaving no clue as to when. Likewise, Cuscuta corymbosa could not have come from
California, and it grows where earlier botanists, except possibly Palmer, have not collected.
For Layia platyglossa and Stephanomeria diegoensis, though first found in 1897 and 1906,
there is no record of where on the island; and the collector of the one evidently, and of

pis}

GUADALUPE ISLAND FLORA

TABLE 2. West American plants possibly or probably introduced on Guadalupe Island: found in
later years where earlier collectors had failed to find them.

Found Notes

Agoseris heterophylla 1885 Found only once

Dichelostemma pulchellum 1885 Widespread and fairly common since
Eschscholzia californica 1885 Local, not seen since 1893

Filago californica 1885 Common ever since

Castilleja exserta 1897 Found only once

Platystemon californicus 1897 Found only once

Lupinus bicolor 1898 Now scarce or locally common
Atriplex californica 1931 Still found at one other place
Lepidium nitidum 1932 Since occasional

Plagiobothrys collinus var. californicus 1932 Scarce, or some years locally common
Spergularia marina 1932 Since occasional

Trifolium gracilentum 1932 Fairly common

Camissonia robusta 1948 Locally common since

Pectocarya linearis ssp. ferocula 1957 Fairly common since

Plagiobothrys acanthocarpus 1958 Apparently seen only that year

the other probably, reached places not visited earlier. Perhaps arbitrarily, I have placed
Lasthenia coronaria with the introduced plants (Table 1) instead of with the possibly
introduced because the evidence seems at least a little stronger.

GOATS AND PLANTS

No one knows when goats first reached Guadalupe Island (Fig. 23), but the earliest mention
I have seen is by Xantus for 1859 (Madden 1949). As on so many islands, some ship,
here perhaps a whaler or sealer, no doubt left a few goats to multiply into a fresh meat
supply for later visits. Multiply they have. An estimate for 1870, perhaps far afield, placed
their numbers at 100,000 or more, with a projected increase, if the current rate continued,
to nearly 400,000 in 10 years (Esperanza 1874).

About that time, however, the goat population seems to have reached the carrying capacity
of the island. Fragmentary notes by the first few naturalists show a disastrous decline of
the vegetation around 1875 to 1892, suggesting that within that span the goats may have
reached their greatest numbers or at least their greatest destructiveness. For example, in
1875 Palmer found junipers all over the “middle” of the island, but just ten years later
Greene (1885) could find there only three that were not quite dead. And again, Greene
described the northeast slopes as covered with sage scrub, dominated by Senecio palmeri:
but four years later Palmer found many dead plants of the senecio and few still alive
(Vasey and Rose 1890), and after only three more years Franceschi (1893c) found the
slopes bare and the shrubs surviving only on cliffs, accessible only to winged creatures.
These three reports, fortuitously spaced in a critical time, are a striking record of how
goats destroyed the sage scrub in those seven years. The carrying capacity of the island
must have declined abruptly then and must continue to decline as the island becomes
more desertic.

Exploitation. When goats became abundant on the island, they were at various times
harvested for hides and tallow as well as for meat, and apparently there was more than
one attempt to improve the stock on the island. However, my information is fragmentary
and almost haphazard. In early years, some Americans hunted goats without license from
Mexico. In 1884 the Mexican government stationed 40 soldiers on the island to prevent

REID MORAN

FIGURE 23. Feral goats above Northeast Anchorage, 25 April 1958.

their wholesale slaughter (Greene 1885, Agraz Garcia 1978); and goathunter Stewart
(1965) told of losing 1200 skins in 1899 to a Mexican gunboat.

American hunters reportedly took thousands of hides in various years from the 1860s
through the early 1900s, in one year as many as 30,000 (Esperanza 1874, Safford MS,
Anthony 1925, Stewart 1965). Anthony (1925) wrote that as long ago as 1887, when he
was first on the island, 15,000 hides were taken yearly with no noticeable effect on the
herds. He said that many concessionaires failed at harvesting goats because the selling
price for hides was too low — 75 each in 1899 according to Stewart. The Coast Pilot (U.
S. Hydrographic Office 1951) mentioned that in 1930 there was a goat meat cannery at
Northeast Anchorage, and in 1948 and later some remnants were still to be seen at the
beach. Stewart (1965) gave a vivid account of the 1899 season, from April to late Sep-
tember, on the island with unlawful goat hunters.

Esperanza (1874) told how the California-based Guadalupe Island Company took control
of the island in January 1873, planning to improve the strain of goats, and began to bring
in Angora goats and to kill or castrate the wild bucks; but this venture failed. Without
details, Agraz Garcia (1978) mentioned an introduction of English goats in 1909.

Agraz Garcia (1978) reported two government-sponsored expeditions to the island, ap-
parently both in 1971, taking 20,800 goats and leaving, he said, 239. He did not say what
239 goats were spared, whether they were carefully selected to found a super-race or
were just those that couldn’t be caught. A concessionaire, with about 150 workers from
southern Mexico divided between the two trips, spent 146 days rounding up goats and
preparing the meat, bones, and hides. Meanwhile, Agraz Garcia studied the goats, and
his report tells some of his observations. From an unexplained zoometric study, he thought

GUADALUPE ISLAND FLORA

the goat population descended from four races: Orenburg, Kirghizian, “Pelo Rudo,” and
Nordska; thus he failed to detect the Angora race brought in a century before. With the
goats gone or mostly gone, according to the plan, the Subsecretaria de Recursos Forestales
y de la Fauna was to reforest the island but in fact did not. (That would have been the
best thing possible had they first completely rid the island of goats!) Visiting West An-
chorage and the airstrip in May of 1971, I saw what must have been the relics of the
first of the two expeditions; and there was now a good road, newly built, from West
Anchorage to the airstrip. Since many goats remained alive, the second expedition must
have been still to come.

Agraz Garcia did not even mention the operation of Pescados, Mariscos, y Carnes, S. A.,
in 1970, which I saw in progress in August. Starting about May, they built a runway at
about 550 m elevation at the north base of El Picacho, built a rough jeep road about 16
km north to their camp in the cypress grove overlooking the spring, built corrals south
of the spring, brought in a jeep and trailer in a C-47, and by August started flying out
live goats to Ensenada. According to Oscar Paulin of COTECOCA in 1981, the 1970
operation took out 14,000 goats. Added to Agraz’s figure, this would make some 34,800
goats in about one year. The next years must have been ideal for plants to recover and
for botanists to work on the island.

Goats and the Flora. Plants of an oceanic island like Guadalupe have evolved in the
absence of herbivores like goats, have developed no natural defenses against them, and
so are particularly vulnerable to them — though some like the cholla (Opuntia prolifera)
may have brought their defenses with them. Because annuals are abundant in spring and
the goats can’t eat them all, their very numbers allow some to make seeds to last over
the dry season. Although the goats may gradually eliminate the rarer and more palatable,
many annuals thus can live with goats. In the long dry season when the annuals are past,
goats must still rely on their dried remains, but any perennials become relatively more
vulnerable. In parts of the island most visited by goats, such as near the spring, no shrub
within reach could withstand them; hence shrubs survive there only out of reach on cliffs.

The fact that any trees survive might suggest that trees are secure against goats, but
cypresses still are dying from goats’ eating their bark; and the record shows that vast
numbers have died. In any case, as Anthony wrote, not even a pine, oak, or palm can
look down on a seedling to replace the aged trees now falling; so in the long run, all
trees will be gone. The pines and oaks are dying off most rapidly; but although many
cypress and palms remain, their danger also is imminent.

Since goats are still eating bark from cypress trees and evidently killing some, we may
wonder how any survive or why the effect is so different from place to place. I suppose
the smaller eastern grove, now gone without a trace from the area around the springs,
was more vulnerable to the goats just because they would be crowded there in dry summers.
But it is hard to understand why in the greater part of the vast main cypress grove “only
the fallen trunks, now far gone in decay” should remain in 1885, as described by Greene,
whereas in other parts of that grove thousands still stand today (Fig. 25). Possibly there
were other springs in that area: it is hard to imagine another cause.

Aside from the slaughter of goats by men, the goat population fluctuates with climatic
cycles, decreasing in a series of dry years and recovering after wet. The plant cover
fluctuates in the other direction: when goats are few, the plants increase. In average years
many seedlings from cliff-dwelling plants like Perityle incana appear on the slopes below
but never reach maturity. When goats are few, the seedlings last longer, showing that

REID MORAN

without goats the slopes would soon be covered; and no one knows what long-lost plants
might reappear. In 1963-65, bad years for goats, Lotus grandiflorus, not seen by botanists
since 1885, became common near the mouth of the arroyo at Northeast Anchorage, but
by 1967 it was gone again, as if retreating back to some mountain refuge. At the same
time two seedlings of Lavatera occidentalis, which on the main island can survive only
on cliffs, got well started on the slopes and lasted a couple of years.

Distribution on the island. Goats generally prefer the northern and higher parts of the
island, with better feed and more water. Some do reach even the extreme south end, a
few even in dry summers, but these are only small numbers. In drier seasons, when plants
are parched and the principal spring becomes the only source of fresh water, they cannot
stay away from it for long. In the dry August of 1970 I found the herbage clipped off
to the ground above about 800 m; and the goats had to go far downslope from the spring
to forage and far to the south. Most goats looked weak and forlorn, and one even rolled
over and died before my eyes: many more must have died that year before rains brought
more feed. Again in August 1981, the upper slopes were bare beyond belief.

That goats prefer the north end is clear to see and is borne out by comparing the vegetation
at the north and south ends. In the northern half of the island, aside from the poisonous
weedy Nicotiana glauca and the vicious cholla, no shrubs survive except on cliffs; in the
southern half, and especially at the low south end, some shrubs show little goat damage
at low elevations but show more and more damage northward and upward, until they
disappear at about 400 to 500m. Ambrosia camphorata, for example, dominates some
southern parts of the island but is rare and only on cliffs farther north. However, other
shrubs that might grow at the south end but for the goats are lacking there. The goats
that do go there doubtless change the vegetation by eliminating more palatable plants:
plants found on the islets but not at the south end suggest what the goats must have
eliminated on the main island. Northward, with more goats desperate in dry summers,
the standard of palatability must become lower and lower and finally include almost all
plants.

Now that people live at the south end of the island, probably fewer goats go there than
before. At my first visit in 1948, in the second year of the weather station, only three
people lived there, but it has gradually grown. At first, goat meat was an important item
of diet, and a hunter went out every few days and later every week or so. Now, despite
less hunting, | suppose the size of the base with its human activity must intimidate goats
and must give some protection to the plants at the south end, or at least the southwest
corner. In later years | thought I could see some change in the vegetation that might result
from fewer goats, allowing some plants to re-colonize the main island from one of the
islets. See, for example, Castilleja fruticosa, p. 150.

Effect on the island. Not only are goats extinguishing species of plants, but they also
continue to destroy the vegetative cover of the island, even if not so rapidly as before.
They may somewhat improve the soil with manure — some 300 kg each per year according
to Agraz Garcia (1978). However, their overgrazing prevents buildup of mulch and organic
soil, leading to heating and drying of the soil and to rapid runoff of rainfall and so to
soil erosion (Coblentz 1980). Also, their destruction of trees eliminates the fog drip that
has added greatly to rainfall and so has helped feed the springs. Thus they have been
changing the island from a “naturalist’s paradise” to desert in a way that can never be
completely reversed and corrected.

Hope for the future. Looking at the bare and degraded slopes of Guadalupe Island today,

GUADALUPE ISLAND FLORA

it is hard to visualize the original cover of trees, shrubs, and herbs, and perhaps harder
to believe that any return towards the pristine state is possible. But we can see some hope
whenever a series of dry years lowers the goat population and the plants begin to recover,
and especially when a long-lost plant reappears.

Hope can be seen also in the recent history of a few of the islands of southern California,
such as San Clemente Island, 385 km (240 miles) north of Guadalupe. These islands were
so devastated for two centuries by various combinations and sequences of cattle, sheep,
pigs, goats, and rabbits, that many native plants were lost and replaced by European
weeds, much of the soil was lost, and the original appearance is almost impossible to
imagine. The sheep were removed from San Miguel Island in 1950 and burros more
recently, so no herbivores remain; but trees and shrubs are gone and the soil is gone,
according to Johnson (1980), from some two-thirds of the surface. On Santa Barbara
Island, the rabbits were wiped out by 1981, so no herbivores remain. On Santa Catalina
Island goats and pigs have been removed from some parts of the island and the removal
continues. On the main part of Santa Cruz Island the Nature Conservancy has systemati-
cally removed herbivores, though that good work seems to have flagged in recent years
(Junak et al. 1995), And on San Clemente under the U.S. Navy, biologists have made
great progress in the last 20 years in removing goats and pigs, until all are gone, and in
propagating and replanting native plants. In fact, the plants are now replanting themselves.
However, European weeds also are thriving, not only competing with natives but also
increasing the hazard of fire.

Ing. José Rico C. told me in 1994 that in recent years thousands of goats have been
harvested on Guadalupe but that perhaps 7000 remained. If the goats are ever all removed,
Iam sure that many of the plants surviving on cliffs will spread on the slopes. Of course
the many aggressive European weeds would also prosper and might tend to crowd out
some natives. But with the removal of the goats, the process of desertification could
slowly be reversed.

Because the goats have economic value, some may not want them totally removed, and
there may be political pressure to keep the island as a goat range. There might then be
pressure to introduce other pasture plants, as Agraz Garcia (1978) suggested, which would
be a further calamity for the native flora. Even if the goats remain as a harvestable
resource, however, important areas can still be protected from them by fences and to
some degree will recover. If the cypress grove or some part of it were completely fenced,
for example, with the goats outside, cypress seedlings could renew the grove, and many
other native plants undoubtedly would reappear. A fence was started about 1981 but never
finished. Fencing all the pines and palms would be harder because of the steep and rough
terrain, but even smaller exclosures around some of these trees would allow them to
reproduce locally. But they must be good fences: those built in 1987 to protect some of
the pines, according to Libby (1990), the goats took only three months to breach! Also,
exclosures would be extremely useful in other places, such as the lower valley below the
cliffs of the Lower Circus: many rare plants on these cliffs, some now on the verge of
extinction, might then seed themselves and survive. In fact, it is crucial to the ecology
of the island, and even to the survival of the goats themselves, that fences be built and
maintained, to rebuild the forest and to sustain and extend the springs.

The ideal, however, would be to remove all goats, as well as dogs, cats, and mice, and
to preserve all the native plants, as Mexican ecologists have long urged (e.g. Ravest Santis
1983). In fact, Ing. José Rico C. tells me in 1994 that all concerned governmental agencies
now agree on that objective. It is important to save even those plants not different enough

REID MORAN

to have been named as endemics and important not to bring in other genotypes as re-
placements, even when identified as the same species. But it is particularly important to
save the unique endemics. Some, like the pine, have great economic importance, but all
have a scientific and even spiritual value because of their uniqueness. Preservation is
urgent because the seed source and gene pool are shrinking, bringing more plants closer
to extinction; and furthermore, the island is becoming more desertic and harder to restore.
Guadalupe Island with its unique flora is a Mexican national treasure that urgently needs
protection.

PAST AND PRESENT VEGETATION

Clearly the goats have profoundly affected the vegetation of Guadalupe Island. We have
no account of the original vegetation but only fragmentary notes on how it appeared,
already depleted, to the few early naturalists. The best early notes are by Greene (1885)
and Franceschi (1893c), with some for individual species by Palmer (Watson 1876, Vasey
and Rose 1890, McVaugh 1956, Safford MS). Also, ornithologist Walter Bryant (1887)
and Dudley (1899) gave a few brief but helpful notes. At best there is little physical
description of the vegetation but only a mention or description of species of trees and
shrubs that clearly would be dominant, usually with no more than indirect hints as to
associated plants. Likewise, of course, there is no full list of the original flora. Hence in
telling of vegetation in different parts of the island, I am largely telling of species distri-
butions.

As local climate varies from one part of the island to another, especially with altitude
and exposure, the plants vary with it. The low south end of the island, with south exposure,
is comparatively hot and dry and its plants desertic (Fig. 32) — though with frequent fogs
there as elsewhere, lichens also are prominent, giving a yellowish cast to the scene. Weber
(1994) in fact remarked that lichens often dominate the landscape and “form the principal
vegetation of the island”. The north slope of the island, being higher, with more cloud
cover and fog, more rainfall and fog drip, and less direct sunlight than the south end, is
comparatively cool and moist and has quite different vegetation, with many trees (Fig.
21). In fact, most Guadalupe trees now are in the higher northern third of the island.
Although some plants grow almost throughout the island, most are more restricted, and
some have been found only in one area or even only at one spot. Of course, present
restriction may sometimes be due largely to destruction elsewhere by the goats or to
desertification.

Trees. Much of the higher northern part of the island was originally covered with trees
(Figs. 24, 25), but without human intervention the trees of the island seem doomed with
the present generation. There are old trees and dead trees but no young ones, for no
seedling escapes the goats. All the trees must be well past the century mark, and each
season sees new windfalls. The scanty records show that many old trees over wide areas,
at least cypresses and junipers, perished within a short time span, about when (or just
before) the goats were doing their greatest damage to the sage scrub. At least two small
trees, Hesperelaea palmeri and Heteromeles arbutifolia var. macrocarpa, are gone alto-
gether.

Franceschi (1893a, 1893c) supposed that all the northwest part of the island had been
clothed in dense pine forest (Pinus radiata var. binata), and he wrote that very large fires
had evidently burned there more than once. The remaining pines are most crowded on
the northeast ridge near North Point (Figs. 19-22) and only sparsely scattered on up the
ridge and on the north slope among the palms. There is no record of associated shrubs;

to
oo

GUADALUPE ISLAND FLORA

FIGURE 24. The endemic Guadalupe Island palm (Brahea edulis) on the north slope of the island,
19 May 1971. Although the biggest palm grove is here, other palms grow in canyons scattered
almost the length of the island.

FIGURE 25. Part of the large cypress grove (Cupressus guadalupensis ssp. guadalupensis) on the
upper west slope of the island, looking southwest from Mt. Augusta, 13 February 1957. The grove
stretches far around from the south (left) to the northwest, though most trees are now gone.

REID MORAN
but Ceanothus, Heteromeles, and Ribes, come to mind as possible.

Franceschi (1893c) described the large palm grove (Brahea edulis) on the northwest slope
of the island, where there are still hundreds or even thousands (Fig. 24). Palms also grow
in canyons almost the length of the island, and at least a few grew scattered on the
uplands, as west of the Northeast Oaks. | remember fewer dead ones than with the other
trees, but they too must be dying of old age. Again there is no mention of associated
shrubs.

Greene (1885) wrote that almost the entire plateau of the northern half of the island
appeared to have been covered by cypress (Cupressus guadalupensis ssp. guadalupensis)
but that in 1885 only fallen trunks remained over most of that area; the cause of the
destruction he could not guess. Although the grove on the upper west slope of the island
evidently is much reduced, remarkably, some thousands of trees still stand (Fig. 25),
though their bark is often severely damaged by goats. Possibly their distance from the
spring, the main source of water for goats in dry seasons, gives some slight protection.
Bryant, Greene, and Franceschi all told of a smaller cypress grove on the plateau around
the springs, on the eastern drainage, where in my time only a single dead tree still stood
that now too is gone. Under the cypress of both the larger and smaller groves, Franceschi
(1893c) found a few last survivors of what “must have [been] a thick and general under-
wood” 12 or 15 feet high of Ceanothus crassifolius; but these are all gone. The one shrub
of Arctostaphylos sp. that Greene found under a cypress may have been another last
survivor of that underwood.

Palmer in 1875 saw groves of juniper (Juniperus californica) in ravines and low valleys
all over the “middle” of the island and occasional southward, but only ten years later
Greene found nearly all dead. A few junipers have survived until recently on the southeast
coast, where fewer goats range (Fig. 30), but at my last visit to that part of the island in
1970 they were under strong attack and many dead or dying.

Palmer found the oak (Quercus tomentella) frequent at the north end with the pines and
occasional to the south in canyons on both sides of the island. At the north end a few
still are scattered with the palms and more with the pines, and farther south also very
few remain — probably not 50 now on the whole island (Figs. 26, 61, 62).

Scrub. By the earliest account, sage scrub covered the slopes in the northeast part of the
island, dominated by the beautiful white endemic Senecio palmeri but also with abundant
Lotus argophyllus ssp. ornithopus, Sphaeralcea sulphurea, Calystegia macrostegia ssp.
macrostegia, and Dichelostemma pulchellum (this last probably introduced). Greene thus
described the northeast slope in 1885, but already the goats were rapidly destroying every-
thing in reach. Less than eight years later Franceschi found the slopes bare and the shrubs
surviving only on cliffs. Because Palmer in 1875 found several plants rare on cliffs that
probably grew before on the slopes, including some never seen again, still others may
originally have been part of the scrub. Possible examples are: Coreopsis gigantea, Cros-
sosoma californicum, Daucus pusillus, Eriogonum zapatoense, Euphorbia misera, Galvezia
speciosa, Haplopappus canus, Lavatera occidentalis, Malosma laurina, Perityle incana,
Phacelia phyllomanica, Rhamnus pirifolia, Rhus integrifolia, Scrophularia villosa, So-
lanum wallacei ssp. clokeyi, and Stephanomeria guadalupensis.

Most of this northern upland once covered with scrub is now reduced to goat pasture
(Fig. 27), with mostly annuals. Predominant are European weeds, especially Erodium
cicutarium, E. moschatum, and Hordeum murinum, but also commonly Avena barbata,
Bromus diandrus, B. rubens, Malva parviflora, Silene gallica, Sonchus oleraceus, and

GUADALUPE ISLAND FLORA

FIGURE 26. Small grove of island oak (Quercus tomentella) in hanging tributary of Esparsa Canyon
at 450 m, east of Mt. Augusta; 2 May 1967.

Vulpia myuros var. hirsuta. However, some native annuals also are often common, like
Calandrinia ciliata, Claytonia perfoliata ssp. mexicana, Crassula connata, Crvptantha
foliosa, Filago californica (probably introduced), Gilia nevinii, Parietaria hespera var.
hespera, and Pterostegia drymarioides. Less general but common locally in good years
are such annuals as Amsinckia menziesii, Lepidum lasiocarpum var. latifolium, L. oblongum
var. insulare, Linanthus pygmaeus ssp. pygmaeus, Pholistoma racemosum, and Trifolium
palmeri.

In northern parts of the island the native shrubs now survive only on cliffs, beyond the
reach of goats and sometimes of botanists as well. Some like Galvezia speciosa and the
endemic Perityle incana (Figs. 28, 51), which are abundant and reproducing well, clearly
are natural cliffdwellers and have a secure future. But those that were rare on cliffs before
have now only become rarer: Malosma laurina has only three or four survivors, all very
old shrubs that may even date back to before the goats (Fig. 29). There must have been
many on the slopes before, making millions of seeds and multiplying into probability the
chance of a rare start on a less suitable place like a cliff. But now that so few remain,
making few seeds, the chance of starting in such a place is much smaller. So the number
of such shrubs ts likely to dwindle until the species dies out on the island; and from the
old records, it has been dwindling. Other shrubs rare on northern cliffs, and one or two
perhaps extinct since my first visits, include Ambrosia camphorata (common at south
end), Artemisia californica, Crossosoma californicum, Euphorbia misera (common on
Outer Islet), Galium angulosum, Haplopappus canus, Lavatera occidentalis (common on
Outer Islet), Phacelia phyllomanica, Rhamnus pirifolia, Scrophularia villosa, and Senecio
palmeri. Other shrubs or subshrubs that are more common on the cliffs and are likely to

REID MORAN

FIGURE 27. Guadalupe upland formerly covered with shrubs but now reduced to goat pasture,
with mainly European weeds though with some native annuals; 17 December 1957. In the northern
half of the island, shrubs survive only on cliffs, beyond the reach of goats.

FIGURE 28. The endemic Perityle incana on cliff of the Lower Circus, 27 January 1960. It is
common on cliffs, a natural cliffdweller with a secure future.

GUADALUPE ISLAND FLORA

FIGURES 29 and 30. Shrubs not at home on cliffs are rare, becoming rarer, dying out in the
northern half of the island. FIGURE 29. Laurel sumac or lentisco (Malosma laurina) on cliff of the
Lower Circus, 29 June 1968. Only three or four are left on the island. FIGURE 30. Juniper (Juniperus
californica, M12062) about 4m high and 8m wide hanging from cliff, at 350m on lower east
slope of El Picacho, 4 March 1965. Juniper is nearly extinct on the island

REID MORAN

survive are Calystegia macrostegia ssp. macrostegia, Hemizonia frutescens (north end),
Sphaeralcea sulphurea, and Stephanomeria guadalupensis, as well as some herbaceous
plants.

For the north slope there is no early description of the vegetation aside from the trees.
Possibly common on the north slope was Hemizonia frutescens, now common there on
cliffs. The lower vegetation now is largely annuals, in spring more lush there than in
drier areas to the south. These include such weeds as Capsella bursa-pastoris, Cerastium
glomeratum, Erodium cicutarium, E. moschatum, Galium aparine, Hordeum murinum,
Hypochoeris glabra, Malva parviflora, and Silene gallica. Native annuals that are mostly
towards the north end include Aphanes occidentalis, Githopsis diffusa var. guadalupensis
(rare), Mvosurus minimus var. filiformis (rare), Ranunculus hebecarpus, and Stellaria nitens
(northeast slope to cypress grove). On cliffs at the north end are Hemizonia frutescens,
Poa secunda ssp. secunda (rare), Polypodium scouleri (rare), and Polystichum munitum
(rare); also Baeriopsis guadalupensis, Eschscholzia palmeri, and Lomatium insulare, which
are also at the south end and on the islets. Especially on shaded cliffs and some in talus
at the bases of cliffs, there and southward, are Cheilanthes newberrvi, Dudleva virens
ssp. extima, Jepsonia malvifolia, Pentagramma triangularis, Polyvpodium californicum,
and Triteleia guadalupensis, none of them very common.

Farther south, near the “middle” of the island, the three characteristic shrubs, according
to Palmer, were the abundant Ambrosia camphorata, “giving the country a greenish white
appearance”; Atriplex barclayana, “scarcely half so abundant’; and Artemisia californica,
less frequent but still “giving character to the vegetation” (Watson 1876). Below 500 m
in the southern half of the island the ambrosia 1s still abundant over wide areas, on the
top and on the southeast slope of the island, still coloring the landscape; and the atriplex
is still abundant in other places on top and on the south end mesa (Fig. 32). The artemisia
is gone except possibly for a few survivors on northern cliffs. Herbaceous plants common
in the southern half of the island include Amblyopappus pusillus, Aphanisma_ blitoides,
Oligomeris linifolia, Parietaria hespera var. hespera, Plantago ovata, and Pterostegia
drymarioides, and common more locally Calandrinia maritima, Crassula connata, Cryp-
tantha foliosa, C. maritima, Eschscholzia palmeri (flowering first year), Lupinus niveus,
Perityle emorvi, Sphaeralcea palmeri, and Trifolium palmeri. Common weeds include
Erodium moschatum, Hordeum murinum, Mesembryanthemum crystallinum, M. nodi-
florum, and Sonchus oleraceus.

For the far south end we have no description of the original vegetation beyond a first list
of the plants, for 1889. However, the vegetation of the islets, undisturbed by goats, gives
some clues (Figs. 34, 35). At the south end, with the more palatable plants doubtless
eliminated, several shrubs and subshrubs still remain common, notably Atriplex barclayana
(Fig. 32), Hemizonia greeneana ssp. greeneana (Figs. 33, 50), H. palmeri (Plate 1D),
Lycium californicum, Mirabilis californica, and Sphaeralcea palmeri (Figs. 31, 69). Such
plants as Castilleja fruticosa and Lomatium insulare (Figs. 31, 77), now found near the
west end of the mesa above Campo Sur, may be new colonists from the islets, given
some protection by the nearness of the village. Likewise, Baeriopsis guadalupensis, now
seen on the steep north slope below the road where it climbs onto the mesa, may have
increased with protection.

The Islets. The offshore islets are a small but fascinating and invaluable sample of southern
Guadalupe Island undisturbed. They are of great botanical interest because their vegetation
must show much how the original south-end vegetation looked before the goats. They
are refugia where rare plants survive, some perhaps to return someday to the main island.

GUADALUPE ISLAND FLORA

FIGURES 31 and 32. At the dry south end, where few goats range so far from water, perennials
and even shrubs survive, even within reach; 18 December 1957. FIGURE 31. Rocky south-end mesa
Just above Campo Sur, with Lomatium insulare, Atriplex barclayana, and Sphaeralcea palmeri.
FIGURE 32. Sparse low cover of Atriplex barclayana on the south-end mesa; looking south towards

South Bluff.

REID MORAN

FIGURE 33. Sparse growth of Hemizonia greeneana ssp. greeneana (with Atriplex barclayana)
on a cinder cone at the south end, 20 August 1970.

Islote Negro. \slote Negro (Fig. 15) is small and rather barren, with few habitats and a
small flora. | found only 27 species of vascular plants:

Amblyopappus pusillus

Antirrhinum watsonii

Aphanisma blitoides

Atriplex barclayana

Baeriopsis guadalupensis (Guadalupe endemic)
*Chenopodium murale

Cistanthe guadalupensis (Guadalupe endemic)
Cryptantha maritima

Dudleya guadalupensis (Guadalupe endemic)
Eschscholzia palmeri (Guadalupe endemic)
Galvezia speciosa (Insular endemic)
Hemizonia greeneana ssp. greeneana (Guadalupe endemic)
Lycium californicum

Mammillaria blossfeldiana var. shurliana
*Mesembryanthemum crystallinum
*Mesembryanthemum nodiflorum

Mirabilis californica

Oligomeris linifolia

Parietaria hespera var. hespera

Perityle emoryi

Perityle incana_ (Guadalupe endemic)
Phacelia floribunda (Insular endemic)

GUADALUPE ISLAND FLORA

FIGURES 34 and 35. Vegetation of Outer Islet, undisturbed and much as vegetation must have
looked at the south end of the main island before the goats; 21 June 1968. FiGURE 34. Floor of
the crater, with Cistanthe guadalupensis (thick stems, leafless in summer), Lyciwm californi-cum.
FiGURE 35. Shallow upper valley at 190 m, looking north to the main island, with Lavatera lindsayi
(right foreground), Stephanomeria guadalupensis (white), Atriplex barclayana, Coreopsis gigantea.

REID MORAN

Plantago ovata
*Sonchus oleraceus

Spergularia macrotheca

Sphaeralcea palmeri (Guadalupe endemic)
Trifolium palmeri (Insular endemic)

All are Dicots. Four (marked with *) are Old World annual weeds, though of these only
Mesembryanthemum crystallinum is common. Seven are Guadalupe Island endemics, and
three more are insular endemics, found only on Guadalupe and others of the islands but
not on the mainland. Various of these plants are common at the south end of the main
island, such as Amb/vopappus pusillus, Lycium californicum, Mirabilis californica, Parie-
taria hespera var. hespera, and the endemic Hemizonia greeneana ssp. greeneana and
Sphaeralcea palmeri (Fig. 69). But Islote Negro is strikingly different from even nearby
and closely comparable areas on the main island because common here, even in flat
places, are plants rare on the main island except on cliffs. Perityle incana and Galvezia
speciosa, both common here, are found on the main island mainly in the northern part,
where they grow only on cliffs (Figs. 28, 51). Mammillaria blossfeldiana var. shurliana,
though widespread at low elevations on the main island, is nowhere common there but
is abundant here (Plate 1A). Baeriopsis guadalupensis (Fig. 47), scarcely to be found on
the main island at all, also is abundant here. Likewise, Cistanthe guadalupensis, seldom
seen on the main island except rarely on seacliffs, is abundant and luxuriant here (FRON-
TISPIECE, Figs. 73, 74). Dudleva guadalupensis has never been found on the main island
but 1s common on the islets (Figs. 57, 58).

Outer Islet. Like Islote Negro, Outer Islet is a remarkable and revealing sample of south-
end Guadalupe Island undisturbed; but this islet is larger and farther offshore (Fig. 17),
and its flora is larger and truly unique. My four visits to Outer Islet, one overnight but
all of them too short, stand out as highlights in my botanical memories. | found 38 species
of vascular plants:

Aphanisma blitoides

Atriplex barclayana

Atriplex californica

Baeriopsis guadalupensis (Guadalupe endemic)

Calystegia macrostegia ssp. macrostegia (Insular endemic)
Castilleja fruticosa (Guadalupe endemic)

Cistanthe guadalupensis (Guadalupe endemic)

Coreopsis gigantea

Crassula connata

Crossosoma californicum (Insular endemic)

Cryptantha foliosa (Guadalupe endemic)

Descurainea pinnata ssp. menziesii

Dichelostemma pulchellum

Dudleya guadalupensis (Guadalupe endemic, islets only)
Eriogonum zapatoense (Guadalupe endemic, Outer Islet only)
Erysimum moranii (Guadalupe endemic, Outer Islet only)
Eschscholzia palmeri (Guadalupe endemic)

Euphorbia misera

Galvezia speciosa (Insular endemic)

Hemizonia greeneana ssp. greeneana (Guadalupe endemic)
Hutchinsia procumbens

GUADALUPE ISLAND FLORA

Lavatera lindsayi (Guadalupe endemic, islets only)
Lavatera occidentalis (Insular endemic)

Lomatium insulare (Insular endemic)

Lotus argophyllus ssp. ornithopus (Insular endemic)
Lycium californicum

Mammillaria blossfeldiana var. shurliana
*Mesembrvanthemum crystallinum

Mirabilis californica

Oligomeris linifolia

Parietaria hespera var. hespera

Perityle emoryi

Perityle incana (Guadalupe endemic)

Phacelia floribunda (Insular endemic)

Pholistoma racemosum

Rhus integrifolia (Not on main island, but on mainland)
Spergularia macrotheca

Sphaeralcea palmeri (Guadalupe endemic)

All but one are Dicots and all but one native. Ten are native annuals, all but two of these
common on the main island. Twelve are Guadalupe Island endemics, and seven more are
insular endemics. Here again Baeriopsis guadalupensis, Cistanthe guadalupensis (Fig.
34), Galvezia speciosa, and Peritvle incana, are common and not confined to cliffs. Com-
mon too are the endemic Castilleja fruticosa and Hemizonia greeneana ssp. greeneana,
otherwise local but common at the south end of the main island, and the insular Lomatium
insulare, local at the south end and on the northwestern cliffs. Also common are Euphorbia
misera and the endemic Stephanomeria guadalupensis (Figs. 53-55), uncommon on the
main island and largely confined there to cliffs, and the insular Lavatera occidentalis
(Figs. 67, 68), rare on the main island, on northern cliffs. The insular Crossosoma cali-
fornicum, rare on northern cliffs is rare here too: I saw only one shrub and only on my
first visit. Lastly, common here, and at least visible on Inner Islet and perhaps common
on top, is Coreopsis gigantea (Fig. 48), not seen on the main island since Palmer found
two plants in the crevices of high rocks in 1875.

Finally, Outer Islet has five plants (marked “islets only” or “Outer Islet only”) never
found on the main island: Rhus integrifolia (Fig. 48), also widespread on the mainland,
and the four Guadalupe Island endemics Dudleya guadalupensis (also on Islote Negro
and Middle Rock), Eriogonum zapatoense (Fig. 72), Erysimum moranii, and Lavatera
lindsayi (also on Middle Rock) (Figs. 65, 66). Although no one ever saw these five on
the main island, they must have grown there, like the coreopsis, until wiped out by goats.
The five species make up over 13 percent of the small native flora of the islet. This is
the only faint suggestion we have of how many plants, and how many endemics, the
goats must have destroyed on Guadalupe Island before any one could see them. That is
a lesson of the islets.

EXTINCT PLANTS AND NEARLY EXTINCT

By the time Dr. Palmer landed on Guadalupe Island in 1875, the goats had made serious
inroads and many plants that might have grown on slopes were found only on cliffs; and
eight that he found have not been seen again. Also, many aggressive foreign weeds were
crowding native plants, perhaps contributing to their extinction. However, weeds are few
in some habitats, as on volcanic cliffs, and there are few on the islets.

)

REID MORAN

Of some 156 native, and some doubtfully native, plants of Guadalupe Island, 26 have not
been found since about 1900 (Table 3). Most of these almost certainly are extinct on the
island, though for some it is hard to be sure. Though considering in the Catalogue the
likelihood of survival for each, and still cherishing hope for a few, I mark each with a
dagger (+) in the Catalogue and officially list them all in Table 3 as probably extinct. As
shown in Table |, eight of the weeds likewise have not been seen since 1900 and probably
are extinct, apparently never having gotten well started.

TABLE 3. Native or possibly native plants probably extinct on Guadalupe Island: mostly not seen
since 1900.

Last Seen Notes

Ceanothus cuneatus 1875 Only three, nearly dead
Dissanthelium californicum 1875 Found once each on Santa Catalina,
Guadalupe, and San Clemente Islands
Hesperelaea palmeri 1875 Only three still living then; endemic
Micropus californicus 1875
Planta species 1875 Only two, sterile; doubtless endemic
Pogogyne tenuiflora 1875 “Very rare” (only one’?); endemic
Ribes sanguineum 1875 Only two
Silene antirrhina 1875 Occurring sparingly
Agoseris heterophylla 1885 Only collection; introduced?
Allophyllum gilioides 1885 Abundant in 1875
Arctostaphylos sp. 1885 Only collection, only one seedling
Epilobium foliosum 1885 First in 1875; few in 1885
Malacothrix clevelandti 1885 Abundant in 1875
Phoradendron densum 1885 First in 1875
Satureja palmeri 1885 Abundant in 1875; endemic
Ceanothus crassifolius 1893 First in 1875; mostly dead in 1893
Eriophyllum lanatum var.
grandiflorum 1893 First in 1875; rare then
Eschscholzia californica 1893 First in 1885; one place only;
probably introduced
Hesperocnide tenella 1893 First in 1875; “very common
everywhere” in 1893
Castilleja exserta 1897 Only collection; doubtfully native
Lavia platvglossa 1897 Only collections
Platystemon californicus 1897 Only collection; doubtfully native
Castilleja guadalupensis 1898 Rare in 1875; endemic
Nicotiana attenuata
(N. petuniaeflora) 1898 First in 1875, in a few places
Stephanomeria diegensis 1906 Only collection
Heteromeles arbutifolia
var. macrocarpa 1958 First in 1957; only one tree

Many other plants are now rare and threatened with extinction — or some of them perhaps
already gone. All trees are threatened, though we may hope that help will come before
too late. Also threatened are seven shrubs that survive only on northern cliffs and that
aren’t known to be reproducing: Artemisia californica, Crossosoma californicum, Galium
angulosum, Haplopappus canus, Malosma laurina, Rhamnus pirifolia, and Scrophularia
villosa. Were it not also on Outer Islet, Lavatera occidentalis would also be on this list;
but otherwise-rare plants that grow in good numbers on the islets seem safe enough for
the present. Plants that | have seen only rarely or not at all and that seem most threatened,

GUADALUPE ISLAND FLORA

some of which may be gone by now, include: Camissonia guadalupensis ssp. guadalu-
pensis, Descurainia pinnata ssp. menziesii, Githopsis diffusa var. guadalupensis, Myosurus
minimus vat. filiformis, Matricaria matricarioides, and Plagiobothrys acanthocarpus. Oth-
ers that, so far as known, are rare or very local include Ariplex californica, Castilleja
attenuata, Chamaesyce pondii, Cistanthe maritima, Cuscuta corymbosa, Daucus pusillus,
Gnaphalium stramineum, Hutchinsia procumbens, Lotus grandiflorus, Melica imperfecta,
Mimulus latifolius, Phacelia phyllomanica, Poa secunda ssp. secunda, Polystichum mu-
nitum, Senecio palmeri, Solanum wallacei ssp. clokeyi, Stipa lepida, Thysanocarpus erec-
tus. Others that are scarcely more widespread include Dodecatheon clevelandii ssp. insu-
laris, Gnaphalium bicolor, Harpagonella’ palmeri, Linaria canadensis, Lupinus
guadalupensis, Lycium fremontii, Solanum douglasii, and Stebbinsoseris heterocarpa.

Many cliffs on the island are inaccessible and unexplored and may still harbor plants not
seen for many years or even some never found. In crowded days ashore while working
from a ship, there was too little time to reach some of these cliffs and work them thor-
oughly. Now that there are roads and permanent camps on the island, a botanist based
on shore and having time and transportation might find it rewarding to work some of
them more thoroughly. On my first dozen trips to the island, the list of perhaps-extinct
species was cut down almost every trip.

Although some cliffs can be scanned at short range from below, others cannot be. The
often fog-drenched sheer cliffs at the north end of the island seem an especially likely
place for some rare plants to persist, and in fact they have yielded a few to some rather
precarious fishing from above. With time and patience these and other cliffs could prof-
itably be explored further, even if never completely. There are, for example, very promising
cliffs at the brink northwest of Mt. Augusta and others on the southwest coast, and in
both places it is not too hard or too hazardous to go partway down from the top and then
work upward. In some places two or more people could use ropes and other climbing
aids to advantage. Travelling alone, I found good use for a tree trimmer with a 24-foot
aluminum pole in six four-foot sections — though this often seemed just too short, and |
would recommend another 7! inches.

The top of Inner Islet always seemed to me the most interesting but unattainable objective.

COMPOSITION AND RELATIONSHIPS OF THE FLORA

Rzedowski (1978, fig. 65) recognized 17 floristic provinces for Mexico, including a
“Provincia de California,” nearly equivalent to the California Floristic Province, which
extends into northwest Baja California, and a “Provincia de Baja California,” for the rest
of the peninsula. However, he made Guadalupe Island a separate “Provincia de la Isla
Guadalupe,” mentioning the relationship to the flora of Alta California and especially to
that of the islands off the coast but stressing the degree of endemism.

No flora is complete without a neat summary of how many plants of each major group
are native and how many introduced, where else they grow, and, especially with an island
flora, what percentage are endemic. Raven (1963, 1967) discussed thoroughly and in
detail the floristics of the California islands plus Guadalupe. There is little to add except
for changes in numbers with newer information. Wallace (1985) checked herbarium records
and made a very useful summary of these island floras, which gave us new numbers, as
the present flora does for Guadalupe Island. Notably, a reconsideration of what plants are
native to Guadalupe slightly reduces the size of the accepted native flora.

The total known vascular flora of Guadalupe Island is 216 species (and subspecies and

REID MORAN

varieties), including plants that once grew on the island but are now extinct there. These
include some weeds that may have grown there only briefly and never become well
established. It does not include the various species discussed under Doubtful and Excluded
Species (p. 168)

All together, then, 216 species of seed plants and ferns have been found on Guadalupe
Island — but not all at one time. Thirty or more species that once grew there are extinct
on the island (Table 3; also Table 1). Meanwhile, as if to fill the void, new weeds keep
coming. Of the 216 listed species, about 45 are relative newcomers and mostly weeds,
mostly from Europe (Table 1); and 15 west American plants not found by the first collectors
also are probably or possibly introduced (Table 2). Thus under a strict yet somewhat hazy
definition, only 156 species remain listed as native plants.

There is no knowing how many species were destroyed by goats before they could be
found by botanists, so the size of the Guadalupe flora before goats is unknown. However,
the number surviving only on goatless islets is suggestive: of the small flora of Outer
Islet, 13 percent (five species) appear to be extinct on the main island. Assuming as a
rough estimate that 13 percent of the Guadalupe flora went extinct without being found,
the flora before goats would be 179 species.

For each of the eight Channel Islands plus Guadalupe and Cedros, Raven (1967) discussed
the size of the native vascular flora in relation to the size of the island. He concluded
that in the Channel Islands group an island the size of Guadalupe might have a flora of
over 400 species. The much smaller size of the actual Guadalupe flora may be related to
the greater distance of the island from the mainland, perhaps making it harder for immi-
gration to keep up with extinction. Since the size of the Guadalupe flora more closely
approaches that of Cedros, a somewhat larger island close to the mainland but at about
the same latitude, the small size of the Guadalupe flora may be related also to ecological
differences of these islands from the Channel Islands much farther north, such as drier
climate (Moran 1967).

Of the 156 native plants, 34 species and subspecies, making 21.8 percent of the native
flora, are endemic to Guadalupe Island. Two of these species belong to monotypic genera,
Baeriopsis J.T. Howell and Hesperelaea A. Gray, which thus are endemic genera. The
unidentified sterile plant (Planta sp.) of Palmer’s collection, which several keen botanists
failed to recognize even to family, was surely an endemic species and possibly would be
another endemic genus. Also, Nesothamnus Rydb. (for Peritvle incana) is a very distinct
endemic, standing well apart from other species placed in Perity/e; and it could arguably
be kept as an endemic genus, as did J. T. Howell (1942). And finally, E. L. Greene proposed
for Eschscholzia palmeri (and his P. frutescens) an endemic genus Petromecon, which
Fedde (1909, 1936) in his world monographs of the family also kept but later students
of the family have not. We can’t even guess how many endemics were already extinct
by Palmer’s day. The endemics are as follows (+ = extinct):

Baeriopsis guadalupensis

Brahea edulis

Camissonia guadalupensis ssp. guadalupensis
Castilleja fruticosa
+ Castilleja guadalupensis

Cistanthe guadalupensis

Cryptantha foliosa

Cupressus guadalupensis ssp. guadalupensis

GUADALUPE ISLAND FLORA

Dudleya guadalupensis
Dudleya virens ssp. extima
Eriogonum zapatoense
Erysimum moranii
Eschscholzia elegans
Eschscholzia palmeri

Galium angulosum

Githopsis diffusa var. guadalupensis
Hemizonia frutescens
Hemizonia greeneana ssp. greeneana
Hemizonia palmeri
+Hesperelaea palmeri
Lavatera lindsayi

Lupinus niveus

Marah guadalupensis ?
Perityle incana

Phacelia phyllomanica

Pinus radiata var. binata
+Planta sp.
+Pogogyne tenuiflora
+Satureja palmeri

Senecio palmeri
Sphaeralacea palmeri
Sphaeralcea sulphurea
Stephanomeria guadalupensis
Triteleia guadalupensis

The other 122 native plants, of course, are wider ranging, or non-endemic. Of these only
eight species, or 5 percent of the native flora, grow on the peninsula of Baja California
and sometimes beyond but not north into Alta California. These are mainly desert plants
and grow mostly at the desertic south end of the island. They are:

Ambrosia camphorata

Antirrhinum watsonii

Atriplex barclayana

Chamaesyce pondii

Cuscuta corymbosa

Lepidium lasiocarpum var. latifolium
Mammillaria blossfeldiana var. shurliana
Thysanocarpus erectus

Thus the majority of the non-endemic plants, some 114 species or 73.1 percent of the
native flora, grow also to the north, in Alta California, and especially on the Channel
Islands off southern California. In fact, 101 of them, or nearly 65 percent of the native
flora, grow also on some of those islands and (a few) on islands off northwest Baja
California. It may be noted that winds and currents reaching Guadalupe are prevailingly
from the north and northwest, and seeds are perhaps most likely to reach Guadalupe from
this direction; and these other islands of course have a somewhat similar maritime or
insular climate. In fact, 19 plants, or 12.2 percent of the native flora, are insular endemics,
growing also on one or more of the islands off southern California and northwest Baja
California but not on the mainland. They are as follows (the other islands being AN,

REID MORAN

Anacapa; BA, Santa Barbara; Ben, San Benitos; CA, Santa Catalina; Ced, Cedros; CL,
San Clemente; Cor, Coronados; CR, Santa Cruz; Mar, San Martin; MI, San Miguel; Nat,
Natividad; NI, San Nicolas; RO, Santa Rosa; SPH, San Pedro Hill; Tod, Todos Santos

—see Fig. 1):

Calystegia macrostegia ssp. macrostegia AN, CA, CR, Mar, MI, RO, SM
Crossosoma californica CA, CL, SPH
+Dissanthelium californicum CA, CL

Eschscholzia ramosa BA, Ben, CA, Ced, CL, Cor, CR, Mar, Nat, NI, RO, Tod
Galvezia speciosa BA?, CA, CL

Gilia nevinii AN, BA, CA, CL, CR, NI, RO

Haplopappus canus CL

Jepsonia malvifolia CA, CL, CR, NI, RO

Lavatera occidentalis — Cor

Linanthus pygmaeus ssp. pygmaeus CL

Lomatium insulare CL, NI

Lupinus guadalupensis CL

Mimulus latifolius CR

Phacelia floribunda CL

Quercus tomentella AN, CA, CL, CR, RO

Rhamnus pirifolia CA, CL, CR, MI, RO

Scrophularia villosa CA, CL, RO

Solanum wallacei ssp. clokeyi CR

Trifolium palmeri BA, CA, CL, NI

Of the Guadalupe endemics, six have their closest relatives on other islands: Camissonia
guadalupensis ssp. guadalupensis, Dudleva virens ssp. extima, Eriogonum zapatoense,
Haplopappus canus, Lavatera lindsayi, and Solanum wallacei ssp. clokeyi.

Clearly, the relationship of the flora of Guadalupe Island is mainly to the north, with the
flora of Alta California and especially the flora of its islands. Therefore, as Raven and
Axelrod (1978) explained, and indeed as Howell (1931) had shown in slightly different
terms before, Guadalupe Island can fairly be called a southern outher of the California
Floristic Province.

Plant Collectors

My aim here is to tell about the plant collectors, especially the early ones, and their trips
to the island: what is known or can be deduced about where they collected, what new
plants they found, where their collections are reported, where the specimens are filed,
and any sources of pertinent information — in short, to bring together any facts that will
help interpret their collections. Herbarium abbreviations follow Holmgren et al. (1990).
Nelson (1921:140-147) concisely summarized scientific explorations in Baja California
up to 1911, including the work of most early collectors considered here.

In those early days when Guadalupe Island was remote, wild, and unknown, every naturalist
visit was both a major undertaking and a botanical landmark. As the world has shrunk,
visits have become easier and more frequent. And the island itself has changed, less now
because of the goats: with colonists and even roads, it is no longer the wild place I first
knew in 1948, even as that was no longer the more nearly pristine island Xantus saw but
failed to describe in 1859. With the coming of cruise ships, when almost anyone can
become a world traveler for a week, it is impossible to keep track of everyone who might

GUADALUPE ISLAND FLORA
have brought back a few plants — and perhaps not worth trying.

JANOS XANTUS. Although the record of collectors rightly begins with Edward Palmer, it
is not quite complete without mention of Janos (or John) Xantus de Vesey, credited by
Madden (1949:104) withthe first recorded scientific exploration of Guadalupe Island”.
Xantus made important early natural history collections in California and Baja California;
but at the same time he was becoming a sort of Baron Munchausen of his native Hungary
by writing extravagant tales of imaginary exploits in the American West, with text and
illustrations largely lifted from U. S. Government reports of the military explorers (Madden
1949, 1977; Zwinger 1986). In a letter from Xantus to Secretary Spencer Baird of the
Smithsonian Institution, quoted by both Madden and Zwinger, he tells of going ashore
on Guadalupe Island for several hours on March 17, 1859. Although his time ashore was
short, we could wish for collections from that time 16 years before Edward Palmer’s first
visit; but Xantus mentioned none and apparently made none. Or at least we might hope
for a naturalist’s description of the island of that day before the goats peaked (he did
mention about 10,000 goats); but he wrote, for example, that it was “covered entirely
with gigantic cactacea of innumerable genera & species,” with specific mention of Cereus
giganteus, where others have found but two small cacti. He also mentioned “a dark colored
Lepus,” where others have found no native land mammals. Very likely his inspiration
here was the black Lepus insularis of Isla Espiritu Santo, just north of La Paz; for he
wrote the letter from La Paz. Such fantasy impugns the whole of his brief account, though
that includes enough truth to suggest that he did see the island before giving free rein to
his imagination.

EDWARD PALMER. Dr. Edward Palmer, a major plant explorer in the American West and
in northern Mexico from 1853 to 1910, was the first naturalist to collect on Guadalupe
Island. McVaugh (1956) gave a thorough and invaluable account of his life, itineraries,
and plant collections; and an unpublished biography by William E. Safford adds further
details, with quotations from Palmer’s notes (Safford MS). Blake (1961) also quoted from
Palmer’s notes and gave further facts.

On reading an account of the goats on Guadalupe Island (Esperanza 1874), Palmer was
inspired to collect plants there before they were all destroyed. On his first trip, in 1875,
he intended to stay six weeks but was on the island from February | to May 16. With
him as assistant was twelve-year-old Harry Bye Stewart (1862—1922) of San Diego. Palmer
lived in a dugout with a roof of earth near the springs on the plateau about a fifth of the
way from the north end. He did not have a boat, so could travel only overland. At first
he collected in the northern part of the island, as far as he could go in a day in every
direction. After the more accessible places, he “invaded the canyons,” often collecting
from the cliffs with a cord and noose at the end of a long pole,*which caused the men
to laugh when they saw me go out to ‘lasso plants’, [but mJany of the species I could
have gotten in no other way.” However, he could scarcely have collected in lower Esparsa
Canyon since he did not find several species which have survived there until recently,
such as Euphorbia misera, Scrophularia villosa, Stephanomeria guadalupensis, Triteleia
guadalupensis, and several others that he found only on the higher cliffs. During the last
part of his stay, when supplies had run out, he was sick and feeble from an enforced diet
of mostly goat meat, losing some 35 pounds, and could do little collecting.

Palmer mentioned visiting the extreme northern end of the island in early May. It appears
that he got south to somewhere near Jacks Bay (West Anchorage), about two-thirds down
the island — perhaps with the help of Jack the Burro; but judging from the south-end
plants he did not find on this trip, he clearly did not reach the far south end. For plant

REID MORAN

distributions he often gave “south end”: but once it was “south end, especially about
Jack’s Bay,” and for Lycium californicum, which does grow at Jacks Bay, he said “extreme
south end.” And since he cited the cypress and various plants of high cliffs as in the
“middle of the island,’ this phrase often seems to mean well up in the northern half of
the island. In quoting Palmer’s distribution notes for his first trip, therefore, | put “middle”
and “south end” in quotation marks.

According to MeVaugh (1956:230, 354, 362), Palmer on his first trip collected some 1200
specimens (vascular plants and cryptogams) under 140 numbers, which were distributed
in Il sets, the first set at GH and some of the others traceable. Watson (1876) gave an
account of the island based on Palmer’s notes and listed his vascular plant collections
under 119 newly assigned numbers, with 15 mostly rare plants unnumbered. Gray (1876b)
and Watson (1876:124-146) proposed two new genera (Harpagonella, Hesperelaea),
twenty-two new species, and two new varieties from Palmer’s collections; and several
other species were named later. Others reported Palmer’s important collections of bryo-
phytes, lichens, birds, land shells, and insects (McVaugh 1956:230). All together, Palmer
found 119 species of vascular plants, about 12 introduced and 107 native. The plants were
succumbing fast to the cresting goat population, and he was just in time to find eight
natives never found on the island again: Ceanothus cuneatus, Dissanthelium californicum,
Hesperelaea palmeri, Micropus californicus, Planta sp., Pogogyne tenuiflora, Ribes san-
guineum, and Silene antirrhina. We can only wonder what Xantus might have found 16
years earlier. (Phoradendron densum was not found by Greene in 1885 and apparently
was not collected again; but Bryant saw it growing on cypress in 1885.)

For his second visit, Palmer chartered a schooner and was at the island March 27 to April
3, 1889, sleeping aboard. On March 29-30 he collected Nos. 656-58 and 858-905 at the
far south end. On April -3 he got Nos. 667-75 and 831-57 in the northeast part (McVaugh
1956:230), collecting in the canyon above the landing but not climbing to the springs. He
reported that rain had been frequent this year, and his notes suggest that various of the
plants were indeed lush and abundant. Vasey and Rose (1890) reported his collections
(US), listing 64 species, ten of them new for the island. Rose named four new species,
all from the south end: Eschscholzia palmeri, Hemizonia greeneana, H. palmeri, and
Sphaeralcea palmeri, all endemic. The other additions were the native Chamaesyce pondii,
Spergularia macrotheca, and Suaeda taxifolia, and the introduced Centaurea melitensis,
Melilotus indica, and Sonchus tenerrimus. All together, on his two trips, Palmer collected
a total of 135 vascular plants, including 15 introduced species and 120 native — out of the
156 species counted here as probably native.

FE. L. GREENE. The second botanist to work on Guadalupe Island was the Rev. Edward
Lee Greene, a keen student of west American plants, who by his own account had “seven
days of incessant rambling and climbing” there April 20-26, I885. He barely mentioned
a fellow naturalist who, according to Jepson (1931), was George Washington Dunn. Dunn
had just visited the island (Science 4:366, 1884) and so perhaps suggested Greene’s trip.
Greene (1885) gave interesting notes on the vegetation and listed his 106 collections, often
with notes; he also included, each marked with a dagger, the 25 species of Palmer’s list
that he failed to find. He added nine native or possibly native plants: Agoseris heterophylla,
Arctostaphylos sp., Dichelostemma pulchellum, Eschscholzia californica, Filago califor-
nica, Lupinus guadalupensis, Mammillaria blossfeldiana var. shurliana, Myosurus minimus
var. filiformis, and Opuntia prolifera, the first two of which were not found again; and
five weeds: Brassica rapa, Chenopodium murale, Hordeum murinum, Mesembryanthemum
crystallinum, and Polypogon monspeliensis, the first of which was not found again but
isn’t missed. He proposed nine new species then and others later. Greene’s mammillaria

GUADALUPE ISLAND FLORA

was from “towards the south end,” but he wrote that in this dry year the southern half
of the island was a sunburnt waste and that most of his additions were from the vicinity
of the springs, the district of highest fertility, where the goats did not now range. He
remarked that the presence of the small garrison at the beach, with soldiers coming daily
for water and sometimes to hunt goats, must already have had a favorable effect on the
vegetation of this very best part of the island.

Greene’s Guadalupe Island collections are scattered, at CAS, DS, GH, ND-G, NY, UC,
US, (and others?). The Greene-Nieuwland Herbarium at ND seems to have just 13. Some
specimens treated as his types are at CAS, GH, ND-G, UC.

F. FRANCESCHI. Dr. Francesco Franceschi (Emanuele Orazio Fenzi) was a noted Italian
horticulturist, newly arrived in California in “1893” (Tucker 1945). In December 1892 and
part of January he visited Guadalupe Island to see the state of the vegetation and to gather
full particulars on trees and shrubs peculiar to the island, “of which a few have been
sparingly introduced in gardens and others well deserve to be.” In an interesting account
of his visit, Franceschi (1893c) listed four lichens, identified by E. L. Greene, and 56
vascular plants, identified by Mrs. Katharine Brandegee (UC; duplicates at DS, POM,
US). Collecting so early in the season, he added no new plants except for mentioning
Cotyledon ____ (probably Dudleva virens), but his observations are astute and often colorful.
He published fuller accounts of the palm elsewhere (Franceschi 1893a, 1893b) and men-
tioned other articles sent for publication that seem not to have appeared yet.

A. W. ANTHONY. Alfred W. Anthony of San Diego made many trips in I887 to 1899 into
northwest Baja California and to the west-coast islands, collecting birds and mammals
(Nelson 1921:143). Plant collectors with him on two trips to the islands, including
Guadalupe, put up many sets of specimens that were distributed with printed labels bearing
Anthony’s name, and he is usually cited as the collector. These are mostly numbered, but
not all chronologically, and most have only inclusive dates for the trip. T. S. Brandegee
probably identified the specimens, a set of which is in his herbarium (UC). Clues to
Anthony’s itineraries come from publications by him and his associates and from his
manuscript catalogue of bird skins (Department of Birds and Mammals, California Acad-
emy of Sciences). In 1896, leaving San Diego July 9 with a chartered schooner, he was
on Guadalupe Island September 17-22. James M. Gaylord was plant collector (Anthony
1896) and on the whole trip apparently made at least 148 numbered collections. In 1897
on his schooner Wahlberg, Anthony visited the west-coast islands and the Revillagigedos,
stopping at Guadalupe March 20-26. Botanist T.S. Brandegee was with him as far as
San Jose del Cabo, collecting for his own herbarium (see next entry). Collecting plants
for Anthony for the whole trip was Alfred L. Stockton, a nephew of Mrs. Brandegee
from a ranch 9 miles from Ramona, San Diego County. Anthony’s numbers 231-262 are
from Guadalupe.

Anthony had visited Guadalupe Island before and went again with Hanna and Slevin in
1922 (Hanna 1925, Hanna and Anthony 1923).

T. S. BRANDEGEE. Townshend Stith Brandegee was a major collector of and student of
Baja California plants (Ewan 1942, Moran 1952). In 1897 he went with Anthony on the
schooner Wahlberg to the west coast islands, visiting Guadalupe Island March 20-26.
Brandegee (1900) wrote that the vegetation of Guadalupe was very rank and green but
not so far advanced as that of the islands they had just visited nearer the coast. He added
that most plants were not in blossom and that only at the lowest elevations could he find
them suitable for specimens. Nevertheless, he collected at least 52 species, including the

REID MORAN

following not found before: Aphanisma blitoides, Capsella bursa-pastoris, Castilleja ex-
serta, Cistanthe maritima, Lasthenia coronaria, Layia platyglossa, Phalaris caroliniana,
Platystemon californicus, Stebbinsoseris heterocarpa, and the endemics Stephanomeria
guadalupensis and Triteleia guadalupensis. The castilleja, lasthenia, layia, and platystemon
have not been found again. Apparently he landed the first day at Northeast Anchorage.
The cistanthe, phalaris, and stebbinsoseris now are local near the middle of the island on
the east side, an area hard to reach afoot from the north, and evidently not visited by
earlier collectors, but easy enough with the Wahlberg; and the aphanisma grows there
and southward. From the plants he did not find, he evidently did not reach the south end.
His herbarium is at UC. He later named Castilleja guadalupensis and Stephanomeria
guadalupensis.

FUR SEAL TRIP. In the last week of June of 1897 Professors Rufus L. Green, Charles B.
Wing, and Wilbur W. Thoburn, of Stanford University, visited Guadalupe Island as part
of a fur seal survey. Landing at the south anchorage, the west anchorage, and the northeast
anchorage, despite the late season they collected 37 species of plants (DS), reported by
Dudley (1899). To the known flora they added Frankenia salina, Phyllospadix torreyi,
and an endemic cistanthe described as Talinum guadalupense.

HARRY DRENT. The Brandegee Herbarium (UC) includes at least 28 specimens collected
on Guadalupe Island in 1898 by Harry Drent. Among these are 15 plants Brandegee himself
did not collect the year before, including such rare ones as Crossosoma californicum,
Phacelia phyllomanica, and Solanum wallacei ssp. clokeyi, the previously unreported
Euphorbia misera and Lupinus bicolor, and perhaps the last specimens taken of Castilleja
guadalupensis and Nicotiana attenuata. According to Stewart (1965:133), “Dutch Harry”
Drent, a Hollander 38 years old in 1899, “uneducated but bright,” was a deep-water sailor
who made at least five trips to Guadalupe Island. He hunted goats there for about five
months in 1896 (San Diego Evening Sun, 30 Sept. 1896), for some time in 1898, and from
April to late September of 1899 (Stewart 1965). In 1898 he brought to San Diego four
live birds of the soon-to-be-extinct Guadalupe caracara, captured “by a trick | had learned
while in South Africa” (Abbott 1933). He died in San Diego in 1910 (MacMullen 1969).

How did this goat-hunting sailor come to collect plant specimens and, even more puzzling,
how could he have such a good eye for rarities? When A. W. Anthony was on Guadalupe
in mid-September 1896, he could scarcely have failed to meet Drent and his fellow hunters.
I suppose that after five months of goats, Drent must have been glad to visit with the
naturalists and that he may have taken some interest in their collecting. Next year, in
1897, Drent was a seaman on Anthony’s schooner Wah/berg, certainly in December (Purpus
1898) and evidently on the spring voyage to Guadalupe with Anthony and Brandegee.
When Brandegee (1900:21) found few plants flowering that March, he must have asked
Drent to get others for him later and shown him what plants to collect. Since Drent did
not get them until the next year, evidently he was not staying on the island in 1897 but
was indeed on the Wahlberg. In January 1898 Carl Purpus wrote Mrs. Brandegee that
Drent was going to Guadalupe to collect palm seeds (Purpus 1898). Doubtless he also
hunted goats; but clearly that was when Harry Drent turned plant collector — and proved
a bright pupil.

W. W. BROWN. Willmott W. Brown, Jr., studied and collected birds on Guadalupe from
May | to June 28, 1906, accompanied by H. W. Marsden and Ignacio Oroso (Thayer and
Bangs 1908, Nelson 1921). The Brandegee Herbarium (UC) includes at least Il plant speci-
mens collected by Brown. A few at UC and US have printed labels whose text suggests
that his first set is at GH: “Presented to the Gray Herbarium by Messrs. John E. Thayer

GUADALUPE ISLAND FLORA

and Outram Bangs.” A check at GH showed eight specimens of Perityle, some with his
numbers between 4 and 84, so clearly he collected many plants. His collection of
Stephanomeria diegensis (UC) is the only one known from the island. Since he was two
months on the island and may have found other unreported plants, a list of his collections
would be of great interest. However, Dr. Reed Rollins, and later Dr. David Boufford,
wrote me they found at GH no fieldnotes or list of the Brown collections.

J.N. Rose. Joseph Nelson Rose, botanist with the Smithsonian Institution, had named
four new Guadalupe Island species in reporting the collections from Palmer’s second trip
(Vasey and Rose 1890). He visited the island himself on March 2-4, 1911, with C.H.
Townsend’s expedition on the U.S. Bureau of Fisheries steamer Albatross (Townsend
1911; Nelson 1921:146; Maxon 1935). According to Townsend “The botanical gatherings
alone [from this long trip] occupied nearly half of the special freight car to which the
ship’s load was transferred in San Francisco.” Rose’s Guadalupe specimens (16001-16040
and 16901?) and his field notebook are at US.

HANNA, SLEVIN, AND ANTHONY. G Dallas Hanna and Joseph R. Slevin of the California
Academy of Sciences and A. W. Anthony of San Diego visited the island July 11-17,
1922 on the Mexican patrol boat Tecate, on a trip to check on marine mammals (Hanna
and Anthony 1923, Hanna 1925, Anthony 1925). They made a small collection of plants
(CAS) and took some valuable 4” x 5” photographs, including some of oaks, pines, and
cypresses published by Eastwood (1929).

MASON AND SOLIS. Herbert L. Mason of the University of California and Octavio Solis
of the botanical garden at Chapultepec Park, Mexico City, visited Guadalupe Island April
19-21, 1925 (Hanna 1926). Mason collected in the canyon above Northeast Anchorage
one day and at the south end the next. He got 43 species of plants, including one weed
not found before, Hypochoeris glabra. In reporting Mason’s collections (CAS), Eastwood
(1929) also gave a useful summary of previous collections. There 1s no report of Solis’
collections. At US I saw one specimen of his No. 12, Hemizonia greeneana, April 20.
The printed label is headed: “Plants of Mexico/Received from Direccion de Estudios
Biolégicos, Secretaria de Fomento, Mexico”; very likely the first set is at MEXU.

J.T. HOWELL. John Thomas Howell of the California Academy of Sciences was on
Guadalupe Island November 14—16, 1931, and March 16-18, 1932. In November he climbed
up Barracks Canyon to the oaks the first day, visited the pines the second, and landed at
Melpomene Cove the third. In March he again visited the pines; then on the second and
third days, with one companion, he made the long overland trip from Northeast Anchorage
to the south end, camping overnight at the southern limit of the cypresses. He made very
good use of his time, on the November 1931 trip collecting numbers 8169-8223 and in
March 1932 numbers 8246-8335 (CAS), including bryophytes as well as seed plants. His
most remarkable discovery was Baeriopsis guadalupensis, which he described as a new
genus and species. Beyond that, he added to the known flora seven native or possibly
native species: Atriplex californica, Hutchinsia procumbens, Lepidium nitidum, Pla-
giobothrys collinus var. californicus, Spergularia marina, and Trifolium gracilentum. He
also added five new weeds: Bromus rubens, Medicago polymorpha, Mesembryanthemum
nodiflorum, Poa annua, and Sisymbrium irio. Howell (1941b) gave an interesting account
of his two trips and an annotated list (1942) of the vascular plants he had found. His
observations on the pines he included in a general account of the closed-cone pines of
the islands (Howell 194la). Howe (1934) reported his Hepaticae.

P. J. REMPEL. Peter J. Rempel, then of the University of Southern California, collected

REID MORAN

about 40 numbers on Guadalupe Island July 18-19, 1937, on a cruise of the Velero III of
the Allan Hancock Foundation (see Allan Hancock Pacific Exp. 1:46, 312.) He was the
first to collect on Outer Islet and so first to find Castilleja fruticosa, Eriogonum zapatoense,
Erysimum moranii, and Lavatera lindsayi; but the specimens (DS, LAM now RSA) were
not identified at the time and there was no report of his collections.

GEORGE LINDSAY. George Lindsay, of the San Diego Museum of Natural History and
later the California Academy of Sciences, visited Guadalupe Island several times between
1948 and 1957 and collected plants on three trips (SD). (See Lindsay 1951, 1966; Lindsay
and Dawson 1952; Moran and Lindsay 1950.) 1948, April 9-16, on Louie and Marcho
Cavanagh’s Marviento with Reid Moran (see next entry); NE Anchorage to Mt. Augusta
and the cypress grove, south end mesa, Outer Islet (2800-2840). 1950, January 27 to
February 4, with Dr. Carl Hubbs, on the Orca; Outer Islet, Melpomene Cove, above West
Anchorage, SE coast (1801-1818). 1955, June 10-16, on the Orca, with Dr. Hubbs, and
with Job Kuyt and Harvey A. Miller (see below); southeast sealing camp, Outer Islet,
Islote Negro (2617-2644); he spent four days circumnavigating the island in a small
launch and observed plants on offshore rocks.

REID MORAN. Reid Moran, of the San Diego Museum of Natural History, visited
Guadalupe Island 20 times between 1948 and 1988, ashore 97 days, some short, some
long (and some nights) (See Moran 1959, 1967, 1968, 1969, 1978). Most of the longer
and more fruitful trips were with Professor Carl Hubbs (Fig. 73) of the Scripps Institution
of Oceanography and on Scripps ships. Moran has named five endemic species and sub-
species: Castilleja fruticosa, Dudleva guadalupensis, D. virens ssp. extima, Eriogonum
zapatoense, and Lavatera lindsayi. To the known flora he (sometimes with botanical
companions) has also added 16 other species treated here as native or probably so, and
one of them as endemic, though a few may possibly be recent arrivals: Antirrhinum
watsonil, Camissonia robusta, Castilleja attenuata, Cuscuta corymbosa, Ervsimum mora-
nil, Gnaphalium bicolor, Heteromeles arbutifolia var. macrocarpa, Lomatium insulare,
Lycium fremontii, Pectocarva linearis ssp. ferocula, Plagiobothrys acanthocarpus, Poa
secunda ssp. secunda, Rhus integrifolia, Scrophularia villosa, Stipa lepida, and Zostera
marina, and seven clearly introduced and mostly weedy species: Bromus hordeaceus,
Erodium brachycarpum, Herniaria cineria, Nicotiana glauca, Phalaris minor, Ruta
chalepensis, and Sisymbrium orientale. His collections from the 1948 trip are at DS, the
rest at SD, as also his notebooks. 1948, April 9-16, with George Lindsay (see previous
entry), on the Marviento; NE Anchorage to pine ridge (over night), south-end mesa, Outer
Islet (2835-2947). See Lindsay (1966), Moran and Lindsay (1949, 1950), Moran (1951).
1957, February 9-14, with Dr. Hubbs; West Anchorage, Islote Negro, Outer Islet, NE
Anchorage to Mt. Augusta with Julio Berdegue (see Berdegue 1957) and camp at spring
(5602-5706). April 17-21, with Dr. Hubbs; West Anchorage, Esparsa Canyon, Islote
Negro, Barracks Canyon, Pilot Rock Beach (5950-5996). October 25—30, with Dr. Hubbs
and George Lindsay on the Orca; cliffs above NE Anchorage, Esparsa Canyon, Islote
Negro, south-end mesa, south palm canyon with George Lindsay, SE sealing camp (61 14—
6166). December 13-18, with Dr. Hubbs and with botanists C. H. Muller (UCSB) and
Gene Newcomb (UC) and C. F. Harbison (SD): up pine ridge to camp at old lookout and
down by spring and Oak Canyon, lobster camp with Newcomb, lobster camp to El Picacho
and south end (6420-6481). 1958, April 22-30, on Scripps boat, to collect pine seeds for
breeding program, with botanists Sherwin Carlquist (RSA, UC), Wallace Ernst, and Ira
Wiggins, (Wiggins & Ernst 1-223, Ernst 259-277, DS); South Esparsa Canyon, Pilot
Rock Beach to pine ridge for pine cones, Esparsa Canyon, NE Anchorage to camp at
spring, then to Mt. Augusta and south to lobster camp, south-end mesa, Islote Negro, up

GUADALUPE ISLAND FLORA

Arroyo Melpomene to flank of El Picacho at 650 m and down to lobster camp, NE An-
chorage (6593-6771). 1960, January 26-30, with Dr. Hubbs and class; NE Anchorage to
Oak Canyon, NE Anchorage to Lower Circus, NE Anchorage to beach cliffs at canyon
rim, NE Anchorage to canyon rim, Arroyo Melpomene to 600 m (7828-7851). 1965,
February 28 to March 6, with Dr. Hubbs and class; NE Anchorage to pine ridge, NE
Anchorage to Lower Circus and Oak Canyon, NE Anchorage to Mt. Augusta and cypress
grove, Juniper Canyon, lobster camp, Arroyo Melpomene to 400 m (1201 1—12078). 1967,
April 30 to May 3, with Dr. Hubbs; SW coast, Arroyo Melpomene to 350 m, NE Anchorage
to Lower Circus, SE Oak Canyon, NE Anchorage to north end (13755-13822). 1968,
June 21—22, with the Pacific Project of the Smithsonian Institution, on the Stella Polaris;
Outer Islet (over night), Arroyo Melpomene to 470m (15113-15128). 1969, February
21-22, a trip for Museum members; NE Anchorage, south-end mesa (15733). 1970, April
11-22, with Dr. Hubbs; pine ridge (over night) to NW slope, Arroyo Melpomene, Long
Canyon to El Picacho to south end, West Anchorage, Islote Negro (17274-17428). August
16-21, with Harold Pringle of La Jolla; NE Anchorage to Mt. Augusta and by jeep to
landing strip, Esparsa Canyon, lobster camp, south end, Outer Islet (18143-18171). 1971,
May 18—23, with Dr. Hubbs; Pilot Rock Beach to pine ridge (over night) and NW slope
to NE Anchorage, West Anchorage to El Picacho and gray cliff just south (18385-18397).
1973, March 25, with tourist trip; NE Anchorage to Lower Circus (20304-20310). 1974,
March 24, with tourist trip; NE Anchorage to Lower Circus (21665-21668). 1978, March
26-29, to collect pine cones for breeding program and for study of inbreeding in natural
population, with William Libby, Ken Eldridge, Daniel Axelrod, Yan Linhart, Cliff Ohmart,
plus graduate students and other helpers; to cypress grove and pine ridge (25374-25394).
1981, May 23—24, Palm Society trip to see Guadalupe Island palm, with botanists Mitchel
Beauchamp, Curtis Clark, Mary Hochberg, Steve Junak (22 specimens, SBBG), Tom
Oberbauer, Ralph Philbrick, Steve Timbrook, Sherry Whitmore, Howie and Eric Wier,
and others; Pilot Rock Beach up pine ridge to camp at old lookout, next day to upper
NW slope (29591-29600). August 19-26, with ecologist Martin Gonzalez and geologist
Carlos Garcia of Ecoterra, range ecologist Don Duncan, and Oscar Paulin of COTECOCA,
making preliminary inventory of natural resources for SARH (Gonzalez 1981), taking
truck on Mexican LCT, driving to cypress grove to camp (unbelievable!), then Lower
Circus, pine ridge, south-end mesa, cliffs NW of cypress (29799-29816) [unfortunately
a very dry summer]. 1988, March 28-30, with Margie Stinson on Pacific Queen, with
several botanists, some collecting: Bruce Baldwin (688-692, DAV), Jim Bartel, Mitchel
Beauchamp, Tony Burgess (7437-7469, ARIZ), Steve Junak (3469-3569, SBBG), Tom
Oberbauer, Chuck Quibell (1741-1817, NCC), Fred Sproul, Robert Thorne (63011-63111,
RSA), Ray Turner; NE Anchorage to spring, next day to Mt. Augusta [some to pines],
south-end mesa (31000-31018).

KUT AND MILLER. With Dr. Hubbs and George Lindsay, Job Kuyt and Harvey A. Miller
visited Guadalupe on the Orca June 10-16, 1955. They collected on the pine ridge, at
West Anchorage, at the old sealing camp on the southeast coast, and on Outer Islet, and
with Lindsay they made the first collections on Islote Negro. Kuyt collected higher plants
(1001-1081, UC). Miller collected bryophytes (5502-5532, DS); and he collaborated on
an account of the bryophyte flora (Crum and Miller 1956).

WEBER AND McCoy, Copp. William A. Weber and C. J. McCoy, Jr., of the University
of Colorado, visited the island April 19-29, 1963 with Dr. Hubbs. Though not reaching
higher elevations, they made the first thorough collection of lichens for the island: NE
Anchorage, lobster camp, south end, Islote Negro (36493-36680). They also collected
vascular plants (11969 etc., COLO) and bryophytes. Scripps technician Joseph F. Copp

REID MORAN

collected vascular plants in the same places (141-188, DS). Weber (1964, 1965, 1994)
named a new genus and another new species of lichen and reviewed the lichen flora of
the island.

ENRIQUE MELING. Alf Enrique Meling Lopez, as a student at Universidad Autonoma de
Baja California, in Ensenada, wrote a professional thesis on the state of the vegetation
of Guadalupe Island (Meling 1985). He did field work there in the spring of 1982, the
spring of 1983, and the summer of 1984. He made three long transects, listing species but
not always collecting vouchers; and some of his identifications are questionable. For check-
ing by Dr. Geoff Levin, he gave 80 specimens to SD, 46 of them numbered but not
chronologically, most of them dated but 20 with only month and year; others are at ENC.
Some of his localities are questionable: for example Camissonia robusta, Galium aparine,
Linaria canadensis, Lotus grandiflorus, and Uropappus lindleyi all from “Extremo Sur,”
the desertic south end. However, he added three or four new weeds to the known flora:
Lamarckia aurea, Raphanus sativus, Sisymbrium irio (perhaps a reintroduction), and
Triticum aestivum. About his specimen of Solanum tenuilobatum, see Doubtful and Ex-
cluded Species, p. 168.

Jose Rico C. Ing. José Rico C. of the Division de Ciencias Forestales, Universidad
Autonoma Chapingo, Chapingo, Mexico, visited the island in 1982 and again July I-15,
1994, collecting many specimens (CHAP). (See Rico. C. 1983).

PHILBRICK AND JUNAK. Botanists Ralph Philbrick and Steve Junak, with Peter Schuyler,
of the Santa Barbara Botanic Garden visited Guadalupe Island April 23-26, 1984, also
taking entomologist David Faulkner of the San Diego Museum of Natural History. They
landed near a fish camp on the southeast coast north of Morro Sur, walking inland and
up the airport road to 1800 ft; they had 2 ' hours on Outer Islet (see Philbrick 1984);
and they climbed from Esparsa Canyon to Mt. Augusta. Junak collected numbers 2055—
2117 and Philbrick some 64 numbers (SBBG). Faulkner also collected some plants (SD).

Junak and Philbrick also were with Moran in May 1981 and Junak again in 1988 (see
above); and Philbrick visited Northeast Anchorage on a cruise boat in 1975.

an
to

GUADALUPE ISLAND FLORA
Catalogue of the Plants

This catalogue aims to include all vascular plants, native and foreign, reported to grow
on Guadalupe Island and its islets. An asterisk (*) before a name shows that the plant
seems to be a late arrival, usually meaning a European weed (see Table | and discussion).
A letter phi () shows that it is a west American plant that has been taken for native but
is possibly or probably introduced (see Table 2 and discussion). A dagger (+) shows that
it is possibly or probably extinct on the island, commonly meaning that no one has seen
it since 1900 (see Table 3 and discussion). The synonymy includes chiefly names used
for the plant as it grows on this island.

The catalogue not only tells the most recently known distribution of each plant on the
island but also quotes or paraphrases such early reports as suggest differences in the past,
thus often telling of the decline of natives and the spread of weeds. These reports thus
sometimes give clues to the drastic but little documented changes in the vegetation that
took place in a very few years, even between Palmer’s visit of 1875 and Franceschi’s of
1892, when the goats apparently were just reaching their greatest numbers. The map
(Fig. 2) shows place names used in this account; for other names see under Place Names,
p. 9.

Because the Guadalupe flora is most closely related to the floras of the other Pacific
islands of southern California and Baja California (Fig. 1), the catalogue also notes oc-
currence on those islands. Useful sources of this information are Wallace’s (1985) tabu-
lation of floras of the Channel Islands, plus the floras of the Islands of Santa Cruz (Junak
et al. (1995), San Nicolas (Foreman 1967), Santa Barbara (Philbrick 1972, Junak et al.
1993), Santa Catalina (Thorne 1967, 1969), San Clemente (Raven 1963, Thorne 1969),
Todos Santos (Junak and Philbrick 1994a), and San Martin (1994b).

The catalogue does not describe most plants but does include some descriptive notes and
even full descriptions when they can tell something new, as well as thumbnail descriptions
for some interesting endemic plants. It cites specimens mainly for those plants that are
rare or seldom collected, and the account of collectors (p. 44) tells for each one where
the first set of specimens is deposited. Herbarium abbreviations follow Holmgren et al.
(1990). Field numbers prefixed by “M” are mine, and my collections since 1948 are at
SD.

Although this is the very latest flora of Guadalupe Island, it cannot be called up-to-the-
minute. My visits to the island, and so my observations of the plants, were scattered over
40 years, from 1948 to 1988 (see p. 50). Where I could go on the island was always
limited by time and by trip goals. On the latest trips I could not revisit all the remote
spots where I had seen rare plants before, to see whether they still survived. Thus some
of my sightings are long out of date, and I can’t know whether the plants are still there.

FERNS

Polypodiaceae s.1.

Cheilanthes newberryi (D.C. Eaton) Domin, Biblioth. Bot. 20 [Heft 85']:133. 1915.
Notholaena newberryi D.C. Eaton, Bull. Torrey Bot. Club 4:12. 1873.

RANGE. — Southern California and NW Baja California; San Clemente Island.

nN
ws)

REID MORAN

Palmer found this fern throughout the island, on rocks in dry exposed places (Watson
1876:122). It is fairly common from Northeast Anchorage and the Lower Circus to Arroyo
Melpomene and the southeast sealing camp. Rolla Tryon (1956:45) cited Guadalupe Island
collections by Palmer, Franceschi, Anthony, Rose, and Mason.

Notholaena californica D.C. Eaton, Bull. Torrey Bot. Club. 10:27. 1883.

RANGE. — Southern California to Arizona, Sonora, and central Baja California; Santa
Catalina Island.

This fern is known on Guadalupe Island from only two collections: locally common in
rock crevices, head of east fork of Arroyo Melpomene at 210 m (46/65); southeast coast
just north of Morro Sur, D. Faulkner 7D (SD). Dr. Dale Benham, who has the specimens
on loan, tells me they are the form with yellow farina, which has been called ssp. cali-
fornica.

Pellaea mucronata (D.C. Eaton) D. C. Eaton var. mucronata

Pellaea ornithopus Hook., Sp. Fil. 2:143. 1858.
Pellaea mucronata (D.C. Eaton) D.C. Eaton in Emory, Bot. Mex. Bound. 2(1):233. 1859.

RANGE. — California to southern Nevada and northern Baja California; Santa Rosa, Santa
Cruz, Santa Catalina, and San Clemente Islands.

Palmer found this fern rare, in crevices of the highest cliffs (Watson 1876:120); and
Franceschi (1893c:139) called it the scarcest fern on the island, seen only at the eastern
side, on basaltic rocks fully exposed to the sun. From Guadalupe Island, A. Tryon
(1957:159) cited only Palmer’s specimen. More recent collections are: two in crevice of
Lower Circus at 950 m (47838); one clump in rock crevice, head of east fork of Arroyo
Melpomene at 210 m (6/66).

Pentagramma triangularis (Kaulf.) Yatsk., Windham, & E. Wollenw., Amer. Fern
Jour. 80:15. 1990.

Gymnogramma triangularis Kaulf., Enum. Fil. p. 73. 1874.
Pitvrogramma triangularis Maxon, Contr. U.S. Natl. Herb. 17:173. 1913.

RANGE. — British Columbia to Nevada, Sonora, and southern Baja California; San Miguel,
Santa Rosa, Santa Cruz, Anacapa, San Nicolas, Santa Catalina, San Clemente, and Cedros
Islands.

Palmer found this fern in crevices of the highest cliffs in the “middle” and at the “south
end” of the island (Watson 1876:120). Franceschi (1893c:139) called this the most widely
spread fern, growing luxuriantly in the crevices of rocks with northern exposure, also in
very dry sunny spots but then much reduced in size. They grow on cliffs in the northeast
part of the island and as far south as Juniper Canyon but are not very common.

The Pentagramma triangularis complex, as now imperfectly understood, is “a confusing
array of morphologically cryptic taxa” (Yatskievych et al. 1990), “a puzzling complex of
intergrading chemical, chromosomal, and morphological variants” (Smith and Lemieux
1993), which is variously subdivided and is still due for more study. Alt and Grant (1960)
included Guadalupe Island in the range of var. maxonii, which they did not study in
detail. Dale Smith has annotated Guadalupe specimens as varieties maxonii and triangu-
laris; and George Yatskievych wrote me that one of the plants under Palmer 10] at MO

GUADALUPE ISLAND FLORA

FIGURE 36. Pentagramma triangularis (M17317) on cliff at north end of the island, 800m, 13
April 1970.

is ssp. viscosa, though a bit more divided-leaved than usual. He said he had seen little
Guadalupe material and most of that was pretty weird. | am a coward, just calling it all
P. triangularis. Figure 36 shows a form with fronds much as in typical “iangularis but
viscid.

Polypodium californicum Kaulf., Enum. Fil. p. 102. 1824.

RANGE. — Central California to NW Baja California; Santa Cruz, Santa Rosa, Anacapa,
San Nicolas, Santa Barbara, Santa Catalina, San Clemente, Los Coronados, Todos Santos,
and Cedros Islands.

Palmer found this fern abundant at the north end, in cracks of rocks in damp places,
sometimes covering large surfaces (Watson 1876:120), but Franceschi (1893c:139) called
it rather scarce, always in shady or sheltered localities. It grows at 600-900 m, mostly in
damp and shady places: on the north slope under the pines and on north cliffs, on the
south wall of Oak Canyon, and with the mimulus on the gray cliff above Campo Oeste,
in a spot that is perhaps often foggy. I have not seen it abundant.

Polypodium scouleri Hook. & Grev., Icon. Fil. 1: pl. 56. 1829.

RANGE. — Vancouver Island along the coast to central California; Santa Cruz and
Guadalupe Islands.

Palmer found this fern encircling the trunk of a single oak in a thick mat of moss and
constantly wet by the fogs, covering the tree with a network of its strong tough roots to
the height of ten feet (Watson 1876:120). Greene (1885:228) feared that “Dr. Palmer’s

nn
n

REID MORAN

gathering of it from ‘the trunk of a single oak’ may have proved the extermination of
the species on this island.” Evidently, though, Greene was looking up the wrong tree, for
Hanna and Slevin again found it on oaks (Eastwood 1929:399). I found it on the trunks
of two pines on the northeast ndge at 530 m (M6422), as well as on northwest-facing
vertical cliffs nearby and farther towards North Point, at 460 m (A16420). The fern-bearing
trees seem to be gone, but the cliff is still there and I hope also the fern.

Polystichum munitum (Kaulf.) C. Presl, Tent. Pterid. p. 83. 1836.

Aspidium munitum Kaulf., Enum. Fil. p. 236. 1824.
Polystichum munitum (Kaulf.) C. Presl ssp. solitarium Maxon, Fern Bull. 11:39. 1903.
Polystichum solitarium (Maxon) Underw. ex Maxon, Contr. U.S. Natl. Herb. 10:493. 1908.

RANGE. — Alaska to Montana and southern California; Santa Rosa, Santa Cruz, and
Guadalupe Islands.

Palmer saw only two large bunches of this fern, at the northern end in a rocky place
inaccessible to goats and constantly damp from the prevalent fogs (Watson 1876:120).
Though seldom collected, it persists on the sheer northwest cliff at 600 m near North
Point (4/2959/), where noted by Sherry Whitmore.

Maxon (1903) wrote that his ssp. solitarium was distinguished by the dark persistent chaff
which very thickly covered the rachis throughout and by the narrower and extremely
coriaceous pinnae, which were more scurfy below and somewhat glaucous in appearance
and possessed of abbreviated decidedly cartilaginous appressed incurved teeth. The holo-
type is Anthony 9 of 1896 from Guadalupe Island (GH); and Palmer /02 is a paratype.
David Wagner annotated my collection (A/2959/) in 1989 as P. munitum. See Wagner
(1979),

GYMNOSPERMS

Cupressaceae

Cupressus guadalupensis S. Watson ssp. guadalupensis
iP g

Cupressus guadalupensis S. Watson, Proc. Amer. Acad. Arts Sci. 14:300. 1879.

Cupressus macrocarpa Hartw. var. guadalupensis (S. Watson) Mast., Gard. Chron, Ser. 3, 18:62.
1895.

Cupressus macrocarpa of Guadalupe Island references.

RANGE. — Endemic to Guadalupe Island.

Tree 15—20 (—25) m tall, the crown dense to rather open, commonly rounded or subrounded,
in isolated trees often broader than high, occasionally pointed, the main axis identifiable
nearly to top or often lost in branches (Figs. 37, 38). Trunk mostly 2—5 dm thick, often
to | and rarely to 2 or reportedly 2.5 m, in age often fluted (Fig. 39); primary branches
mainly ascending at about 45—60°, the lowest 1-2 m above base (Fig. 39); bark of trunk
and branches commonly smooth, peeling annually in thin non-fibrous curling plates two
to several centimeters long and wide, the new bark pink to reddish to greenish, mostly
soon aging various shades of gray to lavender gray, but in a few trees the bark on branches
smooth but on trunks persistent, fibrous or often shaggy, brown, splitting lengthwise into
strips. Foliage green to rich glaucous blue-green, glaucous especially on young trees.
Ovulate cones 2440 (—S0) mm long, nearly as thick, generally with 8 or 10 scales, often

GUADALUPE ISLAND FLORA

UMOIO JO AlIsUDp

“SOHL YOR T “OAc PUL OU] UL SOO1] “SE TUNOL] “SLOT YURPL LZ “Watudiessa Wojsom ay} UO sf “LE AYN}

pue adeys ul uoneuea SulMoys ‘(sisuadnjppons “dss sisuadnjppons snssaidny) ssaid&o adnjepeny ‘g¢ pur /¢ SAYNOL

vay

REID MORAN
with conspicuous horns which may be 6 mm long.

Palmer reported only that the cypress grew in irregular clusters in the “middle” of the
island (Watson 1876:119); he called it a fine widely spreading tree, though varying much
in habit, averaging about 40 feet high. According to Greene (1885:217) “this tree appears
formerly to have occupied almost the entire plateau of the northern half of the island;
but now, upon the greater part of this tract, only the fallen trunks, far gone in decay,
remain. The cause of this destruction | cannot guess.” He mentioned that the cabins near
the springs stood in a fine cypress grove. Franceschi (1893c:138) wrote that the principal
grove on the higher central plateau covered not less than two or three square miles. Dudley
(1899:281) wrote that samples of dead wood from the southern part of the island agreed
in structure with cypress wood and not with juniper wood which had previously been
supposed to occur there. He concluded that “the cypresses, therefore, probably extended
over the southern half of the island until destroyed by the goats: and their disappearance
may have caused the disappearance of subperennial springs, as none exist in that region.”
It is not clear how far south or at how low an elevation the wood was collected, but I
suspect that Dudley’s conclusion is much too sweeping. Furthermore, Palmer saw not
merely dead wood but also living junipers in the “central” and “southern” part of the
island, and indeed Dudley reported dead junipers.

Cypresses now grow mainly on the comparatively gentle western and southern slope of
the island in the vicinity of Mt. Augusta, at about 950 to 1200 m elevation. The main
rove extends from about west of Mt. Augusta around to the south-southeast, with a more
cattered stand to the northwest. The total number of trees must be in the thousands. On
the bench near the spring one old dead tree remained at the time of my first visits but is
now gone. The only trees on the eastern drainage are a few at the head of the canyon
just south of Mt. Augusta. Fallen trees are especially numerous in the area from west to
northwest of Mt. Augusta, and more fallen ones were seen each trip. The southern part
of the grove is still a beautiful dense forest (Fig. 25), but many trees show severe damage
to the bark by the goats. As with the other trees of the island, no seedlings escape the
goats. Although in dense parts of the grove trunks may be as thin as | dm, they must all
be well over 100 years old.

o
o
S

After the visit of Mexican President Lopez Portillo on September 28, 1980, there was a
plan to fence the cypress grove and keep the goats out, to raise cypress seedlings at the
Tecate nursery from Guadalupe seed, and to replant bare areas. However, the fence was
never finished, and the project clearly was abandoned.

Franceschi (1893c) called attention to the great variation of the cypress in habit and color
and also in size and shape of the cones. Howell collected a series of specimens to show
the variation and gave notes on his observations. I also was impressed with the great
variation, as expressed in the description above, the crown dense to open (Figs. 37, 38),
the bark smooth to shaggy and persistent (Figs. 39, 40), the foliage green to very glaucous,
and the cones (Fig. 41) varying greatly in size and shape.

A related tree in Orange and San Diego Counties, California, and in northwest Baja
California was sometimes called C. goveniana Gordon or C. sargentii Jeps. but was then
more commonly called C. guadalupensis, as by Abrams (1923:73). Jepson (1922) named
this mainland tree C. forbesii but did not differentiate it from other species. In his thorough
revision of New World cypresses, Wolf (1948a), after discussing the difficulty of specific
and subspecific delimitation, recognized 15 species and no varieties. He kept C. forbesii
as a species closest to C. guadalupensis but said (p. 5) that a most drastic reduction of

GUADALUPE ISLAND FLORA

YOOWS SUIALY Sv paquosap SI 901) FUL) “TR6l snsny C7 yzeq Adseys YM YUNA ‘Op

Buryoursq SuLMOYsS ‘YUN pany 100J-9 YUM 1],

"6¢ TUNDLA

‘(sisuadnjpppns

dss sisue

(ye

q pal

ALlayo

AOL] “LS6] Ateniqay ¢] yeq Surpaad-yjoours Surmoys pure

adnyppons snssa..dny) ssaiddo adnjependy ‘Op pue 6

vay

REID MORAN

FIGURE 41. Foliage and cones of Cupressus guadalupensis ssp. guadalupensis (M5671).
13 February 1957

species might place it as a subspecies of C. guadalupensis or even reduce it to synonymy.
Little (1970) discussed the problem further and proposed a more conservative treatment.
Thus where Wolf accepted 15 species for the New World, Little had 8 species and 7
additional varieties, with forbesii as a variety of C. guadalupensis. Silba (1981) had 8
species and 9 additional varieties, including C. guadalupensis var. forbesii.

Wolf did not visit Guadalupe Island but had to rely on the herbarium specimens of
Franceschi, Anthony, Hanna, Fleming, and Howell, plus cultivated trees. Possibly the
island trees he saw in cultivation were a biased sample, grown from seeds from trees
selected for their glaucous foliage as more ornamental. For C. forbesii he saw only two
collections from Baja California, whereas well over a dozen localities are now known
(Minnich 1987, fig. 4). Wolf said the two species were alike in having smooth cherry-red
or mahogany-brown exfoliating bark on their trunks and branches; they generally devel-
oped a branched crown with several leaders instead of retaining a single central axis; the
foliage was remarkably alike, despite a difference in color, with the dorsal glands, when
visible, usually inconspicuous and rarely if ever active; and they both had large cones,
with large brown or tan seeds. He said that C. guadalupensis was a larger tree in the
wild as well as in cultivation; usually with bluish green or glaucous blue-green instead
of green foliage; with generally larger ovulate cones, attaining a maximum length of 45
mm, whereas in C. forbesii they are rarely over 30 mm long; with the inner faces of the
cone scales glaucous and the seeds usually glaucous. He added that with the limited
material of C. guadalupensis examined, it seemed that the staminate cones generally had
as many as 18 scales and were therefore much larger than those of C. forbesii, which
had only 12 or 14 scales; and his key shows scales 14-18 for C. guadalupensis and 10-14
for C. forbesii.

GUADALUPE ISLAND FLORA

Wolf described C. forbesii as usually less than 10 m high, with a trunk 3—S dm thick; but
according to Parish, as quoted by Wolf (p. 64), trees on Mt. Tecate had trunks sometimes
nearly | m thick. Since, however, as Wolf pointed out (p. 162), recurrent brush fires have
prevented the survival of old trees, the maximum size of mainland trees is unknown.
Although Wolf's description of C. guadalupensis said the trees varied from slightly blue-
green to rich glaucous blue-green, the notes he quoted on Howell 8299 say “with very
green, non-glaucous leaves”; and I have seen many green trees. With more material, the
suggested distinction in number of scales of the staminate cones does not hold. For ex-
ample, one collection from Guadalupe Island (Ramsay & Peterson s.n. SD) has mostly
8-12, and one from south of Tecate (M//3986) has mostly 16-18. In specimens from
Guadalupe Island, the seeds may lack glaucousness. However, the difference in size of
ovulate cones is impressive. For 42 cones from the island, all from different trees (M/567/,
M25384), | found a range in length of 24-38 mm, with a mean of 29.5 + 0.34 mm. For
mainland trees I measured 79 cones on 28 herbarium specimens (SD) from 14 localities
in Baja California and about 8 localities in southern California. The range in length was
14-29 mm, with a mean of 21.5 + 0.35 mm.

Zavarin et al. (1967) studied the distribution of tropolonic heartwood constituents in various
species of Cupressus. For C. forbesii and C. guadalupensis they found common constitu-
ents separating these two from other U.S. coastal cypresses, but their preliminary data
based on small samples did not show substantial differences between the two. Dr. Zavarin
wrote me in 1994 that he and his associates are continuing their work, using terpenoids
from leaves. They have sampled four U.S. populations out of many populations of C.
forbesii and eleven cultivated trees of undocumented origin for C. guadalupensis. They
have found only small differences between two populations of C. forbesii and for some
terpenes highly significant differences between those populations and their sample of C.
guadalupensis. He writes that the chemical data so far suggest an appreciable gap between
C. forbesii and C. guadalupensis. Considering the great morphological variation in the
island population, however, it might not be surprising to find great chemical variation as
well. At least it would seem to require some large and well-selected samples to show
significant differences between island and mainland trees as a whole.

My impression is that the trees on the island seem more variable than those on the mainland,
and in many respects variation in the island population seems to more than encompass
that in the various mainland populations. However, the remarkable difference in cone size
suggests that mainland trees be separated subspecifically, as C. guadalupensis ssp. forbesii
(Jeps.) R. M. Beauchamp.

Guadalupe cypress grows well in central and southern California, making a handsome
tree. The oldest in cultivation are said to be from seed Palmer collected on his first trip.
Wolf (1948b:389) told of many fine old trees 50 or 60 feet high and 30 or 40 feet wide,
with trunks to 4 feet thick, adding that he had never found cones. Wagener (1948:299)
told of an experimental planting of twenty trees of C. guadalupensis at Stantord University,
with C. macrocarpa Gordon and others. He said they grew almost as fast as C. macrocarpa,
in both height and diameter. Among the test species he found C. guadalupensis outstanding
in character, growth rate, ability to compete with surrounding trees, and freedom from
pests and diseases. He found it apparently not at all susceptible to cypress canker fungus
(Coryneum cardinale Wagener), so destructive of C. macrocarpa. He concluded that
Guadalupe cypress deserves wider use.

REID MORAN
Juniperus californica Carr., Rey. Hort. Ser. 4, 3:352. 1854.

RANGE. — Central California to S. Nevada, NW Arizona, and north central Baja California;
Isla Cedros.

On his first trip, Palmer found the juniper all over the “middle” of the island and occa-
sionally at the “south end,” in the ravines and low valleys (Watson 1876:119). Only ten
years later, Greene (1885:217, 218) found it “upon the very verge of extermination. .. .
Of the grove in the middle of the island [there remained] only three trees that were not
quite dead; and on these three only a few tufts of green twigs gave the feeble sign of
nearly exhausted vitality.” The fur seal party in 1897 found many dead but no living trees
(Dudley 1899:281). About 1.5 or 2 miles south of the cypresses in 1932, Howell (1941b:40)
found a ghost grove which he took to be what was left of the juniper grove described
by Palmer, the dead remains extending for several miles.

However, Greene in 1885 did note one vigorous juniper on a southeastern cliff, hanging
over the sea. In that part of the island, where goats are fewer though still too many, a
few junipers have survived until recent years, showing more damage with each of my
visits. In June 1955 Job Kuyt (7077) and George Lindsay (2635C) collected from a juniper
in a canyon bottom about a mile north of the southeast sealing camp; it was about 5m
tall and wide, only partly alive, the goats having eaten as high as they could reach. In
1957 at 275 m in Juniper Canyon there were two healthy erect trees 5 m tall, with rounded
crowns 6m wide. Near the mouth of the canyon, at 40 to 125 m were many dead junipers
and four living ones, though two were badly damaged. One gnarled sprawling tree was
4m high and 10 m wide, with a trunk 3 dm thick (4/6454). In 1965 two of the four were
dead and only one remained in good condition. In the next canyon south, dead junipers
extended to 350m, where a healthy one 4m high and 8 m wide (4/4/2062) hung from
the south wall of the canyon, out of reach of goats (Fig. 30). In January 1960, when
climbing the southeast slope west of the barracks at Northeast Anchorage to check a
shrub of Malosma at 300 m, | saw beyond it, too far for certain identification and beyond
the reach of man and goat, a healthy shrub of what appeared to be juniper.

Pinaceae

Pinus radiata D. Don var. binata (Engelm.) Lemmon, W. Amer. Cone-Bearers
p. 42. 1895,

Pinus insignis Loud. var. binata Engelm. in W. H. Brewer & S. Watson, Bot. Calif. 2:128. 1880.

Pinus radiata D. Don forma hinata (Engelm.) J.T. Howell, Leafl. W. Bot. 3:3. 1941.

Pinus radiata D. Don forma guadalupensis J.T. Howell, Leafl. W. Bot. 3:3. 1941.

Pinus radiata D. Don ssp. hinata (Engelm.) E. Murray, Kalmia 12:20, 1982. [Basionym incorrectly
given as P. radiata var. binata Engelm.]

Pinus muricata, P. remorata, and P. radiata forma radiata of Guadalupe Island references.

RANGE. — Endemic to Guadalupe Island.

Palmer reported the pine from high elevations at the northern end of the island (Watson
1876:119). Franceschi (1893c) said they extended all over the northern and northwestern
part, which in times past they must have covered in a very thick forest, the finest trees
growing amongst the palms. In my time the main pine grove has been on the northwest
slope of the island near the crest of the northeast ridge, at about 300 to 800 m (Figs. 20,
21, 42). On the hotter and drier southeast side of this ridge, where the fog commonly

GUADALUPE ISLAND FLORA

FIGURE 42. The endemic Guadalupe Island pine (Pinus radiata var. binata) at the old lookout,

800 m, 13 April 1948. This tree was ca. 30m tall with a crown 32m wide and a trunk 2.1 m

thick—the largest pine seen and larger than any Monterey pine measured on the mainland

dissipates as it blows across the ndge, a few trees grew only just below the crest (Fig.
22). The northeasternmost trees were within + | km of North Point (Fig. 19). A few small
groups were scattered above the main grove, up the ridge to the southwest, in 1957 still
to 1160 m, and a few large pines were scattered on the northwest slope among the palms
In a panoramic view from the northwest, Libby et al. (1968, fig. 1) showed most of the
pines on the island in 1957.

From two short days of sampling in March 1964, Libby et al. (1968) counted only 383
living pines but said their census, particularly of trees low on the cliffs and in canyons,
was approximate. A census in 1978 showed that about 5-10 percent of the trees seen in
1964, and all the seedlings, had since died (Libby 1978). Many fallen trees are to be seen
(Fig. 19), and the decline continues rapidly.

Although earlier botanists reported only Pinus radiata var. binata from the island, Mason

REID MORAN

(1932) added P. muricata D. Don, and McMinn and Maino (1935) included Guadalupe
in the range of P. remorata Mason. After studying variation on the island, however,
Howell (1941a) concluded that all the pines belong to one variable species; and Newcomb
(1959), Libby et al. (1968), Axelrod (1980), and Millar et al. (1988) have agreed. Howell
found a few trees with cones very similar to those of typical P. radiata on the coast of
central California — [reflexed and] very asymmetrical, with umbos of the outer scales
much thickened — and a few with [spreading] almost symmetrical cones, with the scales
alike on all sides; but most had intermediate cones, somewhat asymmetrical, with the
outer scales noticeably but not prominently umbonate. (See Figures 43, 44.) Finding com-
plete intergradation among the three types, and finding no other differences correlated
with those in the cones, Howell treated them all as forms of one species. Whereas Engel-
mann had separated the Guadalupe tree as variety binata on the basis of paired needles,
Howell found that on vigorous shoots the needles are mostly in threes, as in typical P.
radiata. So he did not take up the variety but instead named three formae based on the
three types of cones: forma radiata, with very asymmetrical cones, forma guadalupensis,
with nearly symmetrical cones, and forma binata, with intermediate cones like those of
the type collection (Palmer 90, MO).

The pine of Cedros Island was formerly called Pinus muricata D. Don or P. muricata
var. cedrosensis J, T. Howell, though Mason (1930) referred Anthony’s 1896 specimen
(UC) to P. remorata Mason. However, recent students have agreed that the Cedros pine
is another form of P. radiata (Newcomb 1959, Libby et al. 1968, Axelrod 1980, Millar
et al. 1988), and Axelrod named it P. radiata var. cedrosensis (J.T. Howell) Axelrod.

Forde (1964) found that in the Californian populations of P. radiata each needle has zero
to two resin canals; whereas for Guadalupe and Cedros trees Newcomb (1959) reported
the number as two to several.

As in P. radiata, so also in P. muricata, the cones of different trees vary from spreading
and nearly symmetrical, with thin cone scales, to reflexed and very asymmetrical, with
the outer scales (away from the branch) greatly thickened. Linhart (1978) supported the
old view that thick cone scales are adapted to protect the seeds from fire and the newer
view that they also tend to protect the seeds from squirrels. In both species, mainland
trees where there are squirrels have mostly very asymmetrical cones; and the populations
with the greatest proportion of nearly symmetrical cones are on islands with no squirrels,
P. radiata on Guadalupe and P. muricata on Santa Cruz. Axelrod (1980) rejected these
ideas, suggesting instead that larger cones with thicker scales protect the seeds against
summer drought and developed in the Californian populations with the trend to increasing
summer drought. He suggested that ancient types have persisted on Cedros and Guadalupe
Islands, where he supposed there still were summer rains.

Axelrod compared large samples of cones from Guadalupe and Cedros Islands with sam-
ples from the three mainland populations of P. radiata and from all known fossil sites.
He showed (1980, fig. 2) that cone size was significantly different in each of the five
living populations, with Cedros cones smallest (mean length 6.5 + 1.3 cm) and Guadalupe
cones next (8.2 + 1.2 cm), as compared to 9.6 + 1.3 cm, 11.4 + 1.4 cm, and 13.9+ 1.7 cm
for the three Californian populations. The oldest fossils also had small cones, some floras
27 million years old with cones about the size of Guadalupe cones.

The Guadalupe population thus is significantly different in cone size from other living
populations. Also, Linhart (1978:36) called it the only population with a high frequency
(over 30%) of thin-scaled [nearly symmetrical] cones. It is also noteworthy that the maxi-

GUADALUPE ISLAND FLORA

FIGURES 43 and 44. Cones of Guadalupe Island pine (Pinus radiata var. binata) from neighboring
trees, 14 December 1957. FIGURE 43. Cones of the reflexed asymmetrical radiata form (M643/)
FIGURE 44. Cones of the spreading nearly symmetrical remorata form (M6432)

REID MORAN

mum [basal] trunk diameter of 2.71 m noted by Howell is greater than the 1.83 m d.b.h.
recorded for the largest mainland tree measured (Libby et al. 1968). The largest tree |
measured (1/6432), just south of the rock shelters at the old lookout (Fig. 42), had a trunk
diameter at breast height of 2.1 m, a branch spread of 32 m, and a height of + 30m, as
estimated from a photograph showing the full branch spread. The top has since broken
out of this tree.

On the basis of allozymes, Millar et al. (1988) agreed that the five living populations of
P. radiata are well differentiated. They found the Guadalupe and especially the Cedros
populations highly variable. All together then, the Guadalupe population is amply distinct
from other native populations and seems best kept as a geographical variety or subspecies.

Pinus radiata 1s probably the most widely planted tree species in the world, frequent in
both landscape and forest plantings in most countries of Mediterranean climate (Libby et
al. 1968). Under the name of radiata pine, it is especially important in the forest economies
of New Zealand, Chile, and Australia; and worldwide it makes up an increasing share,
now about three percent, of the international trade in wood and wood products (Libby
1995). It is becoming a domesticated plant and one of the best known plants genetically,
and genetic improvement requires bringing in genes of all native populations, including
the Guadalupe trees (Libby 1990). Many cones were therefore collected on Guadalupe in
1958, 1964, and 1978, and seedlings are now growing in California, Australia, New Zea-
land, and elsewhere. Forest geneticists consider that Guadalupe trees are valuable for good
growth form and high wood density and may be more resistant to winter cold and to
relatively hot and dry climates. And they are less susceptible than California trees to some
diseases, such as Western gall rust (Endocronartium harknessii) (Old et al. 1985), that
are serious threats to radiata pine throughout the world, and so can help in breeding for
resistance.

DICOTS

Aizoaceae

*Mesembryanthemum crystallinum L., Sp. Pl. p. 480. 1753.

Gasoul crystallinum (L.) Rothm., Notizbl. Bot. Gart. Berlin 15:413. 1941.

RANGE. — Native to Africa; a widespread weed along the coast of western North America
and elsewhere.

Greene (1885:223) found this weed on the beach at the landing [Northeast Anchorage],
and Dudley (1899:282) reported it from the south anchorage. It is now one of the com-
monest plants in the southern part of the island and on the islets (Fig. 74) and, though
much less common in the northern part, occasional even at the spring and the pine grove.

* Mesembryanthemum nodiflorum L., Sp. Pl. p. 480. 1753.
Gasoul nodiflorum (L.) Rothm., Notizbl. Bot. Gart. Berlin 15:413. 1941.

RANGE. — Native to Africa; a common weed along the coast of southern California and
northern Baja California.

Howell (1942:148) was first to find this weed on the island, at the south end. It is now

GUADALUPE ISLAND FLORA

rather common in the southern part of the island, including Islote Negro, and occasional
farther north, even to the spring.

Anacardiaceae
Malosma laurina Nutt. ex Abrams, Fl. Los Angeles, ed. 2, p. 220. 1917.

Rhus laurina Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:219. 1838.

RANGE. — Southern California to southern Baja California; Santa Catalina, San Clemente,
Todos Santos, and Cedros Islands.

Palmer saw only four shrubs, in the crevices of high rocks (Watson 1876:114). Franceschi
(1893c:135) saw what he took to be the same four, growing not far apart on the basaltic
cliff east of the cabins; and Dudley (1899:282) mentioned the same. Franceschi also saw
another, too high up to be surely identified, on the right bank of the canyon near the
landing. Barkley (1937:417) cited Franceschi’s specimen and also those of Drent and of
Brown. On early trips I saw four shrubs on the vertical cliffs of the Lower Circus at
900 m, large and old and undoubtedly those seen by earlier botanists; but by 1981 only
three remained (Fig. 29). A fourth one, doubtless that mentioned by Franceschi, grows
at 300 m on the southeast slope of Barracks Canyon near the mouth: I checked it with
field glasses from 50 m, which is about as close as anyone but a goat would care to go.

Rhus integrifolia (Nutt. in Torr. & A. Gray) W. H. Brewer & S. Watson, Bot. Calif.
1:110. 1876.

RANGE. — Southern California and NW Baja California; San Miguel, Santa Rosa, Santa
Cruz, Anacapa, Santa Catalina, San Clemente, Los Coronados, Todos Santos, and Cedros
Islands.

Moran (1951:157) reported this shrub from the inner slope and floor of the crater on
Outer Islet, where it is fairly common with Lycium californicum and Cistanthe guadalu-
pensis (Fig. 48). No one has ever found it on the main island.

Boraginaceae

Amsinckia menziesii (Lehm.) A. Nelson & J. F. Macbr., Bot. Gaz. 61:36. 1916.
Amsinckia intermedia, A. spectabilis, and A. vernicosa of Guadalupe Island references.

RANGE. — British Columbia to Idaho, Utah, and NW Baja California; South America;
Santa Cruz, Anacapa, Santa Catalina, and Cedros Islands.

Palmer found this plant very abundant on level ground at the “south end” (Watson
1876:118). It is widespread from low elevations in the northeast part of the island, as at
Northeast Anchorage and in Esparsa Canyon, to the flanks of El Picacho at 600 m and
the mouth of Arroyo Melpomene, but usually just a few scattered plants. Flowers are
February to May.

Dr. Peter Ray wrote me in 1958 that at GH he was able to find only one sheet of Palmer’s
original collection (No. 69), with only one small immature specimen, lacking nutlets. (If
the plant was abundant as reported by Palmer, it seems strange that this should be the
only specimen.) He found it labeled in Gray’s hand as 4. vernicosa Hook. & Am.; but

REID MORAN

since Gray did not cite this specimen in his later work, Dr. Ray suggested that this
identification may have been hasty or without conviction. He referred the specimen to 4.
menziesii in accordance with the treatment of Ray and Chisaki (1957).

Cryptantha foliosa (Greene) Greene, Pittonia 1:113. 1887.

Krynitzkia foliosa Greene, Bull. Calif. Acad. Sci. 1:205. 1885.
Eritrichium muriculatum of Guadalupe Island references.

RANGE. — Endemic to Guadalupe Island.

This is a stiffly erect, divaricately branched annual 0.54 dm high, with leaves 1—6 cm
long, and with 4 homomorphous nutlets.

Palmer found this plant in warm clear places in the canyons of the “middle” of the island
(Watson 1876:118). It is widespread on the island, from the pine ridge and north slope
to Outer Islet, often seen on grassy slopes, one of the commonest native annuals and in
good years so abundant as to color the slopes bluish white. It flowers January to June.

Johnston (1925:63) wrote that this species was readily recognized by its peculiar habit of
branching, congested spikes, coarsely armed calyx-lobes, and small nutlets. Besides the
type collection of Greene (“isotypes” GH, UC), he cited specimens of Anthony, Brandegee,
Palmer, and Townsend.

Cryptantha maritima (Greene) Greene var. maritima

Krynitzkia maritima Greene, Bull Calif. Acad. Sci. 1:204. 1885.

Cryptantha maritima (Greene) Greene, Pittonia 1:117. 1887.

Cryptantha maritima (Greene) Greene var. genuina |. M. Johnst., Contr. Gray Herb. 74:48. 1925.

Eritrichium angustifolium, E. ramosissimum, and Krynitzkia ramosissima of Guadalupe Island ref-
erences.

RANGE. — Southern California to Arizona and southern Baja California; San Nicolas,
Santa Barbara, Santa Catalina, San Clemente, Los Coronados, San Benito, and Natividad
Islands.

Palmer found this plant on level spots at the “south end” and also near the beach on the
eastern side (Watson 1876:118). It is widespread at low elevations, from Northeast An-
chorage to the south end and on Islote Negro, on alluvial fans, in arroyos, and on grassy
slopes, sometimes common.

Greene’s type is from Guadalupe Island (“isotypes” GH, UC). Johnston (1925:48) also
cited Guadalupe collectons by Anthony, Brandegee, and Palmer.

Harpagonella palmeri A. Gray, Proc. Amer. Acad. Arts Sci. 11:88. 1876.

RANGE. — Southern California to Arizona and central Baja California; Santa Catalina
Island.

Palmer found this plant only at the “south end,” in low valleys (Watson 1876:118). Howell
(1942:152) reported it as only in the middle part of the island, where fairly common. |
also have seen it only near the middle of the island: on the east to northeast slopes of El
Picacho and in the Comal area south of the cypress grove, at 275 to 900 m, rather scarce
but locally common.

GUADALUPE ISLAND FLORA

Gray (1876b) described the species and the monotypic genus Harpagonella from Palmer’s
Guadalupe Island collection (No. 70, GH). Further, the very distinctive bur of this plant
inspired Gray (1876a) to write about kinds of burs in the family. All authors have rec-
ognized this genus, and Johnston (1924) in his key kept it under a separate tribal name
Harpagonelleae.

Pectocarya linearis (DC.) Ruiz. & Pav. ssp. ferocula (1. M. Johnst.) Thorne, Aliso
9:191. 1978.

Pectocarya linearis (DC.) Ruiz & Pav. var. ferocula I. M. Johnst., Contr. Arnold Arb, 3:95. 1932.

RANGE. — Central California to central Baja California; Santa Rosa, Santa Cruz, Anacapa,
San Nicolas, Santa Catalina, San Clemente, and Cedros Islands; also Argentina.

This plant was first found on the island in 1957. In 1958 Dr. IL. M. Johnston verified three
of my collections: M5960 and 6633 from the bed of Esparsa Canyon, and M6650, from
the valley below the Lower Circus at 850 m. It is now widespread, from Northeast An-
chorage to the pine ridge to El Picacho and Arroyo Melpomene. In good years it is
common in such places as about the principal spring and in the valley below the Lower
Circus. It could scarcely have been so common in earlier years without having been found
before. Possibly it was rare before and has spread with the drastic changes in the vegetation
of the last century. However, I list it with those west American plants possibly introduced
on Guadalupe Island (Table 2).

Pectocarya recurvata |. M. Johnst., Contr. Arnold Arb. 3:97. 1932.

Pectocarya penicillata of Guadalupe Island references.
RANGE. — SE California and Nevada to Sonora and northern Baja California.

Palmer collected this plant with Harpagonella palmeri, which he found only at the “south”
end, in low valleys (Watson 1876:118). In 1958 Dr. I. M. Johnston wrote me that Palmer’s
specimen and Anthony 237, the only collections of Pectocarva he had seen from Guadalupe
Island, were both P. recurvata. He also verified my collection (4/6594) from the bottom
of Esparsa Canyon, where it was fairly common in 1958. I had also found a single plant
on the northeast slope of El Picacho at 550 m (M6467). I have not since found it on the
island and wonder whether this is just an accident or whether P. recurvata has declined
at the same time that P. linearis ferocula seems to have been increasing.

Plagiobothrys acanthocarpus (Piper) I. M. Johnst., Contr. Arnold Arb. 3:33. 1932.

Allocarya acanthocarpa Piper, Contr. U.S. Natl. Herb. 22:87. 1920.
RANGE. — Central California to NW Baja California; Santa Cruz Island.

Apparently this plant was collected on Guadalupe Island only in 1958: shallow valley
below Lower Circus [?] (Wiggins & Ernst 107); uncommon on dry ground on a gentle
slope northeast of principal spring at 975 m (M6659). (My recollection was that Wiggins
and Ernst collected the plant the same place I did.) This seems an improbable place for
a plant usually found in vernal pools. Remarkably, Palmer, Greene, and Franceschi not
only had visited the area but actually had camped there without finding it — though then
there were even shallow pools in the area, which would seem more hospitable to it!
However, this is a plant that in some places appears only sporadically, apparently de-

REID MORAN

pending on rainfall. I list it (Table 2) with those west American plants possibly introduced
on the island.

Plagiobothrys collinus (Phil.) 1. M. Johnst. var. californicus (A. Gray) L. C. Higgins,
Great Basin Naturalist 34:163. 1974.

Plagioboihrys californicus (A. Gray) Greene, Bull. Calif. Acad. Sci. 2:407. 1887.
RANGE. — Central California to NW Baja California.

Howell (1942:152) found this plant only near the barracks. It grows now also in the
shallow valley below the Lower Circus and on the plateau near the spring, in some years
very scarce but sometimes common. In the wet year of 1978 I also found some with
smaller flowers as in var. gracilis (I. M. Johnst.) L. C. Higgins, the dried corollas 1.5—6
mm wide, varying from plant to plant. The few mature nutlets seen are small, + 1.25 mm
long.

Like P. acanthocarpus, this plant now grows where early collectors would probably have
found it had it been there then, though the vegetation was then very different. The question
again 1s whether it could have spread from some other part of the island. However, | list
it (Table 2) with those west American plants probably recently introduced.

Cactaceae
Note. — For Pachycereus pringlei see Excluded Species, p. 168.

Mammillaria blossfeldiana Boed. var. shurliana (H. E. Gates) Wiggins in Shreve & Wig-
gins, Veg. Fl. Sonoran Des. p. 1030. 1964.

Mamillaria or Neomammillaria goodridgii or goodridgei of Guadalupe Island references.
RANGE. — West central Baja California; Guadalupe Island.

Greene (1885:223) found a single specimen, of fair size and in flower, toward the south
end of the island. On his second trip Palmer found it common at the south end (Vasey
and Rose 1890:24). It is widespread over the mesa at the south end and is on Islote Negro
(Plate 1A) and Outer Islet but is rather scarce in the northern part of the island: cliff on
north rim of Barracks Canyon at the mouth, 450 m (A/6/22).

Lindsay compared living plants from the island with some from the type locality of M7.
blossfeldiana var. shurliana and concluded that they were essentially similar (Lindsay
and Dawson 1952). He did note, however, that the island plant had pitted rather than
smooth seeds, which sounds like an important difference.

Opuntia prolifera Engelm., Amer. Jour. Sci. Ser. 2, 14:338. 1852.

RANGE. — Southern California to NW Baja California and Isla Cedros: offshore islands
except San Miguel and Todos Santos.

sea on both sides of the island, but especially plentiful near the landing. It occurs from
the northeast part of the island and West Anchorage to the south end, generally at low
elevations but on sunny slopes above Northeast Anchorage to 600 m.

Greene (1885:223) found this cholla rather common on precipitous rocky places near the

GUADALUPE ISLAND FLORA

Campanulaceae

Githopsis diffusa A. Gray var. guadalupensis Morin, Syst. Bot. 8:465. 1983.
Githopsis specularioides of Guadalupe Island references.
RANGE. — Endemic to Guadalupe Island.

Palmer found this plant abundant at the “middle” and north end of the island, under sage
brush and dead branches (Watson 1876:116); he said the flowers were white, turning to
blue after gathering. Greene (1885:225) listed it without comment. It is now very rare
among rocks under pines and oaks on the northeast ridge: in 1955 two plants were found
in fruit (Kuijt & Miller 1006) and in 1958 two in flower (66/3, Wiggins & Ernst 43).

Morin (1983) placed the Guadalupe Island variety in G. diffusa ssp. candida (Ewan)
Morin, whose var. candida grows in the mountains of San Diego County, California. She
described it as shorter than var. candida, usually 4-10 cm tall, with smaller upper leaves,
the internodes very short and so the plant densely leafy, the hypanthium shorter and
+ 2-3 times rather than + 5 times as long as wide.

Triodanis biflora (Ruiz & Pav.) Greene, Man. Bay Reg. p. 230. 1894.
Specularia biflora (Ruiz & Pav.) Fisch. & C. A. Mey., Ind. Sem. Hort. Petrop. 1:17. 1835.

RANGE. — Oregon to Mexico and South America; Santa Rosa, Santa Cruz, and Santa
Catalina Islands.

Palmer found this plant rare, in the shade of rocks and sagebrush on hillsides in the
“middle” of the island (Watson 1876:116). It is still uncommon, among rocks under pines
and oaks on the upper northwest slope and also near the barracks and on the grassy slopes
above.

Caryophyllaceae
Note. — For Polycarpon depressum see Excluded Species, p. 169.
*Cerastium glomeratum Thuill., Fl. Paris, ed. 2, p. 226. 1799.
RANGE. — Native to Europe; a widespread weed in North America.

This weed was first found on the island in 1958, on the pine ridge (Wiggins & Ernst 35).
It is now widespread there and on the northwest slope of the island, sometimes lush and
abundant under the trees, and is fairly common to the spring and the slope above the
barracks at Northeast Anchorage.

*Herniaria cinerea DC., Fl. Franc. Suppl. p. 375. 1815.

RANGE. — Native to southern Europe; a rather uncommon and sporadic weed in western
North America.

This weed was first found on the island in 1958: on four trips in 1957 | had not happened
to see it. In April 1958 we found it at a half dozen places scattered from the principal
spring to the mouth of Arroyo Melpomene, mostly just one or a few plants at each place.
Only on the slopes of El Picacho was it more common, and that has remained its main
stronghold on the island. In 1970 it was common on the northeast side from 500 m to

REID MORAN

PLATE |
\. Mammillaria blossfeldiana var. shurliana (M6708A) on Islote Negro, 28 April 1958
B. Flowering branch of the endemic Hemizonia frutescens (M29S807) on cliff at the north end at
800 m, 21 August 1981
C. Flowering branch of the endemic H/emizonia greeneana ssp. greeneana (6/43) on the south
end mesa at 100 m, 28 October 1957
D. Young plant of the endemic Hemizonia palmeri on the south end mesa at 100 m, 27 April 1958
E. Flowers of the endemic Dudleya guadalupensis on plant from Islote Negro, grown in San Diego,
6 June 1957
F. The endemic Lupinus niveus in arroyo on south end mesa, 29 April 1958

GUADALUPE ISLAND FLORA

PLATE 2

G. Flowers of the insular endemic Lavatera occidentalis (left) and the Guadalupe endemic L. lindsayi
on Outer Islet, 11 February 1957

H. The rare endemic Camissonia guadalupensis ssp. guadalupensis in the bed of Arroyo Melpomene
at 450 m (Moran & Ernst 6737), 29 April 1958

I. Young plant of the endemic Eschscholzia palmeri on Islote Negro, 19 April 1957

J. Flower of the endemic Cistanthe guadalupensis on plant from Islote Negro grown in La_ Jolla
30 March 1957

K. The rare insular endemic Mimulus latifolius (Moran & Ernst 6738) in the dry sandy bed of Arroyo
Melpomene at-450 m, 29 April 1958

L. The endemic Triteleia guadalupensis; a plant of the type collection (M/2063), from the east base
of El Picacho, flowering in San Diego 28 April 1973

REID MORAN

the summit and on the south side. Otherwise, it still seems to be scattered.

+Silene antirrhina L., Sp. Pl. p. 416. 1753.

RANGE. — British Columbia to NW Baja California and the eastern United States; San
Miguel, Santa Rosa, Santa Cruz, Santa Catalina, San Clemente, and Todos Santos Islands.

Palmer found this plant only in a canyon on the east side, near the beach (Watson 1876:113).
No one has found it since, and very likely it is extinct on the island. In other places,
however, it appears very irregularly, in some areas commonly following fires. Steve Junak
tells me he found it in 1995 on San Miguel Island, where it was seen before only by
E.L. Greene in 1886.

*Silene gallica L., Sp. Pl. p. 416. 1753.
RANGE. — Native to Europe: a widespread weed in North America.

Palmer found this weed occurring sparingly in the “middle” of the island, in level open
spots (Watson 1876:113). It is now widespread and often common on the northwest slope
and in the northeast part of the island and less common to the south end.

Spergularia macrotheca (Hornem.) Heynh., Nom. 2:689. 1846.

Lepigonum macrothecum (Hornem.) Fisch. & C. A. Mey., Ind. Sem. Hort. Petrop. 3:14. 1837.

Tissa macrotheca (Hornem.) Britton, Bull. Torrey Bot. Club 16:129. 1889.

Tissa talinum Greene, Erythea 1:106. 1893.

Spergularia macrotheca (Hornem.) Fisch. & C. A. Mey. var. talinwn (Greene) Jeps., Fl. Calif.
1:493. 1914.

Tissa pallida of Guadalupe Island references

RANGE. — Washington to NW Baja California: islands of southern California and Los
Coronados.

On his second trip Palmer found this plant common on exposed sides of hills, in arroyos,
and on sides of canyons at the south end (Vasey and Rose 1890:23). Franceschi (1893¢:134)
saw it only on a perpendicular cliff on the right bank of the canyon not far from the
landing, growing there in numbers. In the northern part of the island it survives only on
cliffs, though sometimes locally common: northwest cliff at 600 m (A// 7280); Barracks
Canyon at 480 m (1/7832); Esparsa Canyon at 300 m (6/26). At the south end it grows
not only on seacliffs and on Islote Negro and Outer Islet but also commonly on flat areas
on the mesa. At the southwest corner of the island it extends far out onto barren lava
often exposed to salt spray (Fig. 45).

Greene (1893) based Tissa talinum on the collections of Palmer and Franceschi from
Guadalupe Island (type: Palmer 864a, ND, UC, US). He said it was most unlike all others
in its almost obsolete internodes and consequent dense leafiness: and he described the
seeds as minute and wingless. To Spergularia macrotheca var. talinum, Jepson (1915:493)
also referred specimens from San Clemente Island.

Rossbach (1940:70-76) discussed in detail the great variation in S. macrotheca, seen even
in plants growing side by side. She wrote that at first there seemed some reason to keep
Tissa talinum of Guadalupe Island as an entity because of its extremely short-internoded
habit and crowded, spreading, fleshy leaves, with a leafless strict inflorescence standing
definitely above the foliage (Fig. 46; also see Moran 1959, fig. 20). As she then pointed

GUADALUPE ISLAND FLORA

FIGURE 45. A

FIGURES 45 and 46. Two forms of the polymorphic Spergularia macrotheca

prostrate form (44/3758), on barren lava near the upper limit of salt spray at the southwest corner

of the island, 30 April 1967. FIGURE 46. A form with crowded succulent leaves and with flowers

raised above the leaves (75659), in the crater of Outer Islet, 1] February 1957. E. L. Greene named

this form as the endemic 7issa talinum.

~~
nN

REID MORAN

out, however, not all Guadalupe specimens have this habit: she mentioned variation in
inflorescence and in size, shape, and pubescence of leaves. Later collections show more
variation. In plants of shaded cliffs the stems are pendent and to 7 dm long — longer than
she described for the species as a whole. Internodes are to 13 mm long. Leaves are 1-8
cm long, whereas she described them for the species as to 5 cm long. She described the
petals for the species as rosy to light pink and perhaps occasionally white but noted that
she had never seen any absolutely white in the field; but in five of my Guadalupe col-
lections the color is noted as white and in only one noted as tinged lavender. Rossbach
then wrote that the seeds of Guadalupe plants measured 0.5—0.7 mm long, whereas those
of S. macrotheca are usually 0.7—-0.9 mm long but may be only 0.6 mm; in plants of San
Clemente Island she found the seeds 0.6—0.9 mm long. She concluded that although some
Guadalupe plants have no match in habit elsewhere, 7issa talinum is not consistently
distinct in any character, and she gave it no taxonomic recognition.

For the plants of San Clemente Island, Raven (1963:321) found that those collected late
in the season tend more to approach the form of 7. ¢alinum. He suggested that the complex
needed more study, especially in the experimental garden, since the plants vary so widely
in their ecology in nature and probably also show seasonal variation. My seed measure-
ments of more recent Guadalupe collections agree with Rossbach’s.

Spergularia marina (L.) Griseb., Spicel. Fl. Rumel. 1:213. 1843.

RANGE. — British Columbia to NW Baja California and east coast; Santa Rosa, Santa
Cruz, Anacapa, San Nicolas, Santa Barbara, Santa Catalina, and San Clemente Islands:
also Eurasia.

Howell (1942:149) was first to report this plant, finding it only on the trail to the cypress
erove. It has since been found only in the northeast part of the island, as at Northeast
Anchorage, near the principal spring, and in the valley below the Lower Circus, sometimes
locally common. Because earlier collectors did not find it, the question is whether it could
have spread from some other part of the island with the drastic changes at the end of the
last century. Raven (1963:298) noted it as possibly introduced on Guadalupe, and I list
it with those west American plants probably introduced (Table 2).

Stellaria nitens Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:185. 1838.

RANGE. — British Columbia to Utah and NW Baja California; Santa Rosa, Santa Cruz,
and Santa Catalina Islands.

Palmer found this plant among rocks on hillsides in the “middle” and at the north end
of the island (Watson 1876:113). Greene found it only under oak trees at the north end.
It is now rather scarce, but in small colonies, in open areas from the northwest slope to
the cypress grove, at 600-1150 m.

Chenopodiaceae

Aphanisma blitoides Nutt. ex Mog. in A. DC., Prod. 13°:54. 1849,

RANGE. — Coastal southern California to west central Baja California and offlying islands,
except San Miguel.

First to find this plant on Guadalupe Island was Brandegee (1900:22), without specific

GUADALUPE ISLAND FLORA

locality. It is fairly common on rocky slopes near the shore in the southern part of the
island, from Juniper Canyon and West Anchorage to the south end, and on Islote Negro
and Outer Islet. It seems remarkable that Palmer did not find it at the south end in 1889,
and I wonder if it could possibly be a late arrival.

Atriplex barclayana (Benth.) A. Dietr., Syn. Pl. 5:537. 1852.

Atriplex palmeri 8. Watson, Proc. Amer. Acad. Arts Sci. 11:146. 1876.

Atriplex dilatata Greene, Pittonia 1:264. 1889.

Atriplex rosei Standl., N. Amer. Fl. 21:60. 1916.

Atriplex barclayana (Benth.) A. Dietr. ssp. palmeri (S. Watson) H. M. Hall & Clem., Carnegie Inst.
Wash. Publ. 326:315. 1923.

Atriplex barclayana (Benth.) A. Dietr. ssp. dilatata (Greene) H. M. Hall & Clem., Carnegie Inst.
Wash. Publ. 326:315. 1923.

RANGE. — Baja California, Sonora, and Sinaloa.

Palmer found this plant only at the “south end” of the island (No. 83; Watson 1876:119).
He called it one of the three characteristic perennials of the island, much more frequent
than Artemisia californica but scarcely half so abundant as Franseria [= Ambrosia| cam-
phorata. It is still very common in the southern half of the island below 600 m (Fig. 32),
which is about the level at which the goats totally destroy it. It 1s also on Islote Negro
and Outer Islet (Fig. 35). At the south end in August 1970 T noted this and Hemizonia
greeneana as the only plants looking much alive at that season.

Watson based 4. palmeri on Palmer 83 of 1875 (GH). Greene based 4. dilatata on speci-
mens from the San Benito Islands. Standley based 4. rosei on part of Rose 16022
(US638110), from Guadalupe Island, March 2, 1911.

In their careful revision of Atriplex, Hall and Clements (1923:313) stated: “Eight species
have been described from what is here included under 4. barclavana .. .. After assembling
all of the available material, it seems doubtful if any of these, with the possible exception
of A. lurida, can be advantageously retained even as subspecies. Much individual variation
is apparent and single plants very frequently have fruiting bracts of two or more of the
‘species’. However, the differences between the extremes are so striking that it seems
desirable to give some kind of taxonomic recognition to the more frequently recurring
forms.” Accordingly, they recognized six subspecies, mostly with considerably overlap-
ping ranges. Plants from the southern part of Guadalupe Island they referred to two sub-
species: ssp. dilatata (including A. rosei) and ssp. palmeri. Howell (1942:148) noted some
variation in habit and vesture, and considerable variation in size and shape of fruiting
bracts, but regarded all five of his collections as belonging to a single variable entity. |
agree. Whatever the ultimate best treatment of this species, it does not seem feasible to
recognize two subspecies occurring together on Guadalupe Island.

oAtriplex californica Mog. in A. DC., Prod. 132:98. 1849.

RANGE. — Central California to NW Baja California and Isla Cedros; islands of southern
California.

Howell (1942:48) found this plant on an ocean bluff near Northeast Anchorage. Moran
(1951:54) found it on the north outer slope of Outer Islet, where also it has been seen
later. These are the only known localities on the island. Because earlier collectors did not
find it at the principal landing, the question is whether it was a relatively recent arrival;
and | list it as possibly so (Table 2).

REID MORAN

Note. — *Chenopodium ambrosioides L., “Epazote”, grown in a pot at Campo Sur in
1971 (M29799), seemed a potential weed, as on Isla Socorro.

*Chenopodium murale L., Sp. Pl. p. 219. 1753.
Chenopodium album of Guadalupe Island references?
RANGE. — Native to Europe, a common weed in North America.

Greene (1885:227) found a few plants of this weed near the landing and said it was
evidently a newcomer. It is now widespread on the island, from the pine ridge and north-
west slope to West Anchorage and Islote Negro, in the northern part of the island sometimes
locally common.

Palmer found one plant near the sea on the east side which Watson (1876:227) reported
as C. album L., but according to Wallace (1985:101) the NY specimen is C. murale.
Greene (1885:227), while listing C. mura/e as a new arrival, also listed C. a/bum without
comment; and his specimen, if any, has yet to be checked. Eastwood (1929:403) cited a
collection of Hanna and Slevin as C. a/bum, but I have not found this at CAS. If C.
album ever has been on the island, it clearly has not become well established and persisted
there.

Suaeda taxifolia (Standl.) Standl., Field Mus. Bot. Ser. 8:10. 1930.

Suaeda californica 8. Watson var, pubescens Jeps., Fl. Calif. 1:447. 1914.
Suaeda californica, S. fruticosa, and S. torreyana of Guadalupe Island references

RANGE. — Southern California and Baja California; islands of southern California and
also Los Coronados and San Martin.

Palmer saw suaeda on his second trip: Vasey and Rose (1890:22) mentioned it among
plants that “have very recently gained a foothold here,” although they did not include it
in the list of plants collected. It is fairly common at West Anchorage on the flat behind
the beach at 10 m and at Campo Sur on the talus at the base of the cliffs and out onto
the barren lava flat to the west.

Compositae
+bAgoseris heterophylla (Nutt.) Greene, Pittonia 2:178. 1891.
Troximon heterophyllum (Nutt.) Greene, Bull. Torrey Bot. Club 10:88. 1883

RANGE. — British Columbia and Idaho to NW Baja California; Santa Rosa Island.

is no further record of this plant from the island, and presumably it is extinct. Greene
wrote that this was “of the ordinary form only”, and Quentin Jones annotated Greene’s
specimen of 20 April 1885 (DS) as 4. heterophylla var. heterophylla. Because Palmer
had camped for 15 weeks at the springs and had failed to find this plant and three others
later found there by Greene, it very likely was a new arrival not well established by
Palmer’s time. | therefore list it with the west American plants probably introduced on
the island (Table 2).

Greene (1885:225) found fine large specimens in grassy ground about the springs. There

GUADALUPE ISLAND FLORA
Amblyopappus pusillus Hook. & Arn., Hook. Jour. Bot. 3:321. 1841.

RANGE. — Coast and offlying islands of south central California to central Baja California;
also Peru and Chile.

Palmer found this plant in low ground at the “southern end” (Watson 1876:116). It is
often abundant on flats and hillsides at low elevations in the southern part of the island,
including Islote Negro, less common northward, and rare at high elevations in the north.

Ambrosia camphorata (Greene) W. W. Payne, Jour. Arnold Arb. 45:414. 1964.

Franseria camphorata Greene, Bull. Calif. Acad. Sci. 1:192. 1885.
Franseria bipinnatifida of Guadalupe Island references.

RANGE. — Baja California, including Cedros Island, western Sonora, and southern San
Luis Potosi.

Palmer found this one of the most conspicuous plants at the “south end”, especially about
Jacks Bay [West Anchorage], growing in thick roundish clumps about a foot and a half
high on level spots and among rocks, giving the country a greenish white appearance.
He added that it was not relished by goats but that asses were very fond of it. In the
northern part of the island, even if not relished, 4. camphorata is now rare and confined
to cliffs: north rim of Barracks Canyon near the mouth at 370m (M/7842); Barracks
Canyon at 300 m (M7828); north side of Esparsa Canyon at 300 m (7/6627); north side
of South Esparsa Canyon at 280 m (4/6607). In the southern half of the island it is dominant
in many areas below 500 m. On the southeast coast it is by far the commonest perennial,
giving a grayish color to the landscape. On the northeast flank of El Picacho it extends
to about 500 m, and just to the north I saw a few seedlings to 600 m. In this part of the
island in December 1957 it was little browsed by goats at sea level but showed more and
more damage at higher elevations, the upper limit perhaps being the level of total de-
struction. Similarly, Howell (1941b:40) told how in coming south from the cypress grove
he first found the dead remains of franseria, then a few live stumps, and still farther south
[and lower] some shrubs a foot or two tall. However, on the uplands south of El Picacho,
at about 500 m, some abrupt discontinuities on comparatively level places apparently are
due to differences in soil. From there south on the Melpomene drainage, this plant is
abundant down to about 250 m but is rarely seen much below, except for a few at the
mouth of Arroyo Melpomene. Near Plaza de Toros it is common on the east slope of the
island but stops abruptly near the divide with the Melpomene drainage. It 1s not found
on the islets.

Greene based the species on his collections from Guadalupe and Cedros Islands. Payne
(1964:415) called the Guadalupe collection the type (holotype CAS; Kk, NY, US).

Ll.

ws)

Artemisia californica Less., Linnaea 6:523. 18

RANGE. — Central California to NW Baja California; islands of southern California and
Los Coronados, Todos Santos, and Cedros.

Palmer found this shrub in considerable abundance at the “south end”, in rocky spots,
giving character to the vegetation, about a foot and a half high, of rather loose habit; also
in the “middle” of the island, in crevices of the highest cliffs (Watson 1876:116). He said
it was immune to goats and [after the ambrosia and atriplex] one of the three most char-
acteristic perennials of the island (Safford MS). Franceschi (1893c:136) found it in con-
siderable numbers on the cliff east of the cabins [1.e. the Lower Circus] and also a mile

REID MORAN

or so to the north. Later collectors have not mentioned it, and it seems to be very rare
now, confined to cliffs in the northeast part of the island, if indeed it survives at all. The
only collection in this century was in 1957, from a single shrub at 300m on a cliff in
Esparsa Canyon (4/6/28); | failed to find it there the next year. In 1981 I tentatively
identified with this species a plant seen high out of reach on a south-facing cliff at
750 m, south of the old lookout and thus high above Northeast Anchorage. This may be
the second place mentioned by Franceschi.

Baeriopsis guadalupensis J.T. Howell, Leafl. W. Bot. 3:153. 1942.
RANGE. — Endemic to Guadalupe Island.

This is a dense woody shrublet to 3 dm high and 9 dm wide, having bright green succulent
leaves and solitary yellow heads 1-3 cm wide, with mostly 8 rays. For a full description
and photographs, one in color, see Moran 1968.

Howell (1942:153) found only a single plant of this species, with Hemizonia palmeri and
Sphaeralcea palmeri, on a rocky flat at the south end of the island. Although rare in such
places, accessible to goats, it is common on Islote Negro (Fig. 47), Middle Islet, and
Outer Islet and in some other places protected from them. In fact, it very likely 1s common
in many such places along the west and north shore, sheltered by high cliffs above. On
rocks occasionally exposed to salt spray, as on the west side of Islote Negro, it grows
down close to the sea where little else will grow. In April 1967 it was luxuriant for
perhaps a half kilometer along the talus of a north-facing seacliff, and up to the edge of
the mesa at 100 m, around the corner just north of Campo Sur (47/3760); but in August
1981 plants there showed severe damage by goats. In 1988 it was thriving there at the
top of the cliff beside the road just above Campo Sur. It also grows at 600 m on ledges
on the foggy sheer northwest cliff of the north ridge of the island (Moran & Philbrick
29592).

With his new species Howell proposed Baeriopsis, a monotypic genus that is thus endemic
to Guadalupe Island. He wrote that its affinities were not evident, but he said the characters
of involucre, achenes, and pappus seemed to relate it to Amblvopappus Hook. & Arm.
Carlquist (1959b, figs. 7, 8) studied the wood anatomy of this plant in comparison with
various other Helenieae. He also made a chromosome count of 7 = 8 based on his collection
468 (CAS, RSA).

*Centaurea melitensis L., Sp. Pl. p. 917. 1753.

RANGE. — Native to the Mediterranean region; a widespread noxious weed in western
North America and elsewhere.

On his second trip Palmer reported this weed as introduced on the north end and gradually
finding its way up the island (Vasey and Rose 1890:25). It is now established at a few
places in the northeast part of the island, as at Northeast Anchorage and near the mouth
of Esparsa Canyon, and on the east slope of El Picacho at 450 m (M16757).

Coreopsis gigantea (Kellogg) H. M. Hall, Univ. Calif. Publ. Bot. 3:142. 1907.
Leptosyne gigantea Kellogg, Proc. Calif. Acad. Sei. 4:172. 1872.
RANGE, — Coast and islands of southern California; Guadalupe Island.

Palmer found only two plants of this species, in the crevices of high rocks (Watson

GUADALUPE ISLAND FLORA

1876:115); and no one has since found it on the main island. Moran (1951:159) reported
it from the sheer cliffs of Inner Islet and common on the floor and inner slope of the
crater on Outer Islet (Fig. 48). In 1982 Robert L. Brownell, Jr., of the U.S. Fish and
Wildlife Service, sent me photographs (SD 110590) of the cliffs about half a mile north
of Pillar Point, on the southeast coast, showing plants which he thought might be
C. gigantea. | could not be sure from the photographs, but the colors of foliage and
flowers strongly suggest Peritvle incana.

+Eriophyllum lanatum (Pursh) James Forbes var. grandiflorum (A. Gray) Jeps., FI.
W. Mid. Calif. p. 524. 1901.

Bahia lanata var., and evidently Eriophyllum caespitosum, of Guadalupe Island
references,

RANGE. —SW Oregon and north central California as far south as Mariposa County;
Guadalupe Island.

Palmer found a single plant that had escaped the goats, on a rocky open spot in the
“middle” of the island (Watson 1876:116). Constance (1937:91) described Palmer’s speci-
men and compared it with other collections, saying it was fairly well matched by such
specimens as H. M. Hall 10147 (DS, UC), from Rocklin, Placer Co. He found nothing
sufficient to separate it from var. grandiflorum. However, Palmer described the flowers
as light orange, whereas the rays of this species are always described as yellow.

Franceschi (1893¢:135) reported Eriophy/lum, a woody perennial on a rock near the cabins.
Probably this is the same, but Dr. Constance was unable to find Franceschi’s specimen
at UC, where presumably it should be.

No one has since found this plant, and it probably is extinct on the island.
Filago arizonica A. Gray, Proc. Amer. Acad. Arts Sci. 8:652. 1873.

RANGE. — Southern California to Arizona, central Baja California, and northern Sonora;
Santa Catalina, San Clemente, and Cedros Islands.

Palmer found this plant on level ground at the “south end” (Watson 1876:115). Howell
(1942:154) found it on the slope of the pine ridge as well as farther south. Mostly it
grows in the southern part of the island, sometimes locally common, but generally little
collected. Dr. James Morefield verified two of my collections in 1992.

Filago californica Nutt, Trans. Amer. Phil. Soc. Ser. 2, 7:405. 1841.

RANGE. — Northern California to Utah, Texas, Sonora, and central Baja California; islands
of southern Callifornia except San Nicolas; also Todos Santos and Cedros.

Greene (1885:224) found a fine growth of this plant about the springs, and Franceschi
(1893c:135) called it very plentiful. It is widespread on the island, from the pine ridge
to Arroyo Melpomene, and is more common than the other species. Dr. James Morefield
verified nine of my collections in 1992.

Because Palmer had camped at the springs for 15 weeks without finding this plant, as
well as three others that Greene found there ten years later, and because it is now wide-
spread and common and impossible to overlook, very likely it was a late arrival and not
well established by Palmer’s time. | therefore list it with the west American plants probably

REID MORAN

GUADALUPE ISLAND FLORA
introduced on the island (Table 2).
Gnaphalium bicolor Biolett, Erythea 1:16. 1893.

RANGE. — Central California to southern Baja California; islands of southern California
and Todos Santos, San Martin, and Cedros.

I first found this plant on the island in 1958. It seems to be local below 400 m on the
east side of the island, in Esparsa Canyon (e.g. 4/3793, M18153) and in Long Canyon
(M/2077). It is rather infrequent in a part of the island where no early botanist but Bran-
degee is known to have collected. Possibly it is a recent arrival, but the evidence is not
convincing.

Gnaphalium stramineum Kunth, Nov. Gen. Sp. 4:85. 1820.

Gnaphalium chilense Sprengel, Syst. 3:480. 1826.
Gnaphalium sprengelii Hook. & Arn., Bot. Beechey Voy. p. 150. 1833.

RANGE. — Washington to Montana, Texas, and northern Baja California and Isla Cedros;
islands of southern California except Santa Barbara and San Clemente.

Palmer collected this with Microseris in the “middle” of the island (Watson 1876:116),
Greene (1885:224) found only one, and Palmer on his second trip found three plants in
an arroyo bed at the south end (Vasey and Rose 1890:24). | have seen only a few plants
in the bed of Arroyo Melpomene near the mouth, in 1958 ((/67/5) and again in 1970
(M17350).

Haplopappus canus (A. Gray) S. F. Blake, Contr. U.S. Natl. Herb. 24:86. 1922.
Diplostephium canum A. Gray, Proc. Amer. Acad. Arts Sci. 11:75. 1876.

Hazardia cana (A. Gray) Greene, Pittonia 1:29. 1887.

RANGE. — Endemic to San Clemente and Guadalupe Islands.

Palmer found this shrub in the crevices of high cliffs (Watson 1876:115). Greene (1887:29)
could find but a single specimen, in a niche some 20 feet above the base of a perpendicular
cliff near the summit of the island. He supposed this shrub the only survivor, and it
seemed in the decline of old age. Franceschi (1893c:135) also saw only one, perhaps the
same one, on the cliff of the Lower Circus near the corral.

Despite Greene’s gloomy words, however, | have seen a score of survivors in the last 40
years, on several cliffs in the northern part of the island: one in 1957 on the north rim
of Barracks Canyon near the mouth at 450 m (M/6/20); nine in 1970 on the south side
of Esparsa Canyon at 250 m (M/8/5/); one in 1970 on the pine pinnacle at the north
end, at 550 m; one in 1971 near the middle of the north end, at 675 m; four in 1981 in
Oak Canyon and on the Lower Circus at 850 m (M29804); five in 1981 on a western
cliff northwest of the cypress grove at 1075 m. However, I have not seen seedlings, and
the small total number still suggests endangerment.

Palmer found the plant in flower March 28, but I found the best flowers in August
(29804) and poorer remnants in April, May, and October.

Gray (1876:75) based the species on Palmer 39, from Guadalupe Island (holotype GH;
F, MO, NY). Although Hall (1928:244) treated Hazardia detonsa (Greene) Greene, of
Santa Rosa and Santa Cruz Islands [and Anacapa], as a synonym, he discussed differences

REID MORAN

and suggested that with more information it might be restored. With more collections,
Raven (1963:343) elaborated the differences and separated the northern plant as Haplopap-
pus detonsus (Greene) Raven.

Hemizonia
Note. — For H. streetsii see Excluded Species, p. 169.

Two species and one subspecies of Hemizonia are endemic to Guadalupe Island. These
three belong to a group informally called Section Zonamra by Keck (e.g. 1960:172), with
six shrubby species having much reduced upper leaves. The others are H. clementina
Brandegee on six of the Channel Islands of California, H. streetsii A. Gray of the San
Benito Islands of Baja California, and H. minthornii Jeps. of the Santa Susana Mountains
and Santa Monica Mountains of southern California. Carlquist (1959a) studied the wood
anatomy of this group in comparison with other Madiinae. And Carlquist (1965:115—118)
discussed and illustrated them as an example of adaptive radiation. Bruce Baldwin is
growing the three Guadalupe species plus H. clementina and H. minthornii and tells me
he has fertile hybrids in all combinations, the three Guadalupe species completely inter-
fertile. On the island, H. frutescens seems geographically distinct, high on the north end.
The other two grow together at the south end, where they look quite distinct and different:
but their flowering times overlap and they do hybridize, though rarely.

Hemizonia frutescens A. Gray, Proc. Amer. Acad. Arts Sci. 11:79. 1876.
RANGE. — Endemic to Guadalupe Island.

Small sparse shrub, with stem to 1.5 cm thick, having thin brown bark splitting lengthwise,
and with branches few, erect to arching or decumbent, slender, 4-8 dm long: or according
to Greene flowering while still herbaceous. Stems sparsely glandular and hirsutulous to
closely hirsute with uniseriate trichomes to 2 mm long, later with axillary cottony tufts;
axillary fascicles starting above during first year, those in upper 10 cm later growing out
as closely glandular branchlets to 5 cm long, each with terminal head and with tertiary
branchlets to 1.5 em long in upper 1—2 cm; these sometimes with a few short quaternary
branchlets. Primary leaves linear, thick-margined, 2-8 cm long, 1-5 mm wide, hirsute and
glandular, mostly with 3-4 ascending teeth; secondary linear, entire, 1-2 cm long, closely
glandular; upper fasciculate, filiform, 2-4 mm long. Flowering mainly in summer; heads
thyrsoid-racemose; involucres 3-5 mm high and wide, short hirsute and glandular; ray
florets 8-9, the ligules obovate-oblong, 2—3-toothed, 34 mm long; disk florets 10-12,
the group surrounded by as many nearly discrete linear bracts; pappus of 5 linear or
subulate fimbriate-denticulate paleae, the disk achenes well-formed but apparently sterile.
Mature achenes?

Palmer found only a few small plants among bushes in the crevices of high rocks in the
“middle” of the island (Watson 1876:115). Greene (1885:224) found it common on level
ground and hillsides, in such places strictly annual, and saw only one suffrutescent plant,
on a precipice. Today it is fairly common, as a small shrub, at 500-800 m on cliffs at
the north end of the island (Fig. 49), from North Point at least to the middle of the north
end. Also, I thought I saw it on westside cliffs at 1075 m, northwest of the cypress grove.
Palmer found abundant yellow bloom on May |. | found no flowers in December, April,
or one May, only one plant with a few flowers another May (//38/3), and all plants in
good flower in August (4129807) (Fig. 49, Plate 1B).

GUADALUPE ISLAND FLORA
Hemizonia greeneana Rose ssp. greeneana
Hemizonia greeneana Rose in Vasey & Rose, Contr. U.S. Natl. Herb. 1:24. 1890.
RANGE. — Endemic to the south end of Guadalupe Island and offlying islets.

Dense compact rounded dark green moundlike shrub to 8 dm high and 2.4m wide, with
arching-decumbent branches (Figs. 33, 50), or in exposed places the plant prostrate; herb-
age glandular-viscid throughout. Branches closely leafy below, only the lowermost leaves
opposite, the middle axils often with fascicled leaves, the upper mostly with leafy
branchlets; ultimate branchlets or peduncles with shorter internodes and fascicled leaves.
Lower leaves often glabrous except for the glands, sometimes sparsely hirsute or villous-
hirsute, 24.5 cm long, 5-8 mm wide, the rachis 24 mm wide, seldom entire, mostly
with 4-12 ascending teeth or lobes; upper leaves gradually smaller, entire, those of the
peduncles linear-oblong, + 5-10 mm long and 0.5—1 mm wide. Flowering throughout the
year but mainly in summer; heads often numerous and subcorymbose on main branches,
solitary or few and cymose on branchlets, 11-18 mm wide, mostly 8-rayed, but some
early ones with more rays and the first, terminating the main branches, with as many as
14: involucres campanulate, 4-7 mm high, 4-6 mm wide, sparsely hirsute, beset with
peg-like glands 0.05 mm long and each tipped with a sticky yellowish globule + 0.1mm
thick, the bracts lanceolate, acute, cymbiform and strongly keeled in lower 3/5 and + 2.5
m wide (flattened), with hyaline margins, narrowed to a flatter thickish apex 2—3 mm
long; ray florets 8-14, the tube 1.5—2.5 mm long, stipitate-glandular, the ray oblong to
obovate, truncate and 2—3-crenate at apex, 4.5-6.5 mm long, 2.5—3.5 mm wide, sub-
glabrous, the style branches slender, 2—3 mm long; receptacular bracts 8-13 (about as
many as rays), in one series, weakly united to middle or above, oblanceolate, narrowly
acute, 56 mm high, + 1.5 mm wide, herbaceous with hyaline margins, glandular, the apex
sparsely long-ciliate; disk florets 8-14 (21), the corollas 4-5 mm long, the tube slightly
ampliate above, subglabrous, the lobes triangular-ovate, 0.7—1.0 mm long, thickened and
densely puberulent ventrally on margins, the anthers yellow, ca. 2.5 mm long, the stigma
lobes 2 mm long, the achenes sterile, 1.7—2.5 mm long, sparsely glandular-puberulent, the
paleae 6-11, 1.3-2.4 mm long, free or united at base, unequal, irregularly lanceolate or
oblong, stiff, dull white, subentire to laciniate. Achenes 2—3 mm long, triquetrous, black,
transversely rugose, acute and stipitate at base, with upcurved beak mostly 2—3 times as
long as thick.

On his second trip, Palmer found this shrub common on the south end, in all the canyons
and arroyos and along the beach: he called it a very homely plant, growing in great
clumps in barren places and the most noticeable plant of the region (Vasey and Rose
1890:24). Because of its size and dark foliage, it is indeed the most conspicuous plant
of the south end; and it flowers abundantly in summer, making a beautiful show when
other plants are dormant. Along Arroyo Melpomene it extends only + 3 km from the south
end, to 350 m. However, in April 1970 I saw what seemed to be Hemizonia, probably of
this species, on the western seacliffs a little farther north. It also grows on the east slope
of the island north of Morro Sur (Junak 2069). It is common on Islote Negro, Middle
Rock, and Outer Islet; and in August 1970 I saw many plants in full flower on the sheer
south face of South Bluff. Thus it is abundant but very restricted at the south end of the
island.

The ssp. peninsularis Moran (1969:286) is a slightly taller and mostly more open shrub,
with mostly virgate branches, generally shorter and stiffer trichomes, somewhat longer
and relatively narrower lower leaves with fewer teeth or lobes, more and slightly longer

REID MORAN

FIGURE 49. The endemic Hemizonia frutescens (M29807) on cliff at the north end, 800 m,

21 August 1981

The endemic Hemizonia greeneana ssp. greeneana at the south end—old shrubs,

FIGURE 50
This shrub is local but abundant on the south end

showing the arching branches; 27 April 1958
and its islets and makes a beautiful show with yellow flowers in summer, when other plants are

dormant

GUADALUPE ISLAND FLORA

pappus paleae, and generally shorter beak on the achene; however, both are variable. It
is very local in northwest Baja California, on the Todos Santos Islands, on Punta Banda,
and on north-facing cliffs at the mouth of the Rio San Miguel. Brandegee (1899) first
reported the Todos Santos Island plant as seeming the same as specimens from Guadalupe
Island that were named as H. frutescens but that did not exactly agree with the description
of that species; later (1900) he called the Todos Santos plant H. greeneana.

Rose described the heads of H. greeneana as having 8 rays and 8-10 disk florets, as in
H. frutescens and H. palmeri, and at most seasons they do (Plate 1C). However, early
heads, ending the main branches, may have |2—14 rays, a corresponding number of in-
volucral and receptacular bracts, and a correspondingly large number of disk florets, some-
times as many as 21. These early heads resemble the usual heads of H. clementina Bran-
degee, of the Channel Islands, which regularly have 12—14 rays and 15-30 disk florets
but which differ in having 2 series of receptacular bracts. As the leaves of the two also
are similar, H. greeneana ssp. peninsularis in early flower has been mistaken for H.
clementina. Of the other two species of sect. Zonamra, H. streetsii A. Gray, of San Benito
Islands, has 11—31 rays and 50 or more disk florets, and H. minthornii Jeps., of the
southern California mainland, has 8 rays, 18—23 disk florets, and receptacular bracts in
+ 3 series. | wonder about the number of rays in first heads.

Hemizonia palmeri Rose in Vasey & Rose, Contr. U.S. Natl. Herb. 1:24. 1890.
RANGE. — Endemic to the south end of Guadalupe Island.

Flattened shrub, 0.5—1.5 (—2.5) dm high and 3—6 (—8) dm wide, the woody main stem to
2.5 cm thick; herbage with abundant white silky pubescence. Leaves numerous, opposite
below, linear to narrowly oblanceolate, mostly entire, silky strigose, not at all viscid,
12 cm long, 1.5—3 mm wide. Flowering in winter and spring; heads numerous, somewhat
corymbose; involucral bracts 44.5 mm long, each entirely surrounding achene, hyaline
margined; rays 8, the corolla 4.5 mm long including tube, the tube + 0.7 mm long, villous;
receptacular bracts 8, very weakly united + to middle, the divisions linear-acuminate,
with prominent midvein, pilose to base, the apex ciliate; disk florets 10-12, the corollas
3-3.5 mm long, the lobes lightly villous, the disk achenes 1.2—1.6 mm long, flat, sterile.
Achenes |.7—2.2 mm long, the beak short, appressed to body; paleae 6-12, very unequal,
1.1—2.7 mm long. (Plate 1D).

On his second trip Palmer found this a very common plant on the south end of the island,
in all exposed places (Vasey and Rose 1890:24). Though less common than the larger
and more conspicuous H. greeneana, it is well distributed over the mesa, and sometimes
locally common, within 3 km of the south end and up to 200 m; it has not been found
on the islets. Thus it appears to be highly restricted. It is in full flower in March and
April, when H. greeneana is mostly not flowering; and it is completely dormant, looking
almost dead, in summer, when H. greeneana is at the height of flowering. In 1958 |
found two apparent hybrids between the two species, one intermediate (/6694), the other
(M6695) more like H. palmeri.

*Hypochoeris glabra L., Sp. Pl. p. 811. 1753.
RANGE. — Native to Europe; a widespread weed in North America.

Mason found a single plant of this weed in Barracks Canyon in 1925 (Eastwood 1929:417).
It now occurs the length of the island and from shore to summit, rather common on the
north slope and in some canyon bottoms.

REID MORAN
Lasthenia californica DC. ex Lindl., Bot. Reg. 21: sub. pl. 1780. 1835.

Baeria gracilis (DC.) A. Gray, Proc. Amer. Acad. Arts Sci. 9:196. 1874.

Baeria palmeri A. Gray, Bot. Calif. 1:376. 1876.

Lasthenia chrysostoma (Fisch. & C. A. Mey.) Greene, Man. Bot. San Francisco, p. 205. 1894.
Baeria chrysostoma var. palmeri (A. Gray) J.T. Howell, Leafl. W. Bot. 3:152. 1942.

RANGE. — Oregon to Arizona and NW Baja California; islands of southern California
and Los Coronados, Todos Santos, and Cedros.

Palmer found this plant abundant in warm low spots in the “middle” and at the “south
end” (Watson 1876:116), and Howell (1942:152) called it occasional from the middle to
the south end. It is widespread in the southern half of the island and as far north as
Esparsa Canyon, generally not common but in wet years locally abundant and giving
color to some grassy slopes. Ornduff (1966:58) cited the collections of Brandegee and of
Wiggins and Ernst. Concerning the accepted name, see Johnson and Ornduff (1978).

Although Palmer had found this plant before, Brandegee (1900:22), without explanation,
called it recently introduced. However, it grows also on the northern islands in a familiar
pattern, and in all respects it seems like a native on Guadalupe.

Dr. Ornduff, who used Guadalupe seedlings in his hybridization studies (Ornduff 1966),
wrote me in 1959 that in the greenhouse they were distinctive for their tallness, wiry
stems, and general reluctance to shed their achenes even when ripe.

+*Lasthenia coronaria (Nutt.) Ornduff, Univ. Calif. Publ. Bot. 40:76. 1966.

Baeria coronaria (Nutt.) A. Gray, Proc. Amer. Acad. Arts Sci. 19:23. 1883.

RANGE. — Southern California to NW Baja California; Los Coronados and Todos Santos
Islands.

Brandegee (1900:22) listed this plant as “a recent introduction about the usual landing”
(Northeast Anchorage), though he did not explain further. Since he also called B. cali-
fornica a recent introduction, when that seems clearly native, we cannot accept his state-
ment without knowing his thinking. However, this plant has not otherwise been found on
the island, before or since, suggesting that it may then have been a new arrival that never
became established. Since every botanist to visit the island has gone ashore at the landing
where he found this plant, very likely someone else would have found it had it grown
there for long, whether native or not (though it is true that a few rare native plants have
been found there only once). Furthermore, there is no particular reason to think it a likely
native: it has not reached any of the islands of southern California, all closer to the
mainland, though 89 percent of the Guadalupe natives that occur also in cismontane south-
ern California have reached one or more of these islands. Having no reason to disagree
with Brandegee’s statement, I therefore tentatively accept this species as an introduction
that did not become established. Brandegee’s specimen was cited by Ornduff (1966:79),.

+Layia platyglossa (Fisch. & C. A. Mey.) A. Gray, Mem. Amer. Acad. Arts,
Ser. 2, 4:103. 1849.

Layia elegans (Nutt.) Torr. & A. Gray, Fl. N. Amer. 2:394. 1843.

RANGE. — Northern California to central Baja California; San Miguel, Santa Rosa, Santa
Cruz, Santa Catalina, and San Clemente Islands.

GUADALUPE ISLAND FLORA

Both Anthony and Brandegee collected this plant on Guadalupe Island in 1897, with no
specific locality. There is no other record, and probably it is extinct on the island. Wallace
(1985:50) cited Anthony 251 at US. The specimen of Anthony’s collection at UC was
annotated by David Keck in 1934 as L. elegans. On the same sheet is a specimen collected
20 March 1897 by Brandegee, labeled L. platyglossa A. Gray, and checked by Keck.

t+Malacothrix clevelandii A. Gray, Bot. Calif. 1:433. 1876.
RANGE. — California to Utah and extreme northern Baja California; Santa Rosa Island.

Palmer found this plant abundant among rocks and trees in the “middle” of the island
(Watson 1876:116). Greene (1885:225) noted it without comment. In revising this group,
Davis and Raven (1962:261) noted that the occurrence of M. clevelandii on Guadalupe
Island needed to be confirmed by more material and by a chromosome count. Apparently,
however, no one has found it for over a century, and it probably is extinct on the island.

Matricaria matricarioides (Less.) Ch. Porter, Mem. Torrey Bot. Club 5:341. 1894.

Matricaria discoidea DC., Prod. 6:50. 1837.
Chamomilla suaveolens (Pursh) Rydb., N. Amer. Fl. 34:232. 1916.

RANGE. — Alaska to NW Baja California and NE Asia; Santa Rosa, Santa Cruz, Anacapa,
and Santa Catalina Islands.

Palmer found this plant around springs in the “middle” of the island (Watson 1876:116).
It was still there, but rare, in 1958 (Wiggins & Ernst 113). Although it may still turn up
in a good year, quite possibly it is extinct on the island.

The native range of this weedy plant is unknown, and possibly it is introduced on
Guadalupe Island.

+Micropus californicus Fisch. & C. A. Mey., Index Sem. Hort. Petrop. 1835:42. 1835.

RANGE. — SW Oregon to NW Baja California; Santa Rosa, Santa Cruz, and Santa Catalina
Islands.

Palmer found this plant on dry gravelly slopes in the “middle” of the island (Watson
1876:115). No one has seen it since, and it probably is extinct on the island.

Perityle emoryi Torr. in Emory, Notes Mil. Rec. p. 142. 1848.

Perityle grayi Rose, Bot. Gaz. 15:118. 1890.
Perityle californica of Guadalupe Island references.

RANGE. — Southern California and Nevada to Baja California and Sonora; W South Amer-
ica; islands of southern California and NW Baja California except San Miguel and San
Nicolas.

Palmer found this plant scattered through some of the canyons on the east side (Watson
1876:116). It is widespread on the island, from the north slope to the south end and on
Islote Negro and Outer Islet, more common at lower elevations and southward.

From Guadalupe Island, Powell (1974:258) cited collections by Anthony, Carlquist, Mason,
Moran, Palmer, and Rose. He included here Perityle grayi Rose, based on Palmer 44 of
1875 (US?; isotypes GH, NY) from Guadalupe Island.

REID MORAN

FIGURE 51. Old shrub of the endemic Peritvle incana (M13792) hanging from a cliff in Esparsa
Canyon, with Fritz Ohre for scale; 2 May 1967.

Perityle incana A. Gray, Proc. Amer. Acad. Arts Sci. 11:78. 1876.
Nesothamnus incanus (A. Gray) Rydb., N. Amer. Fl. 34:12. 1914.
RANGE. — Endemic to Guadalupe Island.

This is a dense grayish-white tomentulose shrub 1-2 or even 4m wide and sometimes
trailing to 5 m (Fig. 51), the stem soft-woody, to 4.cm or more thick, with naked corymbs
of 10—-SO or more discoid yellow heads.

Palmer found this shrub very common in the “middle” of the island, in the crevices of
high rocks, hanging in massive bunches of yellow bloom (Watson 1876:116). Franceschi
(1893c:136) called it by far the most abundant of all the shrubs still living on the island
and the most likely to survive, as it seemed quite at home on the more precipitous cliffs
and young plants and seedlings were abundant in the crevices of the rocks. It is certainly
the commonest shrub on the cliffs today (Fig. 28); and on cliffs it extends from sea level

GUADALUPE ISLAND FLORA

to Mt. Augusta at 1220 m and the length of the island, from the north end to the islets
off the south end. On the islets it also grows in less steep places, but Greene (1885:216)
did not mention it among the shrubs that in his day still covered the gentler slopes in the
north. Nor does it grow on gentler slopes in the southern half of the island where Ambrosia
camphorata and Atriplex barclayana are abundant despite the goats or at the south end
where hemizonia thrives. A few seedlings of the perityle appear on slopes below the
northern cliffs but persist even briefly only in years of fewer goats. From February to
October and probably in the other months at least some plants can be found flowering,
but the main flowering period is about April to June.

Gray based this species on Palmer’s collection No. 43 (holotype GH; NY, US). This is
the shrubbiest member of its genus, and Carlquist (1959b, figs. 1, 2) studied its wood
anatomy in comparison with various other Helenieae. It is distinctive also for its tomen-
tulose herbage, rather large naked corymbs, and coarsely pubescent achenes, and it is
nearly unique in its discoid heads. Partly because of its shrubbiness, Powell suggested
that it might be a primitive member of the genus, but presumably this is an example of
insular woodiness. For this species Rydberg (1914) proposed the monotypic and thus
endemic genus Nesothamnus, returned to Peritvle by Blake (1926), Everly (1947), Powell
(1974), and most other authors. However, Powell wrote that it did not belong to any of
the related-species groups and thus stood as an anomalous species. He said that it could,
in fact, be recognized as a monotypic genus with little change in the taxonomic naturalness
of the subtribe. He added that it was similar to Pericome in habit and capitulescence
characters and perhaps shared ancestral affinity with that genus. Howell (1942) did rec-
ognize Nesothamnus. However, Powell suspected that P. incana was distantly related to
the P. crassifolia group and was best treated as a member of section Perityle. Based on
M17426 Powell (1974) reported a chromosome count of 1 = 50-57, hexaploid.

Davies (1980) illustrated a plant grown at Kew from seeds from Islote Negro (\//7426).
From seeds from the same source and from later collections, it is also grown now in
southern California for drylands plantings.

Senecio palmeri A. Gray, Proc. Amer. Acad. Arts Sci. 11:80. 1876.
RANGE. — Endemic to Guadalupe Island.

Dense white shrub + | m high and (on cliffs) to 1.5 or 2m wide, much branched from
base, with trunk to | dm thick. Leaves oblong-spatulate, obtuse, pinnately veined, 2—9 cm
long, 0.5—2. cm wide, entire to sinuate-dentate, narrowed at base to a slender petiole.
Inflorescence a terminal pedunculate few to several headed corymbose cyme. Heads yel-
low, on long bracteate peduncles, 10-14 mm high, radiate; involucre white, barely caly-
culate, of 20-30 linear bracts; ray florets 12-18, the rays oval, 8 mm long; disk florets
numerous; achenes sericeous-pubescent.

Palmer called this shrub “white sage” and found it very abundant on many warm spots,
from the “middle” to the north end: about three feet high, of diffuse habit, a very free
and showy bloomer, beginning to flower early in February and maturing in May, when
the air was filled with its downy seeds (Watson 1876:116). Greene (1885:216) found it
still covering the gentler declivity leading to the plateau from the zigzag trail out of the
canyon; he called it really very handsome, a shrub three or four feet high, with snow-white
foliage and fine clusters of yellow blossoms. According to Bryant (1887:271), an orni-
thologist who spent the first three months of 1886 there, a “small white sagebrush with
yellow blossoms” grew in places “throughout the entire length of the island.” On his
second visit, in 1889, Palmer found that the senecio had decreased, spots where there

REID MORAN

FIGURE 52. The endemic Senecio palmeri from cliff of the Lower Circus (446653), 25 April 1958.
Once the commonest shrub of the northeast part of the island, it survives only on a few cliffs and

is becoming rare

were thickets of it before having now only dead plants, with just a few scattering plants
alive. Four years later, Franceschi (1893c:136) found it only on the eastern cliff, where
he saw perhaps three dozen; he called it very conspicuous and much whiter even than
Perityle incana. | have seen it only in Oak Canyon and around the corner to the south
on the cliffs of the Lower Circus (Fig. 52), where despite the whiter foliage it is sometimes
hard to tell at a distance from the much more abundant Peritvle incana. In January 1960
I saw 12 without having time to cover the length of the cliff. In 1974 I found a young
plant in a crevice of the Lower Circus, showing it still does reproduce on the cliff. In
August 1981, with older eyes and no field glasses, I failed to identify any, but I am
hopeful that they still survive

Carlquist (1962, figs. 26, 27) studied the wood anatomy of this plant in comparison with

various other Senecioneae, based on his collection 260.
*Sonchus oleraceus L., Sp. Pl. p. 794. 1753.
RANGE. — Native to Europe; a common weed in North America.

Palmer found this weed very rare, on warm slopes in the “middle” of the island (Watson
1876:116). Greene (1885:225) reported it now very common on the eastward slope of the
island. It now extends the length of the island, from the north slope to the south end and
on Islote Negro, in some northern areas fairly common.

GUADALUPE ISLAND FLORA
*Sonchus tenerrimus L., Sp. Pl. p. 794. 1753.
RANGE. — Native to Europe; a weed in the SW USA and Mexico.

On his second visit Palmer found this plant in shady canyons at the south end, not common
(Vasey and Rose 1890:25). It is now occasional near the shore in the southern half of
the island.

Stebbinsoseris heterocarpa (Nutt.) K. L. Chambers, Amer. Jour. Bot. 78:1024. 1991.

Microseris heterocarpa (Nutt.) K. L. Chambers, Contr. Dudley Herb. 4:286. 1955.
Microseris lindleyi of Brandegee’s reference.

RANGE. — California and NW Baja California; Santa Rosa, Santa Cruz, Anacapa, Santa
Catalina, and San Clemente Islands.

Brandegee (1900) reported this plant from Guadalupe Island without comment, and Cham-
bers (1955:288) cited his specimen. It seems to be local on the east side of the island
just south of the middle, generally rather scarce but noted in 1970 as fairly common
locally on the northeast slope of El Picacho at 525 m (M/738/) and as occasional in the
bed of Long Canyon at 50 m (M/7362).

+Stephanomeria diegensis Gottlieb, Madrono 21:476. 1972.

RANGE. — Southern California and NW Baja California; Santa Rosa, Santa Cruz, Santa
Catalina, San Clemente, Los Coronados, and Todos Santos Islands.

This plant is known from Guadalupe Island only from a collection (no. 46, UC) by
W. W. Brown in 1906. Gottlieb (1971) showed a dot for the island on his map for this
species. He wrote me that the Brown collection at UC is a poor specimen whose long
peduncles suggest S. exigua Nutt. ssp. deanei (J. F. Macbr.) Gottlieb, to which S. diegensis
is closest. Dr. Reed Rollins wrote me in 1988 that he did not find a specimen at GH.

Stephanomeria guadalupensis Brandegee, Zoe 5:164. 1903.
RANGE. — Endemic to Guadalupe Island.

Perennial with milky-orange latex, closely white tomentose, sometimes subglabrescent,
usually several branched from a woody perennial base to 4 cm thick and with corky bark,
the stems often decumbent at base, thence erect, unbranched and leafy below inflorescence,
to | or even 1.5 m tall and to 2.5 cm thick at base. Leaves oblanceolate to spatulate, acute
to rounded at apex, attenuate below to a broad petiole, entire to sinuate-dentate or often
deeply pinnatifid into a dozen narrow segments, the lower leaves 0.5—3 dm long, 2—9 cm
wide, the upper gradually smaller and becoming entire. Panicle viscid, 1-6 dm tall, 4-20
cm wide, of many ascending racemose branches with heads more crowded at tips, flowering
August to October. Involucres 10-11 mm long, glandular, somewhat wooly at apex, with
graduated small outer bracts, 5—8-flowered (Fig. 54). Ligules deep pink, + 8 mm long and
3—4 mm wide; anthers 4-S mm long. Achenes 4 mm long; pappus 4-6 mm long, of 14—19
bristles, plumose nearly to base.

Brandegee collected the type March 26, 1897, in Sparmanns Canyon (perhaps Esparsa
Canyon?), where the clumps of white leaves were very conspicuous on the nearly per-
pendicular dark colored cliffs. The inflorescence with fruiting heads that he collected was
a remnant from the previous season (holotype UC92373!). Though not so common as
perityle and galvezia, this showy plant grows on cliffs throughout the island, from the

REID MORAN

FiGuRES 53 and 54. The endemic Stephanomeria guadalupensis. FIGURE 53. Colony on Outer
Islet, 21 August 1970. FIGURE 54. Flower head (x 4.5) of plant from Esparsa Canyon (6/35)
grown in Berkeley, 26 November 1960

GUADALUPE ISLAND FLORA

FiGuRE 55. Young branch of Stephanomeria guadalupensis growing up to become a floral stem:
the condensed form on Outer Islet (!//5//9), with subentire leaves; 21 June 1968

north end to Outer Islet (Fig. 35). Earlier collectors must have seen it but would not have
found it flowering in spring.

On shaded cliffs in the northern part of the island, lower and middle leaves are large and
deeply pinnatifid and tend to be glabrescent; for a photograph, see Carlquist (1965, fig.
14-6). On Outer Islet the plant has a very different aspect: more compact, more densely
tomentose, with leaves subentire to sinuate-dentate (Figs. 53, 55). This form has been
grown at the Santa Barbara Botanic Garden. Plants of exposed seacliffs on the main island
(Howell 8175, M6124) are much like those of Outer Islet.

Carlquist (1960) studied the wood anatomy of this plant in comparison with other
Cichorieae.

Uropappus lindleyi (DC.) Nutt., Trans. Amer. Philos. Soc. II, 7:425. 1841.

Microseris linearifolia (Nutt.) Sch.-Bip., Pollichia 22—24:308. 1866.

REID MORAN
Microseris lindleyi (DC.) A. Gray, Proc. Amer. Acad. Arts Sci. 9:210. 1874.

RANGE. — Washington to Texas and NW Baja California to Isla Cedros; islands of south-
ern California and NW Baja California.

Palmer found this plant only in the “middle” of the island, on stony ridges, eaten close
by goats (Watson 1876:116), and Chambers (1955:285) cited his collection. Greene
(1885:225) called it abundant and very rank about the springs and the cypress groves,
“where the goats do not now range.” It is now widespread in the northern part of the
island and as far south as El Picacho, generally scarce but sometimes locally common
on grassy slopes.

Convolvulaceae

Note. — For Cressa truxillensis var. vallicola, see Excluded Species, p. 169.

Calystegia macrostegia (Greene) Brummitt ssp. macrostegia

Convolvulus macrostegius Greene, Bull. Calif. Acad. Sci. 2:208. 1885.

Volvulus macrostegius (Greene) Farw., Amer. Midl. Naturalist 12:130. 1930.

Convolvulus occidentalis A. Gray var. macrostegius (Greene) Munz, Man. S. Calif. Bot. p. 387.
1935.

Calystegia macrostegia (Greene) Brummitt, Ann. Missouri Bot. Gard. 52:214. 1965.

Convolvulus occidentalis of Guadalupe Island references.

RANGE. — San Miguel, Santa Rosa, Santa Cruz, Anacapa, Santa Catalina, Todos Santos,
San Martin, and Guadalupe Islands; intergrading elsewhere with other subspecies.

Palmer found this plant in the crevices of high rocks, hanging down six feet or more,
continuing to bloom from March through the summer, a thousand flowers seen on a single
plant (Watson 1876:118). Greene (1885:208, 216) found it in crevices of basaltic cliffs
and also spreading its long trailing branches abroad among the rocks of declivities at
lower elevations, where he counted it among the most conspicuous plants. Franceschi
(1893c:137) said it was highly relished by goats but was still keeping its hold on the
most perpendicular cliffs, where its drooping green masses formed a striking contrast to
the silvery foliage of Peritvle incana. Moran (1951:159) found it on the inner slope of
Outer Islet. It is still conspicuous on cliffs, such as the Lower Circus, in the northern part
of the island, and on the southeast coast, often trailing to 25 feet. Seedlings often appear
on slopes below the cliffs and elsewhere and in years of few goats may persist, as in the
arroyo at Northeast Anchorage in 1965. Southward, in some years, it forms occasional
large masses in the bed of Arroyo Melpomene (Fig. 56).

Cuscuta corymbosa Ruiz & Pav., Fl. Peruv. 1:69. 1798.
RANGE. — Southern Baja California and central Mexico to Peru.

Stems pale yellow. Inflorescence axillary, of 1-5 flowers, with some bracts; pedicels 24
mm long. Calyx 34.5 mm wide, 2—2.5 mm long, gamosepalous, the lobes broadly rounded,
entire, + 2 mm wide. Corolla white, ovoid, + 5—6 mm long, the segments ovate, rounded
at apex in fresh material examined, + 1.25 mm long and so + one-fourth tube. Scales
narrow, dentate, extending above middle of corolla tube. Filaments short, the free part
+ 0.5 mm long; anthers + 0.4 mm long. Styles 2-3 mm long, the stigmas capitate. Cap-
sules depressed globose, umbilicate at top, dehiscing irregularly at base, capped at apex
by withered corolla. Seeds + 3-4, + 1.5 mm wide.

GUADALUPE ISLAND FLORA

\
L. satias oat

FIGURE 56. Calystegia macrostegia ssp. macrostegia (M12071) in the bed of Arroyo Melpomene
at 300 m, 5 March 1965, a year of fewer goats.

Known on Guadalupe Island only from one collection: locally common on Sphaeralcea,
Mirabilis, Perityle, etc., at 50m on the lava mesa just north of the fish camp at West
Anchorage, 17 April 1970 (77420). In 1957 I collected twice at West Anchorage, though
farther south and not at this spot, without finding the dodder; and I have not been there
since 1970 to look. Various plants are local on the island, often for obscure reasons; but
it seems remarkable that this parasite has been found nowhere else. It grew near a crude
landing strip used probably for flying out lobsters or abalone and possibly for flying in
foreign plants — though there is no knowing whether planes ever came there, directly or
indirectly, from much farther south where the dodder grows. If it should be found spreading
to other parts of the island, or no longer found at all, that might suggest it was introduced:
but I tentatively treat it as native. Among photographs Meling Lopez left with his specimens
at SD was one of cuscuta, with no locality. Although the species is not identifiable from
the photo, presumably this is of C. corvmbosa on the island, showing it was still there
in the early 1980s.

In the treatment of Yuncker (1932; and note figs. 94F and G) this plant seems more or
less intermediate between vars. grandiflora Engelm. and stvlosa Engelm., both cited from
southern Baja California. Styles are included as shown for var. grandiflora. Wiggins
(1964:1132) called the southern Baja Californian plant var. grandiflora, but in his flora
(1980:378) he called it just C. corymbosa.

Crassulaceae

Crassula connata (Ruiz & Pav.) A. Berger in Engler, Nat. Pflanzenfam. ed. 2,
18a:389. 1930.

REID MORAN

Tillaea erecta Hook. & Arn., Bot. Beechey Voy. p. 24. 1830.
Tillaea minima Gay, Fl. Chil. 2:529. 1847.
Tillaea leptopetala Benth., Pl. Hartw. p. 310. 1849,

RANGE. — Oregon to Argentina and Chile; islands of southern California and NW Baja
California to the San Benitos.

Palmer found two varieties, in large patches in a few exposed clear spots in the “middle”
and at the north end (Watson 1876:115). It is widespread on the island, from the north
slope to Outer Islet, and in some places 1s abundant in good years.

Bywater and Wickens (1984, map 2) showed var. connata and also var. erectoides M.
Bywater & Wickens on Guadalupe Island; and they also annotated some Guadalupe speci-
mens as var. eremica (Jeps.) M. Bywater & Wickens. The four varieties they recognized
for western North America have largely overlapping ranges and characters; and some
differences among the specimens probably result from different temperature and moisture
conditions from place to place and from year to year. After considering the matter in
some detail (Moran 1992), I decided it was not practical to distinguish their varieties.

Dudleya guadalupensis Moran, Madrono 11:154. 1951.
RANGE. Endemic to Guadalupe Island; known only on offlying rocks and islets.

Caudex 1.5—3.5 em thick, to 15 cm or more long, branching to form rounded clumps to
5 dm wide, of 80 or more rosettes. Rosettes subglobose, 3-15 em wide, of 25-40 (—75)
leaves, the leaves pale green to farinose, oblong-oblanceolate, apiculate-acuminate, 2.5—6.5
cm long, 8-13 mm wide, 2-3 mm thick, the base 4-8 mm wide, the margins rounded.
Floral stems glaucous, sinuous, curving down first, then up, 2-3 dm long, 4-7 mm thick,
bare in lower 5 cm, with 15—35 leaves above, the leaves glaucous, ascending, triangular-
ovate, acute, the lowermost 1-2 cm long, 7-10 mm wide. Cyme glaucous, of 2—3 ascending
simple or bifurcate branches, the cincinni 4-8 (—12) cm long, with 7-12 (—17) flowers;
pedicels erect, the lower 3—S (—8) mm long, the upper 2—3 mm long. Flowers April-June.
Calyx 5-6 mm wide, 6-8 mm high, rounded below, the tube 1—1.5 mm long, the segments
narrowly triangular-ovate, acute, slightly unequal, 3—S mm long, 2.5—3.5 mm wide. Corolla
pale greenish yellow but glaucous and appearing almost white, 11-13 mm long, + 4mm
wide at base and 7-11 mm wide above, the petals linear-lanceolate, acute, 22.5 mm wide,
connate 1-2 mm, strongly keeled, nearly straight, erect or ascending and not touching
much above calyx. Stamens erect, the filaments + 9mm long, adnate + 1—2.5 mm, the
epipetalous higher; anthers yellow, + 1 mm long. Nectar glands whitish, | mm or more
wide. Pistils nearly erect but separated, or bowed outward with styles connivent, the
ovaries + 4-6 mm long, the styles at first + 2 mm long, becoming 3-4 mm long. Mature
follicles ascending, the ventral margins + 60—75° above horizontal. Chromosomes: 1 =

17.

This plant was first found on Outer Islet in 1948, the type Moran 2947 (DS324267,
324268). It is fairly common from the inner slope of the crater to the upper valley (Fig.
57). Here the leaves are uniformly light green. In 1957 I found it occasional on Islote
Negro, with a green form (A/5620) and a farinose form (A/56/9) (Fig. 58). Here rosettes
are often solitary or few, though they may form clusters to 4dm wide. In August 1970,
looking from a boat, | saw what clearly was this plant on Middle Rock. In 1955 George
Lindsay wrote me he was sure he saw Dudleva plants on rocks off the north end of the
island, and this species seems to me probable.

Dr. Charles H. Uhl made chromosome counts of the type collection, from Outer Islet,

GUADALUPE ISLAND FLORA

FIGURES 57 and 58. The endemic Dudleyva guadalupensis, found only on offlying islets
FiGuRE 57. Plant on Outer Islet (1/5644), with old dried floral stems and with young floral stems
growing down and then up; 11 February 1957. FIGURE 58. Cincinnus of plant from Islote Negro,
grown in San Diego, 6 June 1957.

REID MORAN

and of three collections (6044, M6706, M6798) from Islote Negro, in all four finding
n=17. Thus the plant is diploid relative to the basic number for the genus (Uhl and
Moran 1953).

Funamoto et al. (1985) reported a somatic count of 27 = 102 from a plant supposedly of
this species and supposedly from Guadalupe Island. However, I have had no response
from the authors about the origin of their plant or whether a voucher specimen is deposited
in any herbaraium where their identification can be checked. Although they seem to imply
that their plants were collected in the wild, it is most unlikely that any of the authors got
to this remote island or, if so, was able to find this plant, known only from two inaccessible
islets off the windward side and south end.

A plant in the nursery trade in Southern California as D. guadalupensis proves to be D.
greenei Rose. In 1990 David Grigsby of Vista kindly gave me plants of it (A/3/032, SD)
that he had as Abbey Garden 69-1168, supposedly field collected. Since neither Charles
Glass nor the supposed collector could verify this information, the source of the plant is
uncertain but probably is not Guadalupe Island, where D. greenei is unknown. For this
collection Charles H. Uhl got a chromosome count of = 51, as in other collections of
D. greenei. Since this count corresponds to the somatic count reported by Funamoto et
al., | suspect that their plants came from a nursery and not from Guadalupe Island. From
this circumstantial evidence, I reject the report of Funamoto et al., based on dubious
material and at variance with Dr. Uhl’s four counts of 7 = 17 based on authentic material.

When the type was collected in April 1948, the rosettes had about 35—70 leaves and were
very compact, the outer leaves tightly folded over the inner (Moran 1951, plate 3). Im-
mature leaves less than 2 cm long were many, making up about half the total; and they
were nearly equal in size rather than grading evenly from youngest to mature, as is usual
in Dudleya. In cultivation the rosettes became more open, with 25-40 leaves grading
more evenly from small to large, and thus appeared more typical for the genus. In later
years, plants on the islets also had this more typical appearance. Thus the plants of the
type collection were somewhat abnormal, presumably because of some unusual growing
conditions. Young floral stems of the type collection became etiolated before they were
unpacked; and although they went on to flower in Santa Barbara, the pedicels were elongate
and the flowers somewhat abnormal (Moran 1951: plate 3, below). From flowering plants
later collected on the islets and from plants flowering in San Diego (Figs. 57, 58; plate
IE), the description above is improved over the original description.

Dudleya virens (Rose) Moran ssp. extima Moran, Haseltonia 3:9. 1995,
Dudleyva hassei, D. virens, and probably Cotvledon of Guadalupe Island references.
RANGE. — Endemic to Guadalupe Island.

The first report of Dud/eva from Guadalupe Island was by Franceschi (1893c:135), who
found only one small plant of Cotvledon ___ on a rock along the trail not far from the
landing. Apparently he brought back the living plant, but if it ever flowered, no specimen
or other record seems to remain. Very likely his plant was of this species, which is rare
on cliffs in the canyon back of the landing of Franceschi’s day.

In 1948 I found a sterile plant of this species (A/29/3) at 800m on what I have since
called Hemizonia Cliff at the north end of the island (Moran and Lindsay 1950, with

photo). It grows mostly on cliffs in the north and northeast parts of the island: one plant
(\//7308) on a rock under oaks on the northwest slope at 700 m; north rim of Barracks

100

GUADALUPE ISLAND FLORA

Canyon, 450 m (M6/23); cliff in Barracks Canyon near confluence of south and west
forks, 420 m (M7830); in shade on north-facing cliffs, Esparsa Canyon at 250 m (M5972
type, SD; M6/3/). Though scarce, it seems to be holding its own despite goats and
botanists, mostly high above their reach.

Dudleya virens is insular (Moran 1995), otherwise growing on San Nicolas, Santa Catalina,
and San Clemente Islands, and on seacliffs at the south foot of San Pedro Hill, a former
island related to Catalina and San Clemente Islands but now connected to the mainland
by the alluvial Los Angeles Plain (see Smith 1900). It is smaller from north to south, the
Guadalupe plant conspicuously different from the robust plant on Point Vicente on the
mainland. There and on Catalina Island (ssp. insu/are), clumps are up to 2 m wide, the
caudex 2-6 cm thick, the rosette leaves 1-3 cm wide, the floral stems 2—7 dm tall and
5—15 mm thick, and the many-branched inflorescence 10-25 cm wide. Plants of San Cle-
mente Island (ssp. virens) are intermediate in size. On Guadalupe Island clumps are ca.
12 dm wide, the caudex I—2.5 cm thick, the rosette leaves 6-15 mm wide, the floral
stems 1—2 dm tall and 2—5 mm thick, the inflorescence 2-12 cm wide.

The two collections from Hemizonia Cliff (29/5, M6436; Fig. 59) have oblong-oblanceo-
late farinose leaves, in what happened to be clustered subsessile rosettes. The collections
from cliffs in the canyons farther south have narrower and more sharply acute green
leaves, and in age the stems are pendent to 3 dm. When M29/5 flowered in cultivation
and turned out to be tetraploid, | placed it with the somewhat larger tetraploid D. hassei
(now D. virens ssp. hassei) of Catalina Island and so reported that species from Guadalupe
Island (Moran 1960). When one of the green narrow-leaved plants (1/5972) proved to be
diploid, I thought it might be feasible to distinguish two taxa on the island, different
cytologically as well as morphologically. But then when the second collection from Hemi-
zonia Cliff (146436) also proved to be diploid, it seemed better, if not entirely satisfactory,
to keep all the plants of Guadalupe Island in one subspecies of D. virens.

Crossosomataceae

Crossosoma californicum Nutt., Jour. Acad. Nat. Sci. Philadelphia, Ser. 2, 1:150. 1847.

RANGE. — Endemic to Santa Catalina, San Clemente, and Guadalupe Islands, and San
Pedro Hill, a landlocked island north of Santa Catalina.

Palmer found only nine shrubs, in the crevices of cliffs overhanging the canyons in the
“middle” of the island, out of reach of the goats and accessible only by the aid of a rope
(Watson 1876:112). Franceschi (1893c:133) saw only one, on the almost inaccessible cliff
of the Lower Circus. Moran (1951:159) found one small shrub on the inner slope of Outer
Islet but saw none on later visits. A few others have survived until recently on cliffs in
the northeast part of the island and possibly are reproducing there to a limited extent, in
Esparsa Canyon, in Barracks Canyon, and on the cliffs of Oak Canyon and the Lower
Circus. However, the plants very likely are decreasing in numbers and so seem threatened
with extinction.

Cruciferae

+* Brassica nigra (L.) Koch in Roehl, Deutsch Fl. ed. 3, 4:713. 1833.
RANGE. — Native to Europe; a widespread weed in North America.

Palmer found this weed in considerable quantity in the “middle” of the island, in open

101

REID MORAN

spots and on the best soil; he said it was eaten by the goats (Watson 1876:113). No one
has found it since, and very likely the goats ate it all. Dr. Reed Rollins wrote me in 1988
that he did not find the Palmer specimen at GH, but Wallace (1985:54) cited one at NY.

+*Brassica rapa L., Sp. Pl. p. 666. 1753.
Brassica campestris L., Sp. Pl. p. 666. 1753.
RANGE. — Native to Europe; a common weed in western North America.

Greene (1885:221) found a few plants near the cabins but added that the species apparently
was not yet well established. No one has found it since, and apparently it never did
become established.

*Capsella bursa-pastoris (L.) Medik., Pflanzeng. 1:85. 1792.

RANGE. — Native to Europe; a cosmopolitan weed.

Brandegee collected this weed on the island in 1897, but Howell (1942:149) was first to
report it. He found it occasional on a rocky slope below the upper corral on the trail to
the cypress grove. It grows in the northern part of the island, mostly at higher elevations,
as on the northwest slope and about the spring, in good years locally common and lush,
less commonly in the mouths of the canyons.

Descurainia pinnata (Walter) Britton ssp. menziesii (DC.) Detling, Amer. Midl. Naturalist
22:508. 1939.

Sisymbrium canescens of Guadalupe Island references.

RANGE. — Central California to NW Baja California; Santa Rosa, Santa Cruz, Anacapa,
Santa Catalina, San Clemente, Los Coronados, and Todos Santos Islands.

Palmer found this plant in great abundance in warm sheltered localities (Watson 1876:113).
Howell (1942:149) found it occasional in a meadow in the cypress grove. I found a small
colony on the south inner slope of the crater of Outer Islet. It seems to be generally
scarce on the island, if not now extinct.

Detling (1939:509) cited Brandegee’s Guadalupe Island collection as ssp. menziesii but
cited Howell’s as ssp. halictorum (Cockerell) Detling. Raven (1963:322) noted that the
plants from San Clemente Island and some from Guadalupe Island have been considered
ssp. halictorum but that their affinities with ssp. menziesii were more obvious.

Erysimum moranii Rollins, Contr. Gray Herb. 200:193. 1970.

Erysimum insulare of Guadalupe Island references

RANGE. — Endemic to Guadalupe Island; known only on Outer Islet.

This is a shrub to 5dm high with stems to | cm thick, with densely appressed short

silvery pubescence, yellow flowers in March to June and even August, and straight spread-

ing siliques 2—5 cm long.

Moran (1951:154) found this plant occasional on the inner slope of the crater of Outer
Islet. It also grows in the upper valley of Outer Islet but has not been found on the main
island or elsewhere.

102

GUADALUPE ISLAND FLORA

On the basis of flowering specimens with young fruit, Rossbach (1958:122) referred to
the plant of Outer Islet as an atypical form of £. insulare Greene, otherwise of San
Miguel, Santa Rosa, and Anacapa Islands. In naming £. moranii, Rollins said that in its
shrubby habit it was most like £. insulare and E. suffrutescens (Abrams) G. Rossb. among
American species. He thought it might be more closely related to E. swffrutescens, of the
California coast, and particularly to var. grandifolium G. Rossb., of Santa Barbara and
San Luis Obispo Counties.

Guillenia lasiophylla (Hook. & Arn.) Greene, Leafl. Bot. Obs. 1:227. 1906.

Sisymbrium reflexum Nutt., Proc. Acad. Nat. Sci. Philadelphia 4:25. 1850.
Thelypodium lasiophyllum (Hook. & Arn.) Greene, Bull. Torrey Bot. Club 13:142. 1886.
Caulanthus lasiophyllus (Hook. & Arn.) Payson, Ann. Missouri Bot. Gard. 9:303. 1922.

RANGE. — Washington to northern Baja California; islands of southern California except
San Nicolas; also Todos Santos, Cedros, and Natividad.

Palmer found this plant abundant in the “middle” and at the “south end” of the island,
in low grounds (Watson 1876:113). Payson (1923:306) cited Palmer’s collection and one
by Brandegee. Howell (1942:149) found it occasional in the middle and southern parts
of the island. It is now rather scarce in the northern part of the island and widespread in
the southern part, where it may be locally common in good years.

Hutchinsia procumbens (L.) Desy., Jour. Bot. 3:168. 1814.
Capsella procumbens (L.) Fr., Novit. Fl. Suec. Mant. 1:14. 1832.

RANGE. — British Columbia to central Baja California and the Old World; San Miguel,
Santa Rosa, Santa Cruz, Anacapa, San Nicolas, and Santa Barbara Islands.

Howell (1942:149) found this plant in gravelly soil at the south end of the island. I have
found it scarce and in only two places: north outer slope of Outer Islet (4/5655) and on
talus of north-facing seacliff northeast of Campo Sur at 50 m (\//3766).

Lepidium lasiocarpum Nutt. var. latifolium C. L. Hitche., Madrono 8:136. 1945.
L. lasiocarpum of Guadalupe Island references.
RANGE. — Baja California to Arizona and Sonora; Isla Clarion.

Palmer found this plant in ravines in the “middle” of the island and rarely at the “south
end” (Watson 1876:113). It is widespread on the island, from the pine ridge and Northeast
Anchorage to the mouth of Arroyo Melpomene and in good years locally common. From
Guadalupe Island Hitchcock (1945:136—137) cited collections by Palmer, Howell, Bran-
degee, and Mason.

oLepidium nitidum Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:116. 1838.

RANGE. — Washington to NW Baja California; islands of southern California except San
Miguel; also Los Coronados and Todos Santos.

Howell (1942:150) was first to report this plant from the island; he saw it only on the
trail to the cypress grove. It now also grows near the spring and in the shallow valley
below the Lower Circus. It seems remarkable that the earlier collectors did not find it,
and | list it with the west American plants that are possibly late arrivals (Table 2).

103

REID MORAN

Lepidium oblongum Small var. insulare C. L. Hitche., Madrofo 8:125. 1945.

L. bipinnatifidum, L. menziesii, and L. oblongum of Guadalupe Island references.

RANGE. — San Miguel, Santa Cruz, Anacapa, San Nicolas, Santa Catalina, San Clemente,
Todos Santos, Guadalupe, Cedros, San Benito, and Natividad Islands.

Palmer found this plant generally abundant on warm hillsides throughout the island (Wat-
son 1876:113). It is widespread, from the pine ridge to the south end and in good seasons
fairly common.

The type of this variety is Palmer 7 (NY; isotypes GH, MO) from Guadalupe Island.
From Guadalupe Hitchcock also cited collections by Palmer in 1889, Rose, Drent, Greene,
and Brown. He said that most material from the mainland of California and Baja California
has larger silicles than do the insular plants and is otherwise somewhat intermediate with
the widespread typical variety.

*Raphanus sativus L., Sp. Pl. p. 69. 1753.
RANGE. — Native to Europe; a common weed in western North America.

Meling Lopez (1985:50) was first to find this weed on the island, in the corrals in the
cypress wood. It was still there in 1988 (4737006), uncommon with abundant Sisymbrium
orientale.

*Sisymbrium irto L., Sp. Pl. p. 659. 1753.
RANGE. — Native to Europe; a weed in western North America.

Howell (1942:149) was first to find this weed on the island, seeing it in 1932, only near
the barracks. Apparently the only other collection until recently was from the lobster
camp (Weber & McCoy 11) in 1963. In 1988 it was occasional at Northeast Anchorage
(413/001) and with the more common S. orientale in open ground near the cabin at the
northeast edge of the cypress grove (73/007), perhaps from a new introduction.

*Sisymbrium orientale L., Cent. Pl. 2:24. 1756.
RANGE. — Native to the Old World; a common weed in western North America.

This weed is a recent arrival, not reported before. In 1988 it was occasional at Northeast
Anchorage (3/000) and near the spring (4/3/7004) and locally common in open ground
near the cabin at the northeast edge of the cypress (A/3/005).

Thysanocarpus erectus S. Watson, Proc. Amer. Acad. Arts Sci. 11:124. 1876.
RANGE. — Guadalupe and Cedros Islands; north central Baja California.

Palmer found this plant only between Jacks Bay (West Anchorage) and Mount Augusta,
in clear level spots (Watson 1876:113). Howell (1942) found it not common, on rocky
slopes between the cypress grove and the broad transverse valley. It appears to be local
in the central part of the island: occasional on bare rocky areas of red soil at 800—
900m, south of the cypress grove (M6675); scarce in Juniper Canyon at 100m
(M1205912), scarce in Long Canyon at 200 m (M//2076).

104

GUADALUPE ISLAND FLORA

Cucurbitaceae

Marah guadalupensis (S. Watson) Greene, Leafl. Bot. Obs. 2:36. 1910.

Megarrhiza guadalupensis S. Watson, Proc. Amer. Acad. Arts Sci. 11:138. 1876.
Echinocystis guadalupensis (S. Watson) Cogn., Monogr. Phan. 3:819. 1881.
Micrampelis guadalupensis (S. Watson) Greene, Pittonia 2:129. 1890.

RANGE. — Endemic to Guadalupe Island?

Tuber (the one dug) irregular because among rocks, 47 cm high, 12 x 6 cm wide above,
tapering downward. Stems to 6 m long and 7 mm thick; internodes to 20 cm long. Leaf
blades orbicular to ovate in outline, 10-27 cm wide, mostly 5-lobed + to middle, the lobes
triangular-ovate to oblong, acute to obtuse, the sinus 3—6 cm deep, wider to narrower than
deep, the margins sinuate to denticulate or somewhat dentate, the upper surface slightly
papillate-scabrous, minutely pubescent, especially on veins, the lower surface more nearly
glabrous; petiole 4-13 cm long; tendrils bifid or trifid, the peduncle 3—8 cm long. Staminate
flowers 6—20 in raceme, or in panicle 1-3 dm long of up to 6 racemes, on pedicels 3—20
mm long; calyx lobes linear-subulate, 1.5—3 mm long; corolla white, broadly campanulate,
12-30 mm wide, the tube obconic, 3—S mm long, the lobes unequal, triangular-oblong,
rounded to subacute, 6-14 mm long, 3—7 mm wide; anther head subglobose, 1.5—2.5 [?]
mm wide. Pistillate flowers on pedicels to 2cm long; calyx lobes linear, 2 (—7?) mm
long; corolla white, open-campanulate, 18-40 mm wide, the tube 2-4 mm long, the lobes
oblong-lanceolate, acute to obtuse, 8-14 mm long, 2—5 mm wide. Fruit ovoid, short beaked,
to 6cm long and 5 cm thick, striated from base to apex, short pubescent, rather sparsely
spinose, the spines 1-15 mm long, to 1-2 mm wide at base, glabrous or puberulent; pedun-
cle 5-8 cm long. Seeds 24, ovoid lenticular, 28 mm long, 25 mm wide, 14 mm thick
(immature), olivaceous, the circumferential line inconspicuous. Flowers December to May.
(Description broadened from Stocking 1955.)

Palmer found this plant in crevices of high rocks in the “middle” of the island (Watson
1876:138). Franceschi (1893c:135) saw it only among rocks on the right bank of the
canyon not far from the landing, but he was assured that it grew all over the island. He
noted that young shoots appeared about the middle of January. It is now uncommon, in
the north on cliffs but in the south sometimes growing well within reach of the goats yet
showing no damage (Fig. 60): Barracks Canyon, 475 m; Esparsa Canyon, occasional on
cliffs and banks, 50 m (M5974), on talus to 250 m; narrow talus above beach, south of
Esparsa Canyon (Wiggins & Ernst 56); Juniper Canyon, 50 m (M6455); Long Canyon,
200 m (M12078); canyon behind lobster camp, 50 m; Arroyo Melpomene, 400 m (M6734).

Stocking (1955) kept MM. guadalupensis as an endemic of Guadalupe Island, characterized
by large campanulate corollas, linear calyx teeth, in pistillate flowers “6-7 mm long”, and
weak-spined two-seeded fruit. Besides the type collection (Pa/mer 33, GH, US), he cited
collections by Anthony, Brandegee, and Franceschi. The island plants of southern Cali-
fornia he called M. macrocarpus (Greene) Greene var. major (Dunn) Stocking (= M. major
Dunn), differing from typical mainland MM. macrocarpus in their larger leaves, flowers,
and seeds. Wallace (1985:15) also kept M. guadalupensis as a Guadalupe endemic, placing
other island plants in M. macrocarpus. Schlising (1993) and Junak et al. (1995) treated
some Channel Island plants as M. macrocarpus var. major, thus by implication also rec-
ognizing M. guadalupensis. On the other hand, Ferris (1960:70) broadened M. guadalu-
pensis to include much of M. major but kept some other island plants in MM. macrocarpus;
and Wiggins (1980:393) did the same. Raven (1963:340) treated the San Clemente Island

105

REID MORAN

FIGURE 59. Plant of the endemic Dudleyva virens ssp. extima (M6436) from a cliff high at the
north end, flowering in San Diego, 11 May 1960

FIGURE 60, The endemic Marah guadalupensis (M6455) at the mouth of Juniper Canyon, 50 m,
16 December 1957. It is accessible to goats but 1s bitter and not eaten

106

GUADALUPE ISLAND FLORA

plants under M. macrocarpus but remarked that much more fruiting material was needed
from the islands before definite conclusions could be reached. He thought that although
the extremes were very different, it did not seem useful at that time to recognize the
variety on the islands, and he thought the plants from Guadalupe Island also very probably
conspecific with M. macrocarpus. More fruiting material still is needed.

Stocking (1955) thought the wide-ranging 7. oreganus (Torr. & A. Gray) Howell, with
its comparatively large calyx lobes and sparsely-spined fruit, was primitive in the genus;
and he thought /. guadalupensis much like it in flowers and fruit and perhaps its closest
relative. Although I have found no mature fruit and seeds of M. guadalupensis, immature
fruits on five collections from different places on the island are fairly uniform in size,
with spines not so sparse as in M. oreganus, often as short as 2 mm but sometimes to 15
mm. In two plants growing side by side (M/2078, 12078A), spines were short in one,
long in the other. Greene (1885:223) wrote that the fruit was conspicuously flattened
laterally, but this was not true in those I saw; possibly he meant the seeds.

In its large leaves and large flowers the Guadalupe plant differs from mainland M. macro-
carpus and agrees with the later-named M. major. Calyx lobes are longer than decribed
for other species, though | did not find them so long in the pistillate flowers as both
Watson and Stocking described them. There is still a question whether this is distinct
from M. major and so whether M. guadalupensis is confined to Guadalupe Island. Tim
Ross, who has on loan the three cited cotypes of M. major, writes me that Trask 9/ is
from San Nicolas Island and 7rask 280 and 28/ from San Clemente Island. It is to be
hoped that future collectors will get mature fruit and seeds.

Ericaceae

+Arctostaphylos sp.
RANGE. — Unknown; perhaps endemic to Guadalupe Island.

Greene (1885:225) found a single seedling of not more than two or three years’ growth,
under a cypress; he called the species apparently new. There are no further reports, and
the plant is undoubtedly extinct. Greene’s specimen has not been found.

Euphorbiaceae

Note. — For Chamaesyce melanadenia and Stillingia linearifolia see Excluded Species,
p. 168.

Chamaesyce pondii (Millsp.) Millsp., Field Mus. Nat. Hist. Bot. Ser. 2:411. 1916.

Euphorbia pondii Millsp. in Vasey & Rose, Contr. U.S. Natl. Herb. 1:12. 1890.
Euphorbia guadalupensis J.T. Howell, Leafl. W. Bot. 1:51. 1933.

RANGE. — West central Baja California and Guadalupe Island.

On his second trip Palmer collected this plant (883, F, GH) at the south end of the island.
Although Vasey and Rose (1890:21—27) failed to list it with his Guadalupe Island col-
lections, Millspaugh mentioned Palmer’s Guadalupe specimen in describing the species
on p. 12 of this same paper. Howell (1933, 1942:151) found the plant occasional on
gravelly flats or in coarse sandy soil on dry windswept slopes at the south end. It appears
to be local near the west edge of south end mesa, at 100-200 m, just above Campo Sur.

107

REID MORAN

Howell (1933) described Euphorbia guadalupensis as a Guadalupe Island endemic, based
on his collections 833/ (type CAS) and 8/95, from Melpomene Cove. However, Wheeler
(1934) pointed out that it was the same as E. pondii. Besides the Guadalupe specimens,
Wheeler (1936:12) cited for E. pondii only the type collection, from Playa Maria. He
said E. pondii was close to E. polycarpa Benth. var. polycarpa but was a small annual.

Euphorbia misera Benth., Bot. Voy. Sulph. p. 51. 1854.

RANGE. — Southern California to Sonora and central Baja California; Santa Cruz, Santa
Catalina, San Clemente, Los Coronados, Todos Santos, San Martin, San Benito, and Cedros
Islands.

Drent in 1898 was first to collect this plant on Guadalupe Island, though his find went
unreported. Moran (1951:157) found it common on the floor and occasional on the inner
slope of the crater of Outer Islet. It is rather rare on cliffs in the northeast part of the
island and scarcely more common on the mesa at the south end: two in Esparsa Canyon,
300 m (6/30), one at 230 m (M18/52); lobster camp, 70 m (M6449); Arroyo Melpomene,
350 m (M12072), 170 m (M13767); south end mesa, 100 m (1/3767).

Catalina, San Clemente, Los Coronados, Todos Santos, San Martin, San Benito, and Cedros
Islands.

Drent in 1898 was first to collect this plant on Guadalupe Island, though his find went
unreported. Moran (1951:157) found it common on the floor and occasional on the inner
slope of the crater of Outer Islet. It is rather rare on cliffs in the northeast part of the
island and scarcely more common on the mesa at the south end: two in Esparsa Canyon,
300 m (A16/30), one at 230 m (M78152); lobster camp, 70 m (M6449); Arroyo Melpomene,
350 m (M12072), 170 m (M13767); south end mesa, 100 m ((//3767).

Fagaceae

Quercus tomentella Engelm., Trans. Acad. Sci. St. Louis 3:393. 1877.
Quercus chrysolepis of Guadalupe Island references.

RANGE. — Endemic to Santa Rosa, Santa Cruz, Anacapa, Santa Catalina, San Clemente,
and Guadalupe Islands.

Palmer reported the oak as frequent at the north end and occasional in canyons on both
sides of the island (Watson 1876:119). Greene (1889:45; 1890a:57) saw a few trees at
the “cool foggy summit” of the north end and a few more, somewhat different in appear-
ance, “in entirely different soil, and on the very opposite kind of exposure, namely in
certain dry, heated canyons of the southeastward slope.” “It seems to exist on Guadalupe
only in a few more than middle-sized trees, at the cool, foggy summit of the north end
of the island; and in a scarcely more considerable number of smaller, and smaller-leaved
trees (possibly not of the same species) in the hot and dry canyons of the eastward slope.”
Franceschi (1893c:138) mentioned oaks in the northwestern part of the island with the
palms, as well as a few scattered trees at the north end; he reported oaks in two places
more than a mile apart on the east side under the cliffs of the Lower Circus; and he said
that the oak was the only tree to be seen in the southern part of the island. And Howell
(1942:147) told of a few trees near the head of Barracks Canyon (Figs. 61, 62).

Watson listed Palmer’s two numbers (88, 89) as “Q. chrysolepis Liebm.; fide Dr. Engel-
mann,” and Engelmann (1877) referred both to Q. tomentella when he founded the species.

108

GUADALUPE ISLAND FLORA

FIGURES 61 and 62. The island oak, Quercus tomentella, endemic to Guadalupe and five of the
Channel Islands. FIGURE 61. Two trees at the mouth of Oak Canyon, a tributary of Barracks Canyon
(M6443), 14 December 1957; these trees are now gone. FIGURE 62. Tree on north slope of the
island, 28 March 1978.

109

REID MORAN

Greene, Franceschi, and Howell, however, all pointed out differences between the oaks
at the north end and those of the east side.

Franceschi described the trees of the northwest slope as “fine specimens forty to sixty
feet high, remarkable for the grayish color of the bark and the size of the leaves, which
are glossy dark green on the upper surface and covered with a somewhat rusty tomentum
beneath.” Howell described specimens from the grove illustrated by Eastwood (1929,
plate 34, figs. 1, 2): “The twigs and the lower side of the leaves of the current season are
densely tomentellous with rather loose buff-colored hairs. The older branches are glabrate
or nearly so and the lower side of older leaves are only thinly pubescent and slightly
glaucous.”

Franceschi wrote that the oaks of the east side, “if not specifically distinct appear at least
to be a very different form, not only by the leaves but also by the bark, which is darker
and corky. These trees are rather stunted and branching from the base.” Of specimens
from these trees, Howell wrote: “Although these specimens are not entirely typical of Q.
chrysolepis Liebm., they are more like that species than [like? or are?] the specimens
from the north end. The leaves are more coriaceous; the upper side 1s light green and the
lower side is entirely pale and chalky with a scattering of those peculiar glandular trichomes
characteristic of the canyon live oak of California. The pubescence on the twigs and
petiole is thick and close, quite unlike the relatively shaggy pubescence of [the oak from
the north end]. I was able to obtain acorns and cups, and these resemble the acorns and
cups of that form of Q. chrysolepis that occurs in the mountains of southern California
that has been called Q. crassipocula Torr.”

Greene considered the trees at the north end as typical QO. tomentella. He thought the
trees of Santa Cruz Island corresponded more closely to those on the east side of Guadalupe
Island than to those at the north end.

Dr. C. H. Muller collected the oak at the north end in 1957 and studied my collections
(\16439-M6443) from Oak Canyon on the east side. He wrote me that the trees at the
north end agree more closely with those on the islands of California and are biologically
more typical of the species, whereas those of Oak Canyon show strong introgression by
Q. chrysolepis but agree more closely with the type specimen.

The main stand of oaks apparently was with the pines at about 600 to 700 m on the steep
northwest slope of the main northeast ridge (Fig. 62). Other oaks are scattered along the
northwest slope of this ridge up to perhaps 900 m, and a few more are scattered among
the palms of the northwest slope west of the main stand. The total number of trees at my
first visit was probably less than 100, and they have been falling with the years.

Of the oaks formerly in the canyons of the east side, two small groups remained at the
time of my visits. One must be the more northern of the two groups mentioned by
Franceschi and is where Mason (/537) and Howell (8/73) collected. These trees were at
ca. 850-900 m in the north fork of Barracks Canyon, in the steep gorge through the cliffs
of the Lower Circus or rim of the lower plateau which I call Oak Canyon (Fig. 61). In
1957 there were nine living trees, or perhaps seven, two with trunks double from the
base; they were 8-10 m tall, with trunks to 6 dm thick and bark in thin grayish plates.
Two of these trees, undercut by the torrent that flows after rains, had fallen; and by 1960
one of these was gone. By 1981 only four trees remained and by 1988 only one. Jose
Rico, just back from the island in July 1994, told me that too was gone.

In 1967 there were five oaks (41/3788, M13789) at 450 m in a hanging tributary on the

110

GUADALUPE ISLAND FLORA

north side of Esparsa Canyon about 3 km east of Mt. Augusta, which I call Southeast
Oak Canyon (Fig. 26). The trees were about 8 m tall, with widespreading branches, in
some places decumbent on the rocks. José Rico told me in 1994 that only two remained.

In 1988 a single spreading tree stood at 1100 m at the head of Esparsa Canyon on the
east slope just south of Mt. Augusta (173/008).

In 1970 there were two oaks at 700 m on the steep west slope below the gray peak above
West Anchorage (1/7394). They were erect, about 8 m tall, with trunks to 5 dm thick.

Minnich (1987:124) wrote that Q. tomentella occurs south of the U.S. border only in
central Baja California “in an arroyo 3 km east of Mt. Augusta and on Guadalupe Island.”
So far as known, however, neither the oak nor the mountain occurs on the peninsula.

Frankeniaceae

Frankenia salina (Molina) |. M. Johnst., Contr. Gray Herb. 70:92. 1924.

Frankenia grandifolia Cham. & Schltdl., Linnaea 1:35. 1826.

RANGE. — Central California to Nevada, central Baja California, and South America; is-
lands of southern California except Santa Barbara, and also Todos Santos.

Dudley (1899:281) reported this plant on the side of the bank near Northeast Anchorage,
where no one else has found it before or since. The only other locality known on the
island is at 5m on the low barren lava flat, near the upper limit of salt spray, at the
southwest corner of the island, northwest of Campo Sur (46472, M/3756).

Geraniaceae

*Erodium brachycarpum (Godr.) Thell., Rep. Bot. Exch. Club Brit. Isles 5:17. 1918.
RANGE. — Native to North Africa; naturalized in western North America.

This weed was getting well started in 1978: occasional in shallow grassy valley below
Lower Circus, 800 m (25374); occasional east of the principal spring, 1000 m (4/253 76).
By 1988 it was abundant about the spring (/3/0//) and elsewhere on the upland.

*Erodium cicutarium (L.) L’Her. ex Aiton, Hort. Kew. 2:414. 1789.
RANGE. — Native to southern Europe; widely naturalized in western North America.

Palmer found this weed abundant all over the island, and the principal food for the goats,
covering the rocks; usually three or four inches high by August (Watson 1876:113). Greene
(1885:222) saw very little of either species, but Franceschi (1893c¢:134) said it literally
covered the whole surface of the island. It now extends the length of the island and is
one of the very commonest plants on rocky slopes in the northern half.

*Erodium moschatum (Burm.f.) L’Her. ex Aiton, Hort. Kew. 2:414. 1789.
RANGE. — Native to southern Europe; widely naturalized in western North America.

Palmer found this weed in the “middle” of the island, less common than the last and
starting later in the spring (Watson 1876:114). Franceschi (1893c:134) found it plentiful
all over, chiefly among rocks and stones, but not so common as E. cicutarium. Howell

111

REID MORAN

(1942:150) found it throughout the island, especially at lower elevations, and said the two
species are perhaps the commonest plants after Hordeum; and that is still true.

Grossulariaceae

+Ribes sanguineum Pursh, Fl. Amer. Sept. p. 164. 1814.
RANGE. — British Columbia to NW California; Guadalupe Island.

Palmer found only two plants of this species, in the damp shade of cliffs at the north end
(Watson 1876:114); and no one else has seen it. Quite possibly it is extinct on the island,
and I have marked the name with a dagger. Nevertheless, | am reluctant to declare the
case closed, for the foggy northwest cliff has many inaccessible damp shady ledges, on
some of which it might possibly survive.

Eastwood (1929:407) thought this must be some other species because this one is found
in California only at the extreme north. However, Peter Raven wrote me in 1962 that

Otto Solbrig had checked Palmer’s specimen and found it to be R. sanguineum — with
glabrous styles and glandular fruit.

Hydrophyllaceae

Emmenanthe penduliflora Benth., Trans. Linn. Soc. 17:281. 1835.

RANGE. — Central California to Utah, Arizona, and central Baja California; Santa Cruz,
Santa Catalina, and San Clemente Islands.

Palmer found this plant in rocky ravines in the “middle” of the island (Watson1876:118)
and on his second trip saw it in canyons and on hillsides at both ends of the island (Vasey
and Rose 1890:26). Greene (1885:226) called it abundant, very large and handsome, far
surpassing what one sees of this species in California. It is occasional or locally common
in arroyos and on rocky slopes in the northeast part of the island and south as far as the
lobster camp.

Eucrypta chrysanthemifolia (Benth.) Greene var. chrysanthemifolia
Eucrypta chrysanthemifolia Greene, Bull. Calif. Acad. Sci. 1:200. 1885.

RANGE. — California to NW Baja California; San Miguel, Santa Rosa, Santa Cruz, Ana-
capa, Santa Barbara, Santa Catalina, and San Clemente Islands.

Palmer collected this plant apparently with the following (Watson 1876:118). It is occa-
sional in the northeast part of the island, favoring more shaded places than the following.
Constance (1938:146) cited the collections of Palmer and Mason.

Constance recognized both the species and the variety, with generally different but some-
what overlapping ranges.

Eucrypta chrysanthemifolia (Benth.) Greene var. bipinnatifida (Torr.) Constance, Lloydia
1:147. 1938.

Ellisia chrysanthemifolia of Guadalupe Island references.

RANGE. — SE California to Nevada, Arizona, and central Baja California.

GUADALUPE ISLAND FLORA

Palmer found this plant abundant under sage brush on warm hillsides from the “middle”
to the north end and also rarely at the “south end” (Watson 1876:118). It is occasional
or locally common in canyon bottoms and among rocks in the northeast part of the island.

From Guadalupe Island Constance (1938:148) cited Palmer 74 and Anthony 252. Many
collections he found impossible to assign with any degree of certainty to either the species
or the variety, and he remarked that the rather abundant collections from Guadalupe Island
illustrate admirably the confusing transitions that occur whenever the two occur together.

Phacelia floribunda Greene, Bull. Calif. Acad. Sci. 1:200. 1885.

Phacelia phyllomanica A. Gray var. interrupta A. Gray, Proc. Amer. Acad. Arts Sci. 11:87. 1876.
RANGE. — Endemic to Santa Barbara, San Clemente, and Guadalupe Islands.

Rather dense annual to 6 dm high and wide; flowers lavender, February to May.

Palmer found this plant frequent in warm nooks in rocky ravines in the “middle” and at
the “south end” of the island (Watson 1876:118). Greene (1885:226) found it in the lower
parts of the island. It is occasional in arroyos in the southern half of the island, from the
lobster camp and West Anchorage south and on Islote Negro and Outer Islet, and less
common now in the northern part.

Gray named the variety from Palmer’s Guadalupe Island collection (No. 72, GH holotype).
Greene thought it in no wise resembled the gigantic half shrubby P. phyllomanica, of the
upper precipices, except as regards the pinnately parted calyx lobes. In M5653, from
Outer Islet, Marion Cave found n= 11 (Cave and Constance 1959).

Phacelia phyllomanica A. Gray, Proc. Amer. Acad. Arts Sci. 11:87. 1876.
RANGE. — Endemic to Guadalupe Island.

Leafy subshrub 1-2 m high and to 2m wide, canescent with dense short soft hairs and
above longer villous, not hispid or glandular but somewhat viscid above. Leaves 6-10
cm long, 34.5 cm wide, pinnatisect, with 13-25 crowded oblong-linear pinnatifid seg-
ments, the lobes short, oblong, sometimes few-toothed. Cincinni thyrsoid-congested, to 5
cm long, with 40-55 flowers, at first dense, the flowers sessile. Calyx segments either
all leaflike and 3—5-pinnately parted or 1-3 broadly linear and entire. Corolla blue-violet,
slightly exceeding calyx, the scales short and broad. Annular disk conspicuous, the stamens
subexserted. Chromosomes: n= 11. (Fig. 63)

Palmer found this plant rare, in large compact masses in the crevices of high rocks in
the “middle” of the island (Watson 1876:118). Gray named the species from Palmer’s
collection (No. 72, GH type). Greene (1885:226) said it was shrubby below and often
more than six feet high, the largest species known. Franceschi (1893c:136) saw a con-
siderable number in a limited locality on the cliff east of the corral; he called it a most
elegant shrub, with finely cut foliage, dark green above and whitish below. In January
1960 and May 1967 I found a group of about eight shrubs, not flowering, high overhead
on the cliffs of the Lower Circus (M7836, M/3782). One was + 1.5m high and 2m
wide. In 1981 I saw 6 or 8 large clumps out of reach on ledges of the western cliffs at
1075 m, northwest of the cypress grove. Seeds from 1960 grew at the U. C. Botanical
Garden, flowering September 1962 (Fig. 63), and Marion Cave got a count of n= 11
(apparently unpublished).

REID MORAN

FIGURE 63. The endemic shrubby Phacelia phyllomanica (M7836), grown in Berkeley from seed
from the Lower Circus, 3 September 1962

Pholistoma racemosum (Nutt.) Constance, Bull. Torrey Bot. Club 66:349. 1939.

Nemophila racemosa Nutt. ex A. Gray, Proc. Amer. Acad. Arts Sci. 10:315. 1875.
Vemophila aurita of Guadalupe Island references

RANGE Southern California to central Baja California and Isla Cedros; Santa Cruz,
Santa Barbara, Santa Catalina, San Clemente, Los Coronados, Todos Santos, San Martin,
Cedros, and Natividad Islands.

Palmer found this plant on warm slopes in the “middle” of the island and rarely at the
“south end” (Watson 1876:118). Franceschi (1893c:136) found it plentiful among rocks
all over the island. It is widespread in the northern half of the island, from the north slope
to Juniper Canyon, locally common among rocks, and less common southward but ex-
tending to Outer Islet. From Guadalupe Island Constance (1939:350) cited collections of
Anthony, Franceschi, Greene, Palmer, and Rose.

Labiatae
+Pogogyne tenuiflora A. Gray, Proc. Amer. Acad. Arts Sci. 11:100. 1876.
Hedeomoides tenuiflora (A. Gray) Brig. in Engl. & Prantl, Nat. Pflanzenfam. IV, 3a:295. 1896

RANGE. — Endemic to Guadalupe Island.

Palmer found this plant very rare among sage brush on the eastern side (Watson 1876:117).
No one has found it since, and undoubtedly it is extinct.

114

GUADALUPE ISLAND FLORA

Gray based P. tenuiflora on Palmer’s collection (No. 65, GH holotype) from Guadalupe
Island. This species falls in subgenus Hedeomoides A. Gray, which was made a separate
genus by Briquet (1896), with the upper pair of stamens sterile. Howell (1931) thought
it probably closest to the widespread P. serpylloides, mainly of central California and
now very rare if not extinct in northwest Baja California near San Quintin.

+Satureja palmeri (A. Gray) Briq. in Engl. & Prantl, Nat. Pflanzenfam. IV, 3a:303.
1896.

Calamintha palmeri, A. Gray, Proc. Amer. Acad. Arts Sci. 11:100. 1876.
Clinopodium palmeri (A. Gray) Kuntze, Rev. Gen, 2:515, 1891.

RANGE. — Endemic to Guadalupe Island.

Palmer found this plant abundant among trees and sage brush in the “middle” of the
island (Watson 1876:117). He said it was strong-scented and not eaten by the goats, but
he didn’t know how hungry they would be. Greene (1885:117) listed it without comment.
No one has found it since, and undoubtedly it is extinct.

Gray based the species on Palmer’s collection (No. 66, GH holotype?). He placed this
species in section Acinos and added the following remarks: “Calyx shorter than in C.
acinos, in fruit less declined or ascending. Except for the four fertile stamens this plant
would be referred to Hedeoma. The stamens are too straight and distant for a Calamintha,
but apparently it may be referred to that polymorphous genus.”

Leguminosae

Lotus argophyllus (A. Gray) Greene ssp. ornithopus (Greene) Raven, Aliso 5:326.
1963.

Hosackia ornithopus Greene, Bull. Calif. Acad. Sci. 1:185. 1885.

Syrmatium ornithopum (Greene) Greene, Bull. Calif. Acad. 2:148. 1886.

Lotus argophyllus var. ornithopus (Greene) Ottley, Univ. Calif. Publ. Bot. 10:238. 1923.
Hosackia argophylla of Guadalupe Island references.

RANGE. — Endemic to San Nicolas, Santa Barbara, Santa Catalina, San Clemente, and
Guadalupe Islands.

Palmer found this plant in the crevice of a rock (Watson 1876:114). Greene (1885:185,
216) called it frequent in the middle of the island and mentioned it as one of four con-
spicuous plants of middle altitudes where the trail climbed out of the canyon above the
landing. Moran (1951:157) found it on the inner slope of the crater on Outer Islet. In the
northern part of the island it is occasional on cliffs, with seedlings often seen on the
slopes below. It is fairly common at West Anchorage and southward, often in arroyos.
The type of Hosackia ornithopus is Greene’s collection of April 19, 1885, from Guadalupe
Island (UC 80750, part of type). From var. ornithopus Isely (1981) separated the plants
of San Nicolas, Santa Barbara, San Clemente, and Santa Catalina Islands as var. argenteus
Dunkle, thus leaving ssp. ornithopus only on Guadalupe Island. He wrote that Guadalupe
plants had clusters of greatly exserted pods, some with 3 seeds, whereas plants from the
northern islands have shorter pods, with only 2 seeds, as in other forms of L. argophyllus.

Lotus grandiflorus (Benth.) Greene, Pittonia 2:145. 1890.

Hosackia grandiflora Benth., Trans. Linn, Soc. 17:366. 1837.

115

REID MORAN

Lotus guadalupensis Greene, Pittonia 2:144. 1890.
Lotus grandiflorus (Benth.) Greene var. mutabilis Ottley, Univ. Calif. Publ. Bot. 10:208. 1923.

RANGE. — Northern California to NW Baja California; Santa Rosa, Santa Cruz, and Santa
Catalina Islands.

Palmer found this plant among trees in the “middle” of the island (Watson 1876:114).
Greene (1885:222) saw no trace of it at Palmer’s locality and found only a single large
bushy plant in a nearly inaccessible crevice on the face of a cliff. No one saw it again
until April 1963, when Weber and McCoy (//969) and Copp (/46) found it rather common,
flowering in the arroyo bed at Northeast Anchorage. It was still there in February 1965
(M120/2), but I failed to find it in 1967 and later. It seems unlikely to have been growing
at Northeast Anchorage, the usual landing, all those years without having been found.
Presumably it survived on some cliff above, got started in the arroyo in a year when
goats were few, then disappeared when goats increased again. Meling L. 014, of March
1982, is labeled “Extremo Sur,” the extreme south end, which seems very improbable.
Meling L. (1985:54) reported finding four plants near the “aguaje del bosque” and one
in an arroyo near the south end.

Greene (1885) at first reported his collection as Hosackia grandiflora. He later (1890b)
described Lotus guadalupensis as follows: “Branches stout, erect, 2 feet high, from a
suffrutescent base, and with the other parts minutely velvety: internodes short, not equalling
the sessile leaves: leaflets about 5 pairs, obovate-oblong, mucronulate: peduncles stout,
surpassing the leaves, the 6—10-flowered umbel with a large |-foliate bract: calyx-teeth
subulate-setaceous, rather shorter than the oblong-campanulate tube: corolla 6 to 7 lines
long, yellow. . . . It 1s singular among the species of this group in having numerous
leaflets equally distributed on the rachis; so that only on account of its gland-like stipules
is It retained in this place. But it is very closely related to [L. leucophaeus Greene] though
more remote from the true L. grandiflorus.”

Ottley (1923:208) and Wiggins (1980:680) included L. guadalupensis in L. grandiflorus.

bLupinus bicolor Lindl., Bot. Reg. 13: pl. 1109. 1827.

Lupinus bicolor Lindl. var. microphyllus (S. Watson) C. P. Sm., Bull. Torrey Bot. Club 50:382.
1924.
Lupinus bicolor Lindl. ssp. microphyllus (S. Watson) D. B. Dunn, Aliso 3:151. 1955.

RANGE. — British Columbia to Sonora and central Baja California; islands of southern
California except Santa Barbara.

The first collections of this plant from Guadalupe Island are by Drent in 1898 and Brown
in 1906, both without specific localities (Smith 1923:385). It grows in grassy places in
the northeast part of the island, from Northeast Anchorage and the spring to El Picacho,
generally rather scarce but in a few places locally common in good years. Since earlier
botanists failed to find it even though they collected in the general area where it now
grows, I list it as a possible late arrival (Table 2).

This plant has been reported before as L. bicolor var. or ssp. microphyllus, but Riggins
and Sholars (1993:628) found the named varieties and subspecies + indistinct in geography
and morphology and in need of further study.

116

GUADALUPE ISLAND FLORA

Lupinus guadalupensis Greene, Bull. Calif. Acad. Sci. 1:184. 1885. (Not L. guadalu-
pensis C. P. Sm. 1951.)

RANGE. — Endemic to San Clemente and Guadalupe Islands.

Greene (1885:184) found this plant on the high plateau of the island, in good fruit but
nearly past flowering April 23. It now appears to be rather scarce and local on the east
side of the island just below the middle, at 100-650 m, from the mouth of Long Canyon
to the slopes of El Picacho and the slope south of Mt. Augusta. Besides Greene’s col-
lections, Dunn (1957) cited one by Brandegee.

On San Clemente Island this plant has been named L. clementinus C. P. Sm., L. alicle-
mentinus C. P. Sm., and L. moranii Dunkle. Dunn (1955) considered it an insular relict
connecting L. nanus and L. affinis with certain Mexican species.

Lupinus niveus S. Watson, Proc. Amer. Acad. Arts Sci. 11:126. 1876.
RANGE. — Endemic to Guadalupe Island.

This is a densely white tomentose bush to 4 dm high and 1 m wide, mostly living only
one season, with light blue flowers. (Fig. 64)

Palmer found this plant only in the “middle” of the island, on high cliffs, only one plant
in bloom, March 25 (Watson1876:114). Greene (1885:222) also saw only one flowering
plant and that almost inaccessible; but numerous seedlings of this, or else of an annual
species with the same pubescence, were growing on level ground south and west of the
cabins, where the goats no longer ranged. On his second trip Palmer found it in flower
and fruit March 29 in the sandy bed of a canyon at the south end (Vasey and Rose
1890:23). In the northern part of the island it is rarely found flowering on cliffs, but the
seedlings are occasional, widespread on the slopes and down to the beach. At the south
end in a good year it is abundant and beautiful in sandy arroyo beds (Plate IF).

Smith (1922:205) called this a very distinct isolated insular species, not closely related
to any other known species but better lined up with L. nanus, at least for the time. Dunn
(1960) grew L. niveus in the greenhouse and tried crosses with two subspecies of L.
nanus. Although he got no seed, some pods started to develop, suggesting to him that
the two species had some affinity and that the placement of L. niveus in the Micranthi
was correct. Harmon and Dunn (1968) found in the greenhouse that L. niveus was a
short-lived perennial. (At the south end of the island | noted a few plants that clearly
were living over a second year.) They also crossed it with several shrubby Californian
perennials and a South American annual. They concluded that L. niveus is a relictual
species transitional between these shrubby lupines and the annual Micranthi.

*Medicago polymorpha L., Sp. Pl. p. 779. 1753.
Medicago hispida Gaertner, De Fruct. Sem. PI. 2:349. 1791.
RANGE. — Native to southern Europe; a widespread weed in western North America.

Howell (1942:150) was first to report this weed on the island; he saw it only near the
barracks. I also have seen it only in that vicinity. The var. brevispina (Benth.) Heyn. is
also there.

117

REID MORAN

FIGURE 64. The endemic Lupinus niveus grown in San Diego, 29 May 1981

*Melilotus indica (L.) All., Fl. Ped. 1:308. 1785.
RANGI Native to Eurasia; a widespread weed in North America

Palmer on his second trip found this plant common along the beach at the north end and
ascending into shady canyons (Vasey and Rose 1890:24). I have seen it only at Northeast
Anchorage and in the mouth of Esparsa Canyon, in neither place common

Trifolium depauperatum Desy. var. truncatum (Greene) Martin ex Isley, Brittonia

32:56. 1980

Trifolium minutiflorum Greene, Pittonia 3:215. 1897
Trifolium amplectens of Guadalupe Island references

RANGI Central California to NW Baja California; islands of southern California except
Santa Barbara.

GUADALUPE ISLAND FLORA

Palmer found this plant rare, only on the beach on the east side of the island (Watson
1876:114). Franceschi (1893c:135) saw it only in the canyon near the landing, but not in
large numbers. Howell (1942:150) found it common at higher elevations. It is fairly com-
mon on the north end and about the spring.

Greene in 1897 described 7. minutiflorum as follows: “Annual, very slender, glabrous,
the numerous branches decumbent, a few inches to almost a foot long: leaflets I/2 to 3/4
inch long, the lowest exactly linear, truncate or emarginate, the others oblong-linear and
abruptly acutish, all somewhat serrately and very evenly denticulate: peduncles filiform,
much longer than the leaves: heads not much more than 2 or 3 lines broad in flower;
involucre parted into 6 or 7 oblong entire lobes: calyx teeth subulate-lanceolate, herbaceous
almost throughout: corollas moderately inflated in age.”

“An interesting species, allied to 7. truncatum and T. hydrophilum, which latter it agrees
with in having the leaflets of its earliest leaves narrower than those of the later ones, in
which respect these two differ from all other West American clovers. This one is of
southern California and Guadalupe Island.”

bTrifolium gracilentum Torr. & A. Gray, Fl. N. Amer. 1:316. 1838.

RANGE. — British Columbia to NW Baja California; islands of southern California and
Todos Santos.

Howell (1942:150) was first to find this plant (8247), calling it the common clover at
lower elevations at the north end of the island, as on the slope of the pine ridge. It is
common on North Point and the uplands about the spring, extending to the beach and to
the south end mesa. Because it is so widespread and common now but was not found
until 1932, I list it with those west American plants that probably are introduced on the
island (Table 2).

Trifolium microcephalum Pursh, Fl. Amer. Sept. 2:478. 1814.

RANGE. — British Columbia and Montana to NW Baya California; San Miguel, Santa
Rosa, Santa Cruz, San Nicolas, Santa Catalina, and San Clemente Islands.

Palmer found this clover very abundant at the “middle” and north end of the island
(Watson 1876:114). It is widespread, from the beach to Mt. Augusta and from North
Point to the northwest slope to El Picacho, often rather common, and then less common
southward, as in the bed of Arroyo Melpomene.

Trifolium palmeri S. Watson, Proc. Amer. Acad. Arts Sci. 11:132. 1876.

Trifolium gracilentum Torr. & A. Gray var. palmeri (S. Watson) McDermott, N. Amer. Trifol. p.
300. 1910.

RANGE. — Endemic to San Nicolas, Santa Barbara, Santa Catalina, San Clemente, and
Guadalupe Islands.

Palmer found this clover rather abundant in the “middle” of the island among rocks and
trees on hillsides (Watson 1876:114). On his second trip, in a year of good rainfall, he
called it very common in large masses in canyons and plains at the north end and quite
common on the south end in wet sand in canyons; he added that it was widely distributed
over the island and formed the main food supply of the goats (Vasey and Rose 1890).
In the northeast part of the island it grows mainly in the mouths of the canyons, but from

119

REID MORAN

West Anchorage and El Picacho southward it is more widespread and common. It is also
on Islote Negro.

Zohary and Heller (1984) followed McDermott in treating this as a variety of 7. gracilen-
tum; and it is closely related, differing chiefly in leaf shape. However, Raven (1963:328)
considered that on San Clemente Island the two were completely distinct both ecologically
and morphologically. And they likewise appear distinct on Guadalupe, in South Esparsa
Canyon in 1958 growing side by side (M6598, M6599). Steve Junak tells me he has found
them distinct also on Santa Barbara and San Nicolas Islands.

Vicia hassei S. Watson, Proc. Amer. Acad. Arts Sci. 25:129. 1890.
Vicia exigua var. hassei (S. Watson) Jeps., Fl. W. Mid. Calif. p. 296. 1901.

RANGE. — Southern Oregon to NW Baya California; Santa Rosa, Santa Cruz, Anacapa,
San Nicolas, Santa Catalina, San Clemente, Los Coronados, and Todos Santos Islands.

Palmer in a wet year called this plant common on shady sides of ravines at the north end
(No. 847, Vasey and Rose 1890:24). Ordinarily, it is rather scarce on the island, as on
the slope above Northeast Anchorage and on El Picacho. Commonly this has been called
a variety of 1. exigua, now included in IV’. /udoviciana. However, Lassetter (1975) and
Gunn (1979) have treated it as a monomorphic species of more limited range, similar to
the polymorphic 1 /udoviciana but distinguished by the apical abaxial stylar brush and
the pubescent ovaries and pods. Gunn cited Guadalupe collections by Palmer (847), Wig-
gins and Ernst (200), and Moran (6643).

Vicia ludoviciana Nutt. ssp. ludoviciana

Vicia ludoviciana Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:271. 1838.
Vicia exigua Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:272. 1838.

RANGE. — To SE US and N Mexico: the exigua phase Northern California to NW Baja
California; Santa Rosa, Santa Cruz, Anacapa, Santa Catalina, and San Clemente Islands.

This species seems to be slightly more common than the other, as near the spring, in
Esparsa Canyon, in the arroyo bed of Long Canyon, and at West Anchorage.

According to Gunn (1979), . ludoviciana includes two subspecies, and ssp. /udoviciana
consists of five geographic races (phases) that are not given formal names: phase exigua,
corresponding to V. exigua Nutt., grows mainly in southern California and northwest Baja
California. Lassetter (1984) reached similar conclusions and in his fig. 5 showed his race
4 confined to California and Baja California, including Guadalupe Island.

Loasaceae
Mentzelia micrantha (Hook. & Arn.) Torr. & A. Gray, Fl. N. Amer. 1:535. 1840.

Mentzelia dispersa of Guadalupe Island references.

RANGE. — California to NW Baja California; Santa Cruz, Santa Catalina, and San Cle-
mente Islands.

Palmer found this plant in ravines in the “middle” and at the “south end” (Watson
1876:115); and on his second trip he reported it common in canyons at the south end

120

GUADALUPE ISLAND FLORA

(Vasey and Rose 1890:24). Greene (1885:223), failing to find M. dispersa of Watson’s
list, reported M. micrantha as an addition to the flora, seen only on the beach near the
landing and probably a recent introduction. It is widespread mostly at low elevations,
mostly in arroyo beds, from Northeast Anchorage to Arroyo Melpomene, but nowhere
common.

Although Watson and Vasey and Rose identified Palmer’s specimens as M. dispersa S.
Watson, Darlington (1934:193) cited at least one of his specimens as M. micrantha and
cited no Mexican collections of MM. dispersa. Harry Thompson in 1975 verified my six
collections as M. micrantha.

Malvaceae

Lavatera lindsayi Moran, Madrono 11:158. 1951.

RANGE. — Endemic to Guadalupe Island, and known there only on Outer Islet and Middle
Rock.

Open shrub to 7 dm tall and Il dm wide, the stem to 4 cm thick; herbage velvety with a
dense stellate puberulence. Leaf blades round-cordate with sinus nearly closed, 3-9 cm
wide, 5- to 7-lobed one-third of way to base or less, the lobes rounded, coarsely crenate;
petioles 2-5 cm long. Flowers several in upper axils, pendent on jointed pedicels nearly
equalling flowers, campanulate, 4.5—5.5 cm wide. Involucel 8-15 mm long, of 3 oblong
rounded bracts united in lower third. Calyx 20-25 mm long in flower, to 32 mm long in
fruit, divided nearly to middle into triangular-ovate acute or acuminate segments. Petals
erect or slightly ascending but not widespreading, deep purple throughout, with darker
veins, obovate-spatulate, emarginate, 3—5 cm long, 10-18 mm wide. Staminal column 25—
35 mm long. Fruit 9-12 mm wide, enclosed in calyx; mericarps 8-10, smooth, |-seeded.
This shrub flowers January to August (Figs. 65, 66; Carlquist 1965, fig. 14-7).

This shrub grows on the north outer slope, on the inner slope, and in the upper valley of
Outer Islet, from 20 to 190 m, and I also saw it in 1970 on Middle Rock; it has never
been found on the main island. Flowers were collected in January, February, and April,
with L. occidentalis, and most plants were still flowering in June and August, when L.
occidentalis was past flowering and largely leafless. In the upper valley where the two
occur together, I found a single plant that seemed to be a hybrid (4/5650); the petals are
erect below, spreading from near the middle, whitish with 5 diffuse purple stripes and
suffused with purple between.

Lavatera lindsayi differs from L. occidentalis in its denser puberulence, smaller and more
shallowly lobed leaves, later flowering, and more erect purple petals (Plate 2G). Since
the two grow together and since their flowering times overlap, the apparent rarity of
hybrids is noteworthy. Moreover, plants of L. occidentalis on Outer Islet are even less
puberulent than other collections of that species and agree with them in floral characters,
thus in these respects showing no approach to L. lindsayi. Yet it is remarkable that of all
collections of L. occidentalis, those from Outer Islet (Fig. 68) most closely approach L.
lindsayi (Fig. 66) in leaf shape, having comparatively short and rounded lobes.

Lavatera occidentalis S. Watson, Proc. Amer. Acad. Arts Sci. 11:124. 1876.

Lavatera insularis S. Watson, Proc. Amer. Acad. Arts Sci. 12:249. 1877.

RANGE. — Endemic to Guadalupe and South Coronado Islands.

12]

REID MORAN

FIGURES 65 and 66. The endemic Lavatera lindsayi on Outer Islet. FIGURE 65. Flowering shrub
(M/5118), 21 June 1968. FIGURE 66. Flowering branch (A/5648), 11 February 1957

122

GUADALUPE ISLAND FLORA

Summer-deciduous shrub 0.5—3 m tall and to 2m or more wide, or hanging from cliffs
to 4m, the trunk to | dm thick, the herbage thinly stellate-puberulent. Leaves round-cor-
date, 6-16 cm wide, 7-lobed a third to nearly half way to base, the lobes rounded to
subacute, to 3.5 cm long and 3 cm wide, coarsely crenate, the sinus nearly closed to open
at 110-130°, the petioles 3-23 cm long. Flowers axillary, solitary, on deflexed pedicels
2-3 cm long and jointed just above middle. Involucre of 3 subequal bractlets 7-10 mm
long, 2-5 mm wide, connate |.5—3 mm. Calyx in flower 25—28 mm long, the lobes fo-
liaceous, 18-19 mm long, 12-13 mm wide; in fruit 30-36 mm long or 5.5—6.6 cm wide,
the lobes spreading, ovate, acute, veiny, (15—) 18-25 mm long, (10—) 15-19 mm wide.
Petals ascending in lower third (within calyx), then spreading or somewhat recurved,
narrowly spatulate with a broad naked claw, emarginate, 4-6 cm long, 13—15 mm wide
above, +4 mm wide at base, greenish white below, whitish diaphanous above, with 3
faint narrow diverging purple stripes near center above. Staminal column + 25-27 mm
long, the broad base dark purple. Style + 9 mm long, the lobes + 22 mm long. Fruit 12—15
mm wide, enclosed in calyx, finely pubescent; carpels 6-10, 4.5—5.5 mm high, about
equalling the short conical summit of the axis. Seeds dark brown, 4.5 mm wide. The
shrub flowers January to April and was seen flowerless and nearly leafless in June and
August.

Palmer found this shrub conspicuous on the cliffs in the “middle” of the island, only
rarely and with difficulty accessible, in flower and immature fruit in April (Watson
1876:113). Greene (1885:222) called it a larger shrub than described, the large ones 10
feet high. Franceschi (1893c:134) saw a few scattered specimens, all on the most inac-
cessible rocks east of the island, and found a few seedlings not likely to survive. Moran
(1951:157) found it common on the inner slope of the crater on Outer Islet. There are
also a small colony on the floor of the crater, at 30 m, and a few plants in the upper
valley. Here the plant is likely to survive, but on the main island it is now very rare and
only on cliffs. In 1960 two large dense shrubs grew on cliffs of the Lower Circus (M7835,
7835a), quite possibly in the place meant by Palmer and Franceschi, one near the base,
the other high out of reach (Fig. 67); but by 1981 the lower one was gone. In 1970 two
grew at 300m on the south rim of Esparsa Canyon at the mouth (M//8/45). In 1963, a
year when plants were somehow getting ahead of goats, Weber and McCoy found a single
seedling (//968) near the zigzag trail above the barracks, but I failed to find it there in
1965. And in 1965 I found a single young plant +1 m high on the bank of Arroyo
Melpomene at 400 m (M/2074), which I did not find again in 1967.

The plants of Outer Islet are small, rarely more than | m tall. The leaves are more sparsely
puberulent than in other collections and are lobed only about a third of the way to the
base, with rounded lobes, and the sinus is nearly closed (Fig. 68). The plants of Esparsa
Canyon had leaves with very open sinuses, which are similar to the leaves of Brandegee’s
collection (UC). Moran (1951:158) pointed out the close similarity of L. insularis S.
Watson, of Islas los Coronados, to L. occidentalis, of Guadalupe Island, suggesting that
it be included in the same species, as Fryxell (1988) did; but in 1985 George Krebs
annotated specimens from Coronado as L. insularis. That grows only on the south island,
where it seemed to be one continuous population, and the collections are rather uniform.
Collections from Guadalupe Island are variable in leaf shape, but some closely match the
specimens from the Coronado Islands. Greene (1885) said the flowers of L. insularis were
exactly those of L. occidentalis, and so it appears except that they average a little smaller,
+ 3-5 cm long. There is little to separate the two.

REID MORAN

FIGURE 67 and 68. The insular endemic Lavatera occidentalis, found only on Guadalupe and on
South Coronado Island. FIGURE 67. One of two plants high on cliff of Lower Circus (447835), 27
January 1960. FIGURE 68. Flowering branch (5643) on Outer Islet, 11 February 1957.

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GUADALUPE ISLAND FLORA
*Malva parviflora L., Demonstr. Pl. p. 18. 1753.

Malva borealis of Guadalupe Island references.
RANGE. — Native of Eurasia; a widespread weed in North America and elsewhere.

Palmer found this plant only on the richer open spots in the “middle” of the island, in
dense masses (Watson 1876:113), but after his second trip reported it now introduced all
over the island (Vasey and Rose 1890:23). Greene (1885:221) found it very common on
the eastward slope. It is now widespread over the island and some places locally common,
especially on the north slope.

Sphaeralcea palmeri Rose, Contr. U.S. Natl. Herb. 1:23. 1890.
RANGE. — Endemic to the southern half of Guadalupe Island.

Plant perennial with a stout woody crown to 2 cm thick, densely yellowish canescent with
very short hairs, those of the stems 0.15—0.20 mm long, with ca. 25 rays. Stems erect or
arching to prostrate, conspicuously angled, mostly 2-6 dm long and 2-4 mm thick and
dying back nearly to the thick woody base but in sheltered spots sometimes persisting
and to | m long and | cm thick. Leaf blades thick, rugose, with veins very prominent
beneath, broadly ovate to subrhomboidal, 3-veined from base and sometimes faintly 3-
lobed, at base cuneate, at apex obtuse to rounded or retuse, rarely acute, 2.5—-6 cm long,
2-4.5 cm wide, crenulate-toothed, the petioles stout, 14cm long, 1/5—1/2 as long as
blade. Inflorescence a narrow thyrse with lower branches not more than 5 cm long. Pedicels
stout, shorter than fruiting calyx. Calyx at anthesis 6—9 mm high, with ovate acutish
strongly ribbed lobes equalling in length the angular-ribbed tube. Petals widespreading,
white or pinkish, yellowish or yellowish green at base, conspicuously veined, narrowly
obovate with long narrow claws, 13-17 mm long, 4-10 mm wide, with dense tuft of hairs
on each margin near base. Column sparsely or rather copiously pubescent. Fruit higher
than hemisperical, about 2/3 as high as fruiting calyx. Carpels 12-17, with chartaceous
walls, 4-5 mm high, ca. half as wide, broadly and shallowly notched, the dehiscent part
erect, ovate, at apex rather narrow but obtuse, muticous, the indehiscent part forming
1/4-1/2 of the carpel, rather prominently reticulate; attaching threads short. Seeds 2, very
pubescent. (Description modified from Kearney 1935.)

Palmer, on his second trip, found this plant in all exposed parts on the south end of the
island (Vasey and Rose 1890:23). Moran (1951:159) reported it occasional on the crater
floor, Outer Islet. It is fairly common at West Anchorage and on the south end mesa and
also grows on Islote Negro (Fig. 69) as well as Outer Islet.

Rose based the species on Palmer S867 (type US; GH, K, NY, UC) and 868. Kearney
(1935) kept it distinct from S. su/phurea, his concept of the species based only on five
specimens of Palmer’s two collections. His key characters for S. palmeri were plant yel-
lowish-canescent (vs. whitish-canescent or tomentose), stems conspicuously angled (vs.
terete or nearly so), leaf blades with veins very prominent beneath, strongly cuneate (vs.
with veins not very prominent beneath, subcordate to subcuneate), calyx conspicuously
ribbed (vs. not conspicuously ribbed), and carpels about 17 (vs. about 12); and his de-
scriptions suggest various other differences. With the study of more material of both
species, these differences seem to hold up to some degree, though some of them become
less absolute. In particular, there is considerable overlap in the number of carpels, though
there appears to be some difference in the means. Kearney placed these two species by
themselves in his section Su/phureae, distinguished by the petals less than half as wide

125

REID MORAN

FIGURE 69. The endemic Sphaeralcea palmeri (M5988) on Islote Negro, 19 April 1957.

as long. His chart of hypothetical relationships shows this section as rather isolated but
closest to the Emorvanae. That section includes two species of central and southern Baja
California and one extending from southern Nevada and southeast California to northern
Sonora and northern Baja California. Fryxell (1988) closely followed Kearney’s treatment.

Sphaeralcea sulphurea S. Watson, Proc. Amer. Acad. Arts Sci. 11:125. 1876.
Sida hederacea and S. leprosa hederacea of Guadalupe Island references.

RANGE. — Endemic to Guadalupe Island.

Plant perennial, to + | m high and 1—1.5 m wide, with rather stout woody crown, densely
whitish canescent or tomentose with short hairs, those of stem 0.2—0.4 mm long, with
20-25 rays. Stems numerous, erect or arching from near base, to 5 mm thick. Leaf blades
usually thickish, with veins not very prominent beneath, broadly ovate, deltoid, or nearly
orbicular, at base subcuneate to subcordate, at apex very obtuse to acutish and sometimes
mucronate, 3—S-veined from base, shallowly 3-lobed at or below middle with rounded or
acutish lobes, the margin finely to rather coarsely crenate and often somewhat rugose,
the larger blades 3-9 cm long, 2/3 as wide to wider than long, the petioles slender or
rather stout, 1/3—1 1/2 times as long as blade. Inflorescence an interrupted many-flowered
thryse, narrow or with lower branches up to 12 cm long, and often forked. Pedicels rather
stout, usually much shorter than fruiting calyx. Calyx at anthesis 5-7 mm high, with
lanceolate or deltoid-ovate, acute or short-acuminate lobes 1—2 1/2 times as long as tube.
Petals in bud pale orange at tips, in anthesis spreading, white, greenish or yellowish at
base, conspicuously veined, oblanceolate or narrowly obovate with long narrow claws,
10-15 mm long and 1/3 to 1/2 as wide. Column sparsely pubescent. Fruit higher than

126

GUADALUPE ISLAND FLORA

hemispherical, + 2/3 as high as calyx; carpels + 2, with thin, chartaceous walls, 34.5 mm
high and 1/2 to 2/3 as wide, shallowly to rather deeply notched, the dehiscent part erect
or ascending, ovate or deltoid, obtuse, muticous to short-cuspidate, the indehiscent part
forming about 1/3 of carpel, prominently and somewhat coarsely reticulate; attaching
threads short or rather long. Ovules 2 or 3. Seeds 1—3, sparsely pubescent. (Description
modified from Kearney 1935.)

Palmer found this plant very abundant on rocky slopes and in the crevices of the highest
rocky ridges, from the “middle” of the island to the “southern end,” where it was most
frequent (Watson 1876:113). Greene (1885:216) found it quite abundant on the gentler
declivity leading to the plateau from the zigzag trail out of the canyon. Franceschi
(1893¢:134) called it much more abundant than lavatera, one of the very few plants of
which some meager specimens may be seen scattered about even in places occasionally
visited by goats; he saw seedlings and young plants near the landing, both on the beach
and on the dry lava rock. It is common on cliffs in the northeast part of the island, from
Northeast Anchorage and Esparsa Canyon to the Lower Circus, and south as far as the
lobster camp. In 1981 I saw it on western cliffs at 1075 m, northwest of the cypress
grove. It seems not to be at the north end. Kearney (1935:34) cited specimens of Palmer
(No. 18 GH, type; CM, F, K, MICH, MO, NY, PH, UC, US), Greene, Franceschi, Bran-
degee, Drent, Anthony, and Mason.

From a collection of Thoburn, Green, and Wing in July 1897, Dudley (1899:282) listed
“Sphaeraclea? Material scanty. Near northeast landing.” The specimen (DS 140703) was
annotated by T. H. Kearney as “Sida hederacea ?” and by I.D. Clement in 1948 as
“? Sida leprosa var. hederacea”; but Clement (1957:58) cited it as S. leprosa var. hederacea
(Dougl.) K. Schum., without the query. Most leaves are folded so that it is hard to tell
their shape, but some clearly are ovate and longer than in the sida. There is no question
that this is the sphaeralcea.

Nyctaginaceae

Mirabilis californica A. Gray in Torr., Bot. Mex. Bound. p. 173. 1859.

Hesperonia heimerlii Standl., Contr. U.S. Natl. Herb. 13:412. 1911.
Mirabilis heimerlii (Standl.) J. F. Macbr., Contr. Gray Herb. 56:24. 1918.
Mirabilis laevis of Guadalupe Island references.

RANGE. —SW California and NW Baja California; Santa Rosa, Santa Cruz, Anacapa,
Santa Barbara, Santa Catalina, and San Clemente Islands and islands of NW Baja Cali-
fornia.

Low shrub to 2 m wide and mostly 2—3 dm high, with many prostrate branches, the stem
to 2cm thick, with gray bark, the branches brittle, glabrous, glaucous, much swollen at
nodes. Leaf blades subsucculent, deltoid-ovate, cordate at base, obtuse or acutish, 1-4
(—5.5) cm long, to 4.5 cm wide, + | mm thick, usually glabrate but younger ones glandular
puberulent, the petioles half as long as blades or upper ones much shorter. Involucres
numerous in axils of leaves, on peduncles 0.5—3 mm long, or congested at ends of branches,
campanulate, often purplish, densely glandular-puberulent, 3-6 mm long at anthesis, S—8
mm long in fruit, the lobes deltoid to ovate, 1-3 mm long. Perianth purple, 8-12 mm
long, glandular-puberulent on parts exposed in bud. Anthocarp globular to prolate, 34.5
mm long, 2.5—3 mm thick, dark brown.

Palmer found this shrub in crevices of the walls of canyons on the east side (Watson

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REID MORAN

1876:118) and on his second trip reported it common at the south end (Vasey and Rose
1890:26). In the northeast part of the island it is not uncommon on beach cliffs, ascending
to 450 m on the north rim of Barracks Canyon at the mouth, and occasional on cliffs in
the canyons. In the southern part it is more common, especially near the shore but also
inland in arroyos and on mesas, at least to 350m, as well as on Islote Negro and
Outer Islet.

Standley (1911) based Hesperonia heimerlii on Palmer 886 (US22626, holotype), from
the south end of Guadalupe Island, and on collections of Franceschi and Rose. He named
it for Dr. Anton Heimerl, a student of the family, who had marked specimens of it as a
new species. Standley said that the form of the fruit (“spherical or slightly depressed
vertically”) placed it near M. cedrosensis Standley, from which it differed in its pubescence,
the plants “glandular pubescent, never scabrate.” Likewise, Macbride (1918) separated
Mirabilis heimerlii by its “truly globose fruit.” Wiggins (1980:149) and Wallace (1985:21)
also recognized M. heimerlii as a Guadalupe endemic. Recent collections, however, show
that the fruit varies and is often somewhat longer than thick.

Since Ferris (1964) has clarified the distinction of M. californica A. Gray from M. laevis
(Benth.) Curran, the Guadalupe plant clearly belongs to the M. californica complex, agree-
ing in the form of the involucre. It is perhaps woodier than typical M. californica and
tends to differ in its shorter anthocarp and perhaps larger leaves. From var. cedrosensis
it differs in its pubescence, which is fairly uniform in all collections. No combination of
characters is evident to separate it from all other members of this complex, and it is
similar to some plants of San Clemente Island.

Oleaceae

+Hesperelaea palmeri A. Gray, Proc. Amer. Acad. Arts Sci. 11:83. 1876.
RANGE. — Endemic to Guadalupe Island.

This was a rather compact glabrous tree 20 to 25 feet high, with entire coriaceous mostly
opposite leaves and terminal panicles of perfect 4-merous lemon-colored flowers, and
with fruit undoubtedly drupaceous (Fig. 70).

Palmer found only three live trees, in a canyon on the east side; he saw no young trees
but many dead ones (Watson 1876:118). Brandegee (1900:22) remarked that goats were
not the only exterminators of endemic species, “for | searched in vain for Hesperelaea at
the exact locality where Dr. Palmer found it and could only find a goat corral made from
trees chopped down in the vicinity.” No one has seen it since, and it is undoubtedly
extinct.

Gray called this genus remarkable in the Oleaceae for its wholly distinct and unguiculate
petals, imbricate above, and its apparently uniformly isomerous stamens. The genus has
remained monotypic and so is endemic to Guadalupe Island.

Onagraceae

Camissonia guadalupensis (S. Watson) Raven ssp. guadalupensis
Oenothera guadalupensis S. Watson, Proc. Amer. Acad. Arts Sci. 11:137. 1876.

Camissonia guadalupensis (S. Watson) Raven, Brittonia 16:284. 1964.
Oenanthe guadalupensis of Guadalupe Island references.

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GUADALUPE ISLAND FLORA

cm 2 3 4 5 6 7 8 9 10

FIGURE 70. Flowering branch of the remarkable Hesperelaea palmeri, a small tree of the Oleaceae,
now extinct. In 1875 Palmer found many dead but only three still alive; no one has found it since.
Hesperelaea was an endemic monotypic genus of Guadalupe Island.

RANGE. — Endemic to Guadalupe Island.

Palmer found only two plants, in a ravine on the east side near the beach (Watson
1876:115). In 1958 Ernst and I found seven plants widely scattered in the bed of Arroyo
Melpomene at 450 m (Moran and Ernst 6737) (Plate 2H), and in 1965 I found two more
in the same arroyo at ca. 300 and 400 m (M/2070). Raven studied plants raised from the
seeds of the first collection, as Raven 17530 and 18175 (DS).

In a list of plants known only from Guadalupe Island, Vasey and Rose (1890:22) included
“9. Oenanthe guadalupensis Wat.” There is no such name in Oenanthe, and no member
of that genus has been found on Guadalupe Island or seems likely to be. The similarity

129

REID MORAN

of the generic name, the use of the same specific epithet and author, and the position in
the list, together show that this is an error for Oenothera guadalupensis S. Watson. Joseph
Rose’s field notebook for March 1911 (US) shows Rose 16901 as Oenanthe guadalupensis,
suggesting that he may have found the rare endemic camissonia in the same part of the
island where Palmer found it. However, Dr. Raven wrote me that he was unable to find
Rose’s specimen at US.

Raven (1963:332) described a ssp. clementina from San Clemente Island, where he later
(1969:275) reported it as common on dunes around the north end and down the west
shore, up to 100 feet elevation. He found it to differ in its spreading rather than appressed
strigose pubescence, in the presence of short glandular trichomes in the inflorescence,
and in the presence of a red dot near the base of each petal. Raven (1969) found both
plants autogamous, with a gametic chromosome number of 7 = 7. Reciprocal Fy hybrids
showed no evidence of chromosomal structural heterozygosity, but reduced stainability
of pollen suggested a substantial reduction in fertility in the hybrids, although seed-set
did not appear to be impaired.

dCamissonia robusta Raven, Brittonia 16:284. 1964.

Oenothera micrantha of Guadalupe Island references.

RANGE. — Southern California to NW Baya California; San Miguel, Santa Cruz, Santa
Catalina, and San Clemente Islands.

Moran (1951:160) found this plant in the canyon mouth at Northeast Anchorage, where
it has remained fairly common in later years (Fig. 71). It has also been found near the
spring (Lindsay in 1948), rather scarce on the northwest slope at 550 m (1/7295), and
near the north rim of Barracks Canyon at 500 m (A//3804).

Because this plant was fairly common at the landing where every botanical visitor to the
island had come ashore without finding it, Moran (1951:160) thought it very likely a new
arrival. Raven (1963:298) at first also called it probably introduced. Judging later from
the phytogeographic relationships of Guadalupe Island and from the scarcity of the plant
on San Clemente Island, however, Raven (1969:305) thought it equally likely that it might
simply have gone undetected on Guadalupe before and have become more common re-
cently. He added that it probably will not be possible to resolve this question definitely.
I list it with those west American plants that are probably introduced (Table 2).

A missing piece to this puzzle may possibly be the Rose collection (/690/) of “Oenanthe
guadalupensis,” mentioned under the previous species. Was that really C. guadalupensis,
or could it possibly have been C. robusta? If C. robusta, it would show that this plant
was already on the island by 1911 and then perhaps more likely native.

+Epilobium foliosum (Torr. & A. Gray) Suksd., Deutsch. Bot. Monatss. 18:87. 1900.

Epilobium minutum Lindl. var. foliosum Torr. & A. Gray, Fl. N. Amer. 1:490. 1840.
Epilobium minutum of Guadalupe Island references.

RANGE. — Vancouver Island and Idaho to southern California; Guadalupe Island.

Palmer found this plant only at the north end, among rocks and sage brush (Watson
1876:115). Greene (1885:223) found two or three plants in the same area. No one has
seen it since, and it is doubtless extinct on the island.

Seavey, Wright, and Raven (1977) found £. foliosum distinct from E. minutwm Lindl. in

130

GUADALUPE ISLAND FLORA

several respects, including chromosome number. Palmer’s Guadalupe Island collection
(K, MO, NY, UC) was hard to assign to either species, but from the flower size they
tentatively called it E. foliosum.

Papaveraceae

Dr. Wallace E. Ernst collected poppies on Guadalupe Island in April 1958 and reported
some chromosome counts (Ernst 1958). He helped me straighten things out.

+bEschscholzia californica Cham. var. californica

Eschscholzia californica Cham. in Nees, Hort. Phys. Ber. p. 73. 1820.
Eschscholzia gigas Fedde, Repert. Nov. Sp. 3:28. 1906.

RANGE. — Northern to southern California; San Miguel, Santa Rosa, Santa Cruz, and
Guadalupe Islands.

Greene (1885:221) found this plant “luxuriating in one place only, near the edge of a
precipice, northeast of the cabins,” i.e. near the top of the Lower Circus. Franceschi
(1893c:133) reported it plentiful in the same limited locality, its leaves clipped pretty
closely by the goats. No one has found it since, and it doubtless is extinct on the island.

Greene described the plant as follows: “Root perennial: stem robust, two feet high: petals
orange, two inches long. The true E£. californica.” Franceschi affirmed that it was perennial.
Greene (1885:183) wrote that E. californica did not appear in southern California except
at considerable altitudes in the mountains, as at San Gorgonio Pass and Tehachapi Pass,
and he mentioned his collection from Guadalupe Island. He said that in all these localities
it was the same robust large-flowered perennial so well known from San Francisco to the
borders of British Columbia. Later, however, Greene (1905a) restricted E. californica to
the San Francisco Peninsula but did not refer the Guadalupe specimen to any other species.
Based only on Greene’s collection (GH holotype?), Fedde in 1906 named FE. gigas, thus
a Guadalupe endemic, which he thought doubtless related to E. crocea and E. rigida but
different in habit and in the form of the hypanthium. Dr. Ernst wrote me in 1962 that
Greene’s Guadalupe Island collection of 24 April 1885 at GH was annotated by Fedde
as E. gigas Fedde. Ernst called this the giant coastal form of £. californica, the only
specimen of it he had seen from Guadalupe Island.

Greene thought this plant was “possibly of recent introduction.” Although conceivably
the goat hunters may have beautified the premises by planting California poppies, the
ability of Eschscholzia to reach this island is shown by the presence of three other species;
and the finding of this unexpected northern one is consistent with the occurrence here of
such other northern plants as Polypodium scouleri, Ribes sanguineum, and Eriophyllum
lanatum. var. grandiflorum. On the other hand, since Palmer had camped nearby for 15
weeks without finding this conspicuous plant, or three other plants later found by Greene,
very likely it was a late arrival and not well established. I therefore list it with the west
American plants probably introduced on the island (Table 2).

Eschscholzia elegans Greene, Bull. Calif. Acad. Sci. 1:182. 1885.
Eschscholzia californica var. hypecoides of Guadalupe Island references.
RANGE. — Endemic to Guadalupe Island?

Erect annual 3-6 dm high, branching above base, glabrous, very glaucous. Leaves finely
dissected, their ultimate divisions long and parallel or shorter and divergent, linear or

131

REID MORAN

oblanceolate, acute. Calyptra ovoid, apiculate, 10 mm long. Torus cylindrical, the outer
rim corky, the inner erect and hyaline. Corolla to 35 mm wide, the petals spreading with
margins hardly meeting, 8-20 mm long, greenish yellow when fresh, turning orange on
drying. Stamens 16-20, the filaments and anthers about equal. Pods thin-walled, slender,
6-9 cm long. Seeds slightly elongated, apiculate, tuberculate, the raphe obvious. Chromo-
somes: 1 = 17 fide W. Ernst.

Palmer found this plant only at the “south end” in ravines and in the “middle” of the
island on level ground (Watson 1876:112). Greene (1885:183) called it. common in the
middle and southern parts of the island, the type his collection at CAS? I found it common
on the alluvial fan at West Anchorage (M5604, M5956).

Eschscholzia palmeri Rose, Contr. U.S. Natl. Herb. 1:23. 1890.

Petromecon palmeri (Rose) Greene, Pittonia 5:293. 1905,
Petromecon frutescens Greene, Pittonia 5:294. 1905.
Eschscholzia frutescens (Greene) J. T. Howell, Leafl. W. Bot. 3:149. 1942.

RANGE. — Endemic to Guadalupe Island.

Shrub to 5 dm high and | m wide, erect or with decumbent branches, with soft-woody
stem to 4 cm thick, the bark corky, golden brown to gray, orange in fissures; but flowering
the first season when still tiny and herbaceous. Foliage commonly dense, very glaucous
when young, the leaves alternate, pedately dissected into moderately divergent long nar-
row-linear subterete acute to obtuse segments, the petiole linear, to 4cm long. Pedicels
erect, rising above foliage, persisting stiff and conspicuous, to 17 cm tall. Torus a hemi-
spherical subligneous cup, the outer margin a thick herbaceous ring, the inner elevated,
thickish, corky. Calyptra almost spherical to ovate-conic, with short blunt point, 5-15 mm
long. Corolla 1.5-6 cm wide, the petals spreading, bright yellow with orange basal spot,
cuneate-obovate, 8-26 mm long. Stamens 50-80 or more, the filaments 3-10 mm long,
filiform above a flattened base, the anthers linear?, 2.5-3 mm long. Stigmas 4, not greatly
elongated, clavellate. Pods 3—7 cm long, hard and subligneus, the valves fluted. Seeds
with no superficial papery coat, appendage, or reticulation, but surface uneven by short
interrupted subsinuous rugosities. Cotyledons linear, 1.5—3 cm long, + | mm wide. Chro-
mosomes: n= 11 fide W. Ernst.

Palmer found this plant at the south end of the island, where he saw it only on a rocky
ledge, but there common (Vasey and Rose 1890:23). In the northern part of the island it
is occasional on seacliffs and cliffs in the canyons and on the northwest cliff at 600 m
(M17281). It is more common on the mesa at the south end and on Islote Negro and
Outer Islet, where many seedlings may be seen flowering in a good year (Plate 21).

Greene (1905b) proposed the endemic genus Petromecon for E. palmeri and his new P.
frutescens. He said the two species had something of the habit of the dendroid insular
eschscholzias but a pedatifid foliage and the flowers of Stvlophorum as to the excessive
number of stamens, 80 or more, on long filaments. He added that the torus and the stigmas
approached those of Eschscholzia but were different, as also were the seeds. Fedde (1909,
1936) continued to recognize Petromecon as an endemic genus of Guadalupe. On the
other hand, Howell (1942), Ernst (1958), and Wiggins (1980) returned it to Eschscholzia.
Eschscholzia palmeri clearly is a very distinct species but I think no more.

Greene (1905b) based Petromecon frutescens on Franceschi’s collection of January 1893
from “rocky ledges on the northern slope of Guadalupe,” the holotype at CAS. Evidently,

132

GUADALUPE ISLAND FLORA

however, this is the plant mentioned by Franceschi (1893c:134) under £. elegans var.
ramosa: “perennial on a nearly inaccessible cliff of lava detritus on the right bank of the
canyon 500 or 600 feet from the landing.” Howell (1942:149) reported it from ocean
bluffs inaccessible to goats near Northeast Anchorage. Greene described the species as
much larger than P. pa/meri, a foot high and as broad; growing branches and their foliage
less succulent; older and naked branches wholly and solidly woody, the wood firm, com-
pact, and hard; leaves ample, on long and slender petioles, pedately divided and cleft into
long narrow-linear moderately divergent segments; calyx ovate-conic; corolla cruciform,
the cuneate-obovate petals not meeting; stigmas 4, rather short, equal; pods and seeds not
known.

Ernst (1958) reported n= 11 for E. palmeri from the south end. Considering plants on
cliffs in the canyons to be E. frutescens, that may be a shrub to | m wide and 5 dm high,
with a trunk to 4cm thick. In May 1967 I compared fresh material of M@//3797, from
Esparsa Canyon, and M/3762, from the talus of the seacliff northeast of Campo Sur,
presumably typical E. palmeri, with various herbarium specimens. M/3797 was less com-
pact, and the ultimate segments of the leaves more slender, as might be expected from
the habitats. The petals were larger, to 26 mm long as compared to 19 mm, and the orange
spot was less pronounced. The stigmas were four in both, contrary to the distinction
implied by Greene. Plants on Islote Negro, are larger and woodier than Greene supposed,
so that difference doesn’t hold. Dr. Ernst got a count of n= 11 for M/3797, as he did
for plants from the south end. He wrote me in 1958 that he thought E£. frurescens was
just E. palmeri growing perhaps a bit shaded and out of the wind on the east side of the
island, and I certainly do not see any way to distinguish it.

Eschscholzia ramosa (Greene) Greene, Bull. Torrey Bot. Club 13:217. 1886.

Eschscholzia elegans var. ramosa Greene, Bull. Calif. Acad. Sci. 1:182. 1885.

RANGE. — Islands of southern California and northern Baja California, except San Miguel
and Anacapa.

Erect compact treelike annual, 2-4 dm high, very glaucous, the stem simple below, com-
pactly branching above. Leaves finely divided, the ultimate lobes short and divergent,
often blunt. Torus somewhat urceolate, the rim membranous, and outer rim lacking. Ca-
lyptra ovoid, 5mm long. Corolla smaller than in £. elegans, reddening less in drying,
12-25 mm wide, the petals 6-13 mm long. Stamens 16—20?, the filaments 1—2 mm long,
the anthers 2-3 mm long. Pods stout and straight, 6-9 cm long, strongly few-striate. Seeds
nearly globular, distinctly reticulate. Cotyledons linear, entire, 2.54.5 cm long, ca. | mm
wide. Chromosomes: 1 = 12 fide W. Ernst.

Greene (1885:183) based this species on plants found under high cliffs near the landing
at the northeast end of the island. He wrote that the erect compact tree-like habit was
peculiar. It is the more common annual species on the beach and near the shore on both
sides of the island.

Dr. Ernst wrote me in 1958 that although £. elegans and E. ramosa are both variable,
he thought he could distinguish all the material of them from Guadalupe Island, and he
sent me an illustrated key. He said that once he recognized that there were two entities
with different chromosome numbers and could release himself from some previous preju-
dices, he found these differences more easily seen. He said that, in fact, he felt more and
more like a true spiritual son of the late E. L. Greene. He added, however, that until he
could see fresh material from the other islands, he would be very cautious about extra-

133

REID MORAN

Guadalupe determinations. According to Raven (1963:322), Dr. Ernst then considered E.
elegans possibly restricted to Guadalupe Island, while plants resembling £. ramosa were
more widely distributed, having been collected on Santa Cruz, Santa Barbara, Santa
Catalina, San Clemente, Guadalupe, Coronado, Todos Santos, San Benitos, and Cedros
Islands, though not on the mainland. Wiggins (1964, 1980) included £. ramosa in E.
elegans, with a range from Santa Rosa Island to Guadalupe and Cedros Islands.

+Platystemon californicus Benth., Trans. Hort. Soc. London, II. 1:405. 1835.

RANGE. — Oregon to Nevada, Arizona, and NW Baja California; islands of southern Cali-
fornia except San Clemente.

Brandegee (1900:22) reported this plant from Guadalupe Island but with no specific lo-
cality. No one has found it since, and it probably is extinct on the island.

Without explanation, Brandegee called this plant a recent introduction. The fact that it
has not been found before or since may suggest that it was a waif that never became
established. However, Brandegee made the same statement about Lasthenia californica,
which seems clearly to be native. Furthermore, although the absence of the poppy from
San Clemente, the most oceanic of the Channel Islands, is not reassuring, its presence
on the other seven islands does show some ability to reach islands. The question is im-
possible to resolve, and I list this plant as possibly introduced on the island (Table 2).

Plantaginaceae

Plantago ovata Forssk., Fl. Aegypt.-Arab., p. 31. 1775.

Plantago insularis Eastw., Proc. Calif. Acad. Sci. Ser. 3, 1:112. 1898.
Plantago patagonica of Guadalupe Island references.

RANGE. — Southern California and offshore islands to central Baja California and Cedros
Island; also Old World.

Palmer found this plant in level spots at the “south end” (Watson 1876:116). In favorable
seasons it is common in the southern half of the island, including Islote Negro, and
occasional farther north.

Stebbins and Day (1967) showed that P. insu/aris, described from western North America,
was more closely related to the Old World P. ovata than to other American plants. On
the basis of chromosome differences and apparent small morphological differences, they
kept it a separate species, and they argued that it must have reached the New World by
natural means, probably at the beginning of the Miocene. Bassett and Baum (1969) treated
P. fastigiata (including P. insularis) as conspecific with P. ovata and thought it probably
introduced into the New World comparatively recently by man. When it arrived remains
uncertain, but | tentatively follow Dempster (1993) in not separating the American plants
taxonomically. There is more to learn.

Polemoniaceae
Note. — For Navarretia hamata see Excluded Species, p. 169.
+Allophyllum gilioides (Benth.) A. D. & V.E. Grant, Aliso 3:105. 1955.
Collomia gilioides var. glutinosa and Gilia divaricata of Guadalupe Island references.

134

GUADALUPE ISLAND FLORA
RANGE. — Oregon to southern California and Arizona; Guadalupe Island.

Palmer found this plant abundant under brush and in protected places in the “middle” of
the island (Watson 1876:118). Greene (1885:225) listed it without comment, but Grant
and Grant (1955:105) cited Greene’s collections. No one has found it since, and very
likely it is extinct on the island.

Gilia nevinii A. Gray, Syn. Fl. N. Amer. ed. 2, 2':411. 1886.

Gilia multicaulis Benth. ssp. nevinii (A. Gray) H. Mason & V.E. Grant, Madrono 9:209. 1948.
Gilia multicaulis and var. millefoliata of Guadalupe Island references.

RANGE. — Endemic to Santa Rosa, Santa Cruz, Anacapa, San Nicolas, Santa Barbara,
Santa Catalina, San Clemente, and Guadalupe Islands.

Corolla lavender to deep blue-violet, not spotted.

Palmer found this plant very abundant under brush and in protected places in the “middle”
of the island (Watson 1876:118). Franceschi (1893c) reported it very common among
rocks over the whole island. It is widespread, from the northwest slope and pine ridge to
the mouth of Arroyo Melpomene. It grows especially on grassy slopes, where in good
years it is often abundant, especially in the higher northern part of the island.

Linanthus pygmaeus (Brand) J. T. Howell ssp. pygmaeus

Gilia pygmaea Brand, Pflanzenreich aaeles 8 1907.

Gilia guadalupensis Brand, Pflanzenreich 4~":134. 1907.
Linanthus pygmaeus (Brand) J. T. Howell, Leafl. W. Bot. 2:100. 1938.
Gilia pusilla and var. californica of Guadalupe Island references.

RANGE. — Endemic to San Clemente and Guadalupe Islands.
Corolla lobes blue to lavender, throat yellow or white, and tube yellow.

Palmer found this plant abundant under brush and in protected places in the “middle” of
the island (Watson 1876:118). It now grows in grassy and rather bare places, mostly at
400 to 1000 m but lower in the canyons, from the pine ridge and cypress grove to El
Picacho, in good years locally abundant.

The type is Palmer 79 in part (where?), from Guadalupe Island. Grant (1959:137) also
referred here G. guadalupensis Brand, based on Palmer 79 in part. Raven (1963:334)
separated the mainland material as ssp. continentalis.

Polygonaceae
Note. — For Harfordia macroptera see Excluded Species, p. 169.
Eriogonum zapatoense Moran, Trans. San Diego Soc. Nat. Hist. 15:267. 1969

Eriogonum molle of Guadalupe Island references.
RANGE. — Endemic to Guadalupe Island: known only on Outer Islet.

This is a shrub, with trunk to 1 dm thick, densely tomentose herbage, and 4-8 times
trichotomous cymes of yellow flowers (Fig. 72). In protected places it may grow erect,

135

REID MORAN

bse.
ai

FIGURE 71. Camissonia robusta above Northeast Anchorage, 25 March 1973. Now common where
every early botanist came ashore, it was not found until 1948. Can it be native?

FIGURE 72. The endemic Eriogonum zapatoense (M18170) on Outer Islet, 21 August 1970. This
shrub has never been found on the main island.

GUADALUPE ISLAND FLORA

but most plants are prostrate, commonly with the trunk or the few main branches running
zigzag up slope, so that the plant is strongly asymmetrical, in many larger plants the
lower branches dead and the foliage clustered at the top.

This shrub is common on the inner slopes of the crater, especially the south slope, and
occasional in the upper valley, of Outer Islet, sometimes known as Islote Zapato. From
sterile plants found in 1948, Moran (1951:154) reported it as E. molle Greene. It is most
similar to E. molle, which is endemic to the north end of Isla Cedros, and to E. giganteum
S. Watson, of Santa Catalina, San Clemente, and Santa Barbara Islands, California. In
leaf shape especially, it resembles E. giganteum ssp. formosum (K. Brandegee) Raven,
of San Clemente Island. Both £. molle and E. giganteum grow erect, and E. giganteum
often much taller; and both have somewhat thinner leaves, commonly taller and stouter
peduncles, and much larger and, especially in £. molle, denser cymes, with pubescent
pedicels, and with white or pink rather than yellow flowers. The holotype is Moran 15114,
collected 21 June 1968 (SD67852).

Pterostegia drymarioides Fisch. & C. A. Mey., Ind. Sem. Hort. Petrop. 2:48. 1835.

RANGE. — California to Utah, Arizona, and NW Baja California; islands of southern Cali-
fornia and south to Cedros.

Palmer found this plant in the shade of rocks in the “middle” and more rarely at the
“south end” of the island (Watson 1876:119). It grows among rocks throughout the island,
from the north slope to the south end and from sea level to the top of Mt. Augusta and
EI Picacho, often rather common but less so at the south end.

Portulacaceae

Calandrinia ciliata (Ruiz & Pav.) DC., Prod. 3:359. 1828.

Calandrinia menziesii Hook., Fl. Bor. Amer. 1:223. 1834.
Calandrinia ciliata (Ruiz & Pav.) DC. var. menziesii (Hook.) J. F. Macbr., Field Mus. Bot. Ser.
11:20. 1931.

RANGE. — British Columbia to Arizona, Sonora, and central Baja California; South Amer-
ica; islands of southern California and Islas Todos Santos and San Martin.

Palmer found this plant in moist spots in the open valleys all over the island, growing in
masses (Watson 1876:113). Greene (1885:221) remarked that it was much smaller than
in California, always prostrate, an albino form very frequent. It is now one of the common
plants of grassy slopes from shore to summit in the northern part of the island and extends
to the south end in damp arroyo beds.

Cistanthe guadalupensis (Dudley) Carolin in Hershkovitz, Phytologia 68:269. 1990.

Talinum guadalupense Dudley in Jordan, Fur Seals and Fur-seal Islands N. Pacific Ocean, Part 3,
p. 282. 1899.

RANGE. — Endemic to Guadalupe Island.

Succulent shrub to 8 dm high when not in flower and to 18 dm wide, summer-deciduous
in dry sites, the stem to 11 cm thick at base, branching from base and above, the lower
branches in large plants sprawling; bark smooth, brown, usually with dull silvery sheen,
with slight tendency to peel in papery layers; ultimate branches blunt at apex, crowned

137

REID MORAN

with leaf rosette, 1.5—4 cm thick below rosette, the leaf scars irregular, lunate to rhombic,
6-14 mm wide, 3-6 mm high, the bundle scar elliptic, 2-3 mm wide, 1.5—2 mm high.
Rosettes lax, 0.5—2 (—3.5) dm wide, of 10-35 (—S5) leaves, only the youngest close-set,
the older separated by areas of stem greater than those they cover. Rosette leaves obovate
to obovate-cuneate, rounded to truncate or emarginate at apex and usually apiculate, 3-10
(—17) cm long, 3—5 (1.5—8.5) em wide, 3—7 mm thick, bluish glaucous, the margins obtuse,
often purple-red. Floral stems terminal, 1—S (—9) dm tall, 3-12 mm thick at base, bearing
12-20 scattered bracts and, from axils of upper ones, 3-10 spreading to ascending mostly
simple branches 3-10 cm long, each ending in a crowded cincinnus or sometimes two or
rarely several; panicle 1-3 dm high, 1—2.5 dm wide. Lower bracts like reduced foliage
leaves, those of inflorescence thin, broadly ovate, acutish, cupped, keeled, yellow-green
with conspicuous purple-red veins, 3-10 mm long, 3-8 mm wide, usually drying persistent.
Cincinni circinate in bud, of 5-13 flowers, the axis conspicuously jointed, its segments
mostly 3—5 mm long, or the lower longer, 1.5—3 mm thick at base, 2-4 mm thick at apex,
each seated in hollow in axil of sheathing bract. Pedicels 2—S mm long, 1—3 mm thick at
base, thickening upward, each with small basal bract and shallow basal constriction, where
abscission may later take place. Buds ovoid to subglobular, rounded at apex, to 9 mm
long and 8 mm thick. Flowers (Plate 2J) at least Jan—June, 2.54.5 cm wide, opening
mostly two days and partially closing at night, subperigynous, the stamens and petals
inserted on edge and outer shoulder of cup 1-2 mm high closely surrounding base of
ovary. Sepals 2, broadly ovate to reniform, rounded at apex, imbricate in bud, 6-10 mm
long, 7-13 mm wide, colored and marked like bracts, persisting in fruit. Petals 5 (—7),
imbricate, rotately spreading to slightly upcurved, rose pink to almost white, obovate,
often cuneate at base, subtruncate to emarginate, erose, 13—23 mm long, 10-22 mm wide,
connate + 0.5 mm. Stamens 70-105, the filaments pale to deep rose, 8-14 mm long, the
outer filiform, glabrous, the inner thickened and connivent in lower fourth and there
bearing colorless clavate unicellular hairs to | mm long, concealing ovary; anthers cream
to deep rose, oblong, 1.2—2 mm long, 0.5—0.7 mm wide, maturing in centrifugal order,
the two thecae rather weakly united, free in lower half and upper fourth. Gynoecium
mostly 3-merous; ovary free, subglobular, 2-3 mm thick, with 3 (2-4) low vertical ridges
alternating with faint grooves; style white, 10-16 mm long, tapering from + 0.4 mm thick
at base to + 0.8 mm thick at apex; stigmas 3 (2-4), spreading, oblong, rounded at apex,
1.5-2 mm long, ventrally papillose, the papillae + 0.1 mm long and a fourth as thick.
Capsule cartilaginous, brown with oily lustre, triangular-ovoid, 6-8 mm long, 5—6 mm
thick, capped nearly to maturity with dark purple corolla remnant, splitting nearly to base,
the valves triangular, subacute. Seeds numerous, lenticular, strophiolate, finely low-mam-
millate, dark red-brown and somewhat shining but largely covered with scurf, 0.9-1.2
mm long, 0.8-1.0 mm wide, 0.5—0.6 mm thick, the raphe rather acute, the opposite edge
rounded. Chromosome number: 7 = 48 fide Sherwin Carlquist.

Botanists of the fur seal expedition were first to find this curious succulent shrub (Dudley
1899): the specimen is noted as “in valley, south end of island.” Thirty years later a single
living plant, found by Steve Glassell and Bill Pemberton, reached Eric Walther, who
showed first the sterile plant (1932), then (1937) the flowering plant with an identification.
Howell (1942:148) saw what he took to be this plant “on rocks inaccessible to goat and
man” near Northeast Anchorage. Moran and Lindsay (1950) found it plentiful on Outer
Islet (Figs. 34, 75) and on a black lava rock just off the landing at the southwest corner
of the island; and Lindsay (1951) showed several fine photographs of it on this rock and
told its remarkable story. It is common also on Islote Negro (FRONTISPIECE, Figs. 73,
74)(Moran 1959, figs. 21-23). In fact, after four days circumnavigating the island in a
small launch in search of fur seals, George Lindsay wrote me in 1955 that it grows on

138

GUADALUPE ISLAND FLORA

FiGuRES 73 and 74. The endemic Cistanthe guadalupensis, rare on the main island, common on
Islote Negro (see FRONTISPIECE). FIGURE 73. The largest shrub seen (/599/), with Dr. Carl Hubbs,
19 April 1957. FIGURE 74. The abundant seedlings (with a few of Mesembryanthemum crystallinum),
10 February 1957.

REID MORAN

FIGURE 75. Cistanthe guadalupensis on Outer Islet, leafless for the summer, 2] June 1968.

rocks pretty well around the island. On the main island itis occasionally seen on seacliffs,
but it is very rare in more accessible places such as the edge of the south-end mesa.

Dudley (1899) said nothing about the generic position of this plant, and until recently no
one has questioned his placing of it in Ta/inwm. From the dried specimen he called the
thickened stem a fusiform fleshy root, perhaps thinking it analagous to the tuberous stems
of various American species of Ta/inum, and called the leaves all radical. Wilson (1932)
and von Poellnitz (1934) could add little to the description and kept the species in Talinum.
The true nature of the stem became known only when the plant was rediscovered and
was illustrated and variously redescribed (Walther 1932, 1937; Moran and Lindsay 1950;
Lindsay 1951; Byles 1957). In fact, Hershkovitz (1992) still called the leaves all basal.

Although Eastwood (1929:404) formally listed this plant as Talinum guadalupense, she
also mentioned it (p. 396) as a species of Calandrinia. Walther (1937) commented on
the similarity of the flowers to those of “the well-known Calandrinia discolor Schrad.
from Chile,” and I have been impressed with its overall similarity to the Chilean plant
grown in California as C. grandiflora Lindl. Carolin decided its relationships were indeed
with the Chilean plants (Hershkovitz 1990). Surprisingly, the superficially very different
annual Calandrinia maritima, which happens also to grow on Guadalupe Island as well
as on the mainland, also belongs to Cistanthe section Cistanthe.

Carlquist (1965, fig. 8-13) showed a large plant and a flower of C. guadalupensis and
briefly discussed its unusual growth form. Carlquist (1974:399, fig. 10.15) showed the
plant and flower and, with a transection and a tangential section of the secondary xylem,
discussed parenchymatization of the xylem.

140

GUADALUPE ISLAND FLORA

Cistanthe guadalupensis throve in the garden of Dr. Carl Hubbs in La Jolla, California,
within sight of the sea, flowering handsomely most of the year and coming freely from
seed. It is widely admired among fanciers of succulent plants and still sometimes grown,
though they often consider it hard to grow (Rowley 1978, 1987).

Cistanthe maritima (Nutt.) Carolin in Hershkovitz, Phytologia 68:269. 1990.

Calandrinia maritima Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:197. 1838.

RANGE. — Coastal southern California to central Baja California; most islands of southern
California and NW Baja California, to Isla Cedros.

Brandegee (1900:22) was first to report this plant from Guadalupe Island but gave no
specific locality. It seems to be scarce and local on the east side of the island just below
the middle, as near the lobster camp. Others of Brandegee’s additions to the known flora
are rather local in this part of the island.

Claytonia perfoliata Willd. ssp. mexicana (Rydb.) J. M. Mill. & K. L. Chambers,
Novon 3:269. 1993.

Limnia guadalupensis Rydb., N. Amer. Fl. 21:311. 1932.
Claytonia perfoliata and Montia perfoliata of Guadalupe Island references.

RANGE. — Central California to Guatemala; islands of southern California to Cedros.

Palmer found this plant all over the island, in masses on the shaded side of rocks or logs
or in deep ravines (Watson 1876:113). Franceschi (1893c:134) found it quite common
from centre to north, most luxuriant under the palms, where it was in flower early in
December. It is now rather common among rocks and in other shaded spots, widespread
from shore to summit in the northern half of the island and to El Picacho, sometimes
lush and abundant.

Rydberg based Limnia guadalupensis on Palmer’s collection (No. 15) of 1875 (NY). From
the key and description it would appear to differ from related species in the shape of the
basal leaves and in the small size of the plant as a whole and especially of its stem leaves,
inflorescence, and seeds. However, the type sheet has not only three very small plants
but also one large one, clearly of the same collection. Howell (1942:148) remarked that
this plant was almost invariable on the island, and clearly there is only one species though
here as elsewhere the size varies greatly. Large plants (e.g. M6600) may be as much as
3.5 dm tall, with basal leaves to 4 cm wide, the disk of stem leaves 7.5 cm wide, and the
inflorescence 14 cm long. Greene (1885:221) remarked that the corolla of the island plant
was smaller and more purple than in California plants. Corollas are about 2—3 mm long,
which is a little small for the species, and I would say deep pink rather than purple. The
seeds are about 0.95—1.3 mm long, which again is rather small for the species.

Miller (1978) reported from chromosome counts of 235 populations that C/laytonia per-
foliata s.\. is a polyploid pillar complex based on three morphologically different and
geographically defined diploids and having five levels of ploidy.

Primulaceae

+*Anagallis arvensis L., Sp. Pl. p. 148. 1753.

RANGE. — Native to Europe; a widespread weed in North America.

141

REID MORAN

Palmer found only three plants, in a gravelly place near the beach on the east side (Watson
1876:117). Greene (1885:225) saw only one plant, on top of the island. No one has found
it since, and evidently it never became well established.

Dodecatheon clevelandii Greene ssp. insularis H. J. Thomps., Contr. Dudley Herb.
4:134. 1953.

Dodecatheon meadia of Guadalupe Island references.

RANGE. — Inner south coast ranges of California to San Miguel, Santa Rosa, Santa Cruz,
Anacapa, San Nicolas, Santa Catalina, San Clemente, and Guadalupe Islands.

Palmer found this plant very abundant on moist rocky slopes at the “south end” and
“middle” of the island and noted that goats were very fond of it (Watson 1876:117).
Franceschi (1893c:136) found robust large-leaved specimens, most abundant only between
the trail to the cabins and the cliff, the finest on the very ridge. He remarked that goat
hunters, short of tobacco and attracted by the leaves, used them as a substitute and said
they had a most pleasant flavor.

Between the goats and the goat hunters, the plant is now scarce, a few small colonies
seen scattered on the northeast slope of El Picacho at 500-800 m and one or two plants
on the cliff of the Lower Circus and on its brink. Most plants seen were sterile, only one
on a cliff having some old flowers 21 March 1974 (A/2//66). From Guadalupe Island,
Thompson (1953:133) cited Anthony 245 and Palmer 55.

Ranunculaceae
Myosurus minimus L. var. filiformis Greene, Bull. Calif. Acad. Sei. 1:277. 1885.
Myosurus minimus of Guadalupe Island references.

RANGE. — Northern California to NW Baja California.

Greene (1885:220) found this plant at the middle of the island and also at the north end,
near springs. Campbell (1952:394) cited Greene’s specimen of April 19, 1885, (GH) and
called Guadalupe Island the type locality. Apparently it has since been collected on the
island only once, in 1958 (Wiggins and Ernst 33), on the upper northwest slope of the
pine ridge at 600m. Thus it is rare in accessible places, but the foggy northwest cliffs
probably have suitable habitats where it may survive.

Ranunculus hebecarpus Hook. & Arn., Bot. Beechey Voy. p. 316. 1840.

RANGE. — Washington and Idaho to NW Baja California; Santa Catalina and Cedros
Islands.

Palmer found this plant abundant on warm slopes in the “middle” of the island (Watson
1876:112). Greene (1885:220) found it only in the shade of a large tree. It is now wide-
spread and locally common on the northwest slope at 500-800 m and more rarely seen
as far south as the spring.

Greene thought the island plant shorter and stouter than the California plant, the achenes
more numerous. However, Benson (1948:111) cited the collections of Palmer and Greene
as R. hebecarpus.

142

GUADALUPE ISLAND FLORA

Resedaceae

Oligomeris linifolia (Vahl) J. F. Macbr., Contr. Gray Herb. 53:13. 1918.
Oligomeris glaucescens and O. subulata of Guadalupe Island references.

RANGE. — Southern California to Texas and Mexico; islands of southern California to
Cedros, San Benitos, and Natividad; also Old World.

Palmer found this plant in deep warm canyons and ravines in the “middle” of the island
and occasionally at the “south end” (Watson 1876:113). It is occasional to locally common
at low elevations throughout the island but especially in the southern half, including Islote
Negro and Outer Islet. Some authors treat it as an introduction in the New World.

Rhamnaceae

+Ceanothus crassifolius Torr., Pacif. Railr. Rep. 4:75. 1857.

Ceanothus crassifolius Torr. var. planus Abrams, Bull. New York Bot. Gard. 6:415. 1910,
Ceanothus perplexans of Guadalupe Island references?

RANGE. — Southern California to NW Baja California.

Palmer found only three alive, at the base of Mt. Augusta (Watson 1876:114). Greene
(1885:222) saw only a small seedling, near the cabins. Franceschi (1893c:134) found one
plant alive near the center of the island west of Mt. Augusta among the cypresses, “sur-
rounded by what appeared to be the dead stumps of thousands of its brethren, which must
have formed a thick and general underwood not only in the larger cypress grove but also
in the smaller near the springs and cabins.” “Twelve to fifteen feet high.” Later he found
three or four more living plants in the upper grove. No one has found this shrub since,
and undoubtedly it is extinct on the island.

Under var. planus, McMinn (1942:234) cited Guadalupe Island, Dr. F. Franceschi 1332
in 1893.

+Ceanothus cuneatus (Hook.) Nutt. in Torr. & A. Gray, Fl. N. Amer. 1:267. 1838.
RANGE. — Oregon to NW Baja California.

Palmer found only three nearly dead specimens, among rocks in the “middle” of the
island (Watson 1876:114). No one has seen it since, and undoubtedly it is extinct on the
island.

Rhamuaus pirifolia Greene, Pittonia 3:15. 1896.
Rhamnus crocea Nutt. ssp. pirifolia (Greene) C. B. Wolf, Mon. Rancho Santa Ana Bot. Gard. 1:45
1938.

Rhamnus crocea of Guadalupe Island references.

RANGE. — Endemic to San Miguel, Santa Rosa, Santa Cruz, Santa Catalina, San Clemente,
and Guadalupe Islands.

Shrub to 2m high and 5 m wide or hanging from cliff to 4m; trunk to 1.5 dm thick.

Palmer found only six shrubs, growing in the crevices of high cliffs in the “middle” of

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the island, in bloom April 6 (Watson 1876:114). About that many have survived until
recent times, scattered on cliffs in the northeast part of the island: two in Oak Canyon
at 850 m (AM//2032) and 900 m (M/2036); Barracks Canyon at 420 m (M7829); north rim
of Barracks Canyon at 500 m (M/3803); Esparsa Canyon at 250 m (M5964). In May 1971
I saw what appeared to be this shrub on a high cliff at 675m near the middle of the
northwest end of the island. From Guadalupe Island Wolf (1938:46) cited Anthony 261,
Hanna (CAS), and Mason 1528.

Rosaceae

Aphanes occidentalis (Nutt.) Rydb., N. Amer. FI. 22:380. 1908.
Alchemilla occidentalis Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:432. 1840.

RANGE. — Washington to NW Baja California; Santa Rosa, Santa Cruz, Santa Catalina,
and San Clemente Islands.

Palmer found this plant among rocks and sage brush at the north end and also around a
spring (Watson 1876:114). This inconspicuous little plant is widespread on the northwest
slope of the island and the northeast ridge, from 500 to 1100 m, and fairly common
locally.

Robertson (1974) wrote that [subgenus] Aphanes “probably was confined to Europe before
western man began his explorations of the world.” However, there seems no question
that this is native in western North America and on Guadalupe Island.

+Heteromeles arbutifolia (Lindl.) M. Roem. var. macrocarpa (Munz) Munz, Aliso
4:92. 1958.

Photinia arbutifolia Lindl. var. macrocarpa Munz, Bull. S. Calif. Acad. Sei. 31:64. 1932.
RANGE. — Endemic to Guadalupe and San Clemente Islands?

The late Guadalupe Island population of one may be described as follows. Tree + 5m
tall, with one arching branch 7 m long; trunk 2 dm thick at base. Young stems and young
leaves densely tomentose with simple slender crisped white hairs + 1 mm long, a few of
which persist in age. Leaf blades elliptic, mostly obtuse and mucronate at apex but some

times longer than wide, the margins revolute, subentire or mostly weakly crenate-dentate,
with 10-18 crenations on each side, when young with subulate red glandular teeth + 0.4
mm long; petioles 12-27 mm long. Inflorescence to 16 cm wide. Petals white, spreading
to spreading-ascending. Young fruit obovoid-truncate, slightly angled, to 9.5 mm thick.

Wiggins (1980:793) reported the species as once common on Guadalupe Island but now
virtually exterminated by the goats, a few spindly shrubs persisting on inaccessible cliffs,
mostly at the north end of the island. However, I have seen no specimens or other records
to support this statement. | have seen only a single tree, arching out from a sheer arroyo
wall, at 70m, just northwest of the lobster camp on the southeast coast. Specimens of
December 1957 (M6452; Newcomb 184, UC) bear a few flowers and some immature
fruit, to 9.5 mm thick according to Newcomb’s notes, though I noted them only as 6 mm
long; specimens of April 1958 (Carlquist 484, Wiggins and Ernst 202) have immature
inflorescences. The tree had disappeared by 1970, and the species probably is now extinct
on the island. Dr. Wiggins wrote me in 1958 that Dr. Hanna said he had seen the same

144

GUADALUPE ISLAND FLORA

tree in 1922, but no specimen could be found at CAS.

times longer than wide; an exceptional leaf is 8.2 cm long by 5.7 cm and so less than 1.5
times longer than wide. In typical mainland specimens the leaf blades commonly are
narrower and acute and almost always are sharply serrate. Also, the young herbage in
the Guadalupe specimens is exceptionally pubescent. All together, these specimens thus
look quite different from typical mainland material: in fact, several able botanists (R.
Bacigalupi, L. Benson, A. Carter, J.T. Howell, P. A. Munz, R. F. Thorne), all well ac-
quainted with H. arbutifolia, failed to recognize sterile specimens.

Mainland collections of H. arbutifolia vary widely in leaf shape. The blades commonly
are acute, 1.5—3 cm wide, and 34 times as long, though specimens with broader or more
obtuse blades occur sporadically but rather widely through the range. The teeth vary in
size but are rarely so reduced as in the Guadalupe plant; and generally no correlation is
evident between width of leaf and size of teeth. Pubescence also varies, usually with no
evident correlation with leaf width or size of teeth; very few specimens are as tomentose
as those from Guadalupe Island.

However, some collections of H. arbutifolia from the Cape Region of southern Baja
California have leaves obtuse to rounded, to 6 cm wide (R. K. Peters 248, UC, from La
Laguna), some with the teeth small or subobsolete; at least some have young growth
tomentose; and at least some have large fruits. At La Laguna I collected from two shrubs
growing side by side (M/74344), both with young growth tomentose, one with leaves 2—3
cm wide, the other with leaves mostly 3—5.7 cm wide.

Munz (1932) proposed var. macrocarpa for trees of San Clemente and Santa Catalina
Islands, on the basis of fruits 8-10 mm long rather than 5-6 mm long. Raven (1963:324)
reserved judgement on the variety, unconvinced that the fruit was uniformly large in the
populations of these two islands; and he reported large fruit on some specimens from
Santa Cruz and Santa Rosa Islands. Although Thorne (1967:63) likewise did not recognize
the variety, he wrote that in general the specimens of Catalina Island did seem to have
larger fruits. These authors also referred to tree-like size.

Discussion of var. macrocarpa thus far had centered on size of fruits and had not mentioned
the leaves. However, after Dr. Munz had seen the Guadalupe specimens, he wrote
(1974:747), “There is a tendency for insular plants to have lvs. subentire, [and he went
on] the frs. red, 8-10 mm long, and these constitute a possible var. macrocarpa (Munz)
Munz. . . . Judging from their behavior in the botanic garden, they are not only more
showy, but less readily eaten by birds.” In the holotype, from San Clemente Island (Munz
6759, POM 18981), the leaves are obtuse and 2-3 times as long as wide; leaf serrations
on one branch are rather small and on the other subobsolete. A near topotype (Raven
17985, RSA, SD) is very similar in leaf shape and margins to the Guadalupe specimens.
In most San Clemente specimens the leaves are rather wide for their length, some 5 cm
wide, but a few are strongly serrate. Most are like average mainland specimens in pu-
bescence, but a few are markedly pubescent.

Trask (1899) remarked that on Catalina Island the leaves of H. arbutifolia were usually
entire, sharply serrate ones occurring but rarely, often not one such leaf on a slope covered
with this species. In fact, this seems an overstatement; for many collections, including
some with large fruit, have sharply serrate (and rather narrow) leaves, similar to average
mainland material. Although some have broad leaves and reduced or subobsolete teeth,
the tendencies seem weaker here than on San Clemente Island. Among specimens from

145

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Santa Cruz and Santa Rosa Islands, where Raven noted some large fruit, some have
narrow and sharply serrate leaves; but again there is some tendency to reduction of teeth
and a greater tendency to greater leaf width. One Ralph Hoffmann collection (SBBG)
from Santa Rosa Island has one leaf blade 15 cm long and 7.9 cm wide! On Isla Cedros
most plants, from canyons on the east side, have the leaves narrow, acute, and sharply
serrate; but one collection (4/4/8432), from under the pines 3 km from the north end, has
some leaves to 4cm wide and obtuse, some with much reduced teeth. Since many of
these insular collections lack fruit, it is not clear what proportion of the plants have large
fruit or to what extent leaf characters are correlated with large fruit. One specimen from
Sierra de San Francisquito in the Cape Region (7. S. Brandegee 205, UC) Tatemi Shimizu
annotated as var. macrocarpa.

Although the known population of Guadalupe Island may be called remarkably uniform,
nothing is known of former variation. Perhaps the Guadalupe plant is a relict that persisted
in a relatively pure state on this most isolated island, persists with some admixture of the
mainland form on less isolated San Clemente Island, and has more nearly been swamped
out on the continental Santa Catalina, Santa Cruz, Santa Rosa, and Anacapa Islands. Dr.
Daniel Axelrod wrote me that he knew of no fossils of this form. It seems well treated
as a subspecies (or variety) on Guadalupe and San Clemente Islands, even though it
becomes more of a taxonomic embarrassment on the northern islands and in the Cape
Region.

Rubiaceae
Galium angulosum A. Gray, Proc. Amer. Acad. Arts Sci. 11:74. 1876.

RANGE. — Endemic to Guadalupe Island.

Dense, much branched, probably dioecious, dark green shrub to 8 dm wide and projecting
nearly as far from cliff, with long slender woody branches spreading to pendent, bearing
dense annual tufts of new vegetation at nodes; stems 4-8 ribbed, the stems and leaves
sparsely hispid with short or long straight spreading hairs. Leaves 4 or often 6 in a whorl,
mostly 3-7 mm long, |-nerved, spreading or often reflexed, elliptical or obovate, narrowed
gradually to base, obtuse or round at apex, tipped with long weak hair; glandular cells
apparently none. Flowers in April and May, apparently few, solitary at ends of lateral
branches; corolla rotate, yellow-green, glabrous or rarely hispid; ovaries glabrous or rarely
with a few hairs like those on stems and leaves. Fruit in October, fleshy, glabrous, whitish,
translucent, 2.5-3.5 mm thick, the mericarps not separating, becoming shiny black and
wrinkled when dry. Chromosomes: 2” = ca. 66. (Description based on that of Dempster
1978:10)

Palmer found only a single small scrubby plant, in a crevice of a high cliff in the “middle”
of the island (Watson 1876:115). Though not reported since, it has persisted until recently
as a very rare plant on cliffs in the northeast part of the island. In 1957 I found two
plants on a cliff at 450m on the north rim of Barracks Canyon at the mouth (4/6/17)
but in 1960 failed to find them again. Also in 1957 | found a single plant at 250 m on
a cliff on the south side of Esparsa Canyon (7/6/36), and it was still there in 1970 on
my last visit to that spot.

Dempster (1978:10-11) thought this plant probably closest to the fleshy fruited dioecious
species of California. In buds collected from the Esparsa Canyon plant in May 1967
(M13802), she found 2n = + 66; she was rather sure that it was hexaploid. She noted that

146

GUADALUPE ISLAND FLORA

all flowers seen had well-developed pistils and abortive stamens. She added that most
material had 4 leaves at the great majority of nodes, though the type had 6 at most nodes.
And the type was unique also among the six collections she had seen in having hispid
corollas.

*Galium aparine L., Sp. Pl. p. 108. 1753.

RANGE. — Widespread in North America, commonly thought to be introduced from
Europe; islands of California.

Palmer found this plant common on warm shady hillsides in the “middle” and more rarely
at the “south end” (Watson 1876:115). It is widely distributed in the northern half of the
island, as far south as Juniper Canyon, and is fairly common about rocks in the canyons,
at the foot of cliffs, and on the northwest slope of the island.

Rutaceae

*Ruta chalepensis L., Mant. p. 69. 1767.
Ruta graveolens of Guadalupe Island references.
RANGE. — Native of southern Europe; widely grown and often escaping.

In 1948 three sterile plants grew in the canyon mouth at Northeast Anchorage (Moran
1951:160). In 1965 a colony of a dozen grew on the south bank of the arroyo at 20m
(M12019), so the plant is reproducing but not spreading rapidly.

Saxifragaceae

Jepsonia malvifolia (Greene) Small, Bull. Torrey Bot. Club 23:19. 1896.
Heuchera ? and Jepsonia parryi of Guadalupe Island references.

RANGE.— Endemic to Santa Rosa (type locality), Santa Cruz, San Nicolas, Santa
Catalina, San Clemente, and Guadalupe Islands.

Woody caudex 5—7 cm high, branching horizontally to form a clump to 15 cm long and
10 cm wide, the branches mostly 1.5—2 cm thick, abruptly narrowed upward to short stem
+5 mm thick bearing leaves and peduncles. Leaves 1—2, appearing in October, fragrant,
the petiole at first pink, 3-7 cm long, 1.5—3 mm thick, the blade at first funnelform and
to 6cm wide. Peduncles I—3 from one stem, at different stages, 14 cm tall, 2 mm thick
at base, densely glandular-pubescent, with bract + 5 cm above base; cyme 2—3 cm wide,
of +20 flowers on several lax cincinni, or with subsidiary cymes below. Flowers August
to October, on pedicels 1-2 mm long. Hypanthium subcampanulate, subpentagonal, sub-
truncate at base, glandular-puberulent, the tube + 2 mm long, greenish with 10 dark red
veins, the calyx lobes erect, triangular-ovate, rounded, I—1.2 mm long by 1.2—1.5 mm.
Corolla 7-8 mm wide, the petals outcurved, ovate, rounded, 3—3.5 mm long, + 2.5 mm
wide, white veined dorsally with pink becoming purplish red, the stipelike base + 0.4 mm
wide. Filaments white, erect, 2—3 mm long, the anthers purplish. Gynoecium whitish,
2mm high, the styles short, indefinite. (Fig. 76)

Palmer found a single plant, not in flower, in the crevice of a rock (Watson 1876:114);
Franceschi (1893c:135) likewise found but one; and Greene (1885) omitted it from his

147

REID MORAN

FIGURE 76. The insular endemic Jepsonia malvifolia (M12064), from lower east slope of El
Picacho at 350 m; flowering in San Diego | October 1972. It grows also on five of the Channel
Islands

list, in which he intended to include Palmer’s findings as well as his own. Moran
(1951:160) found sterile plants on a cliff at the north end, which flowered in cultivation.
Despite early reports or lack of them, the plant is widespread and sometimes locally
common, especially on north cliffs and in deep soil among rocks on north slopes, from
the north end to the gray peak south of El Picacho and to the east slope above the
southeast sealing camp, at least from 100 to 800 m.

Scrophulariaceae
Note. — For Galvezia juncea see Excluded Species, p. 169.
Antirrhinum nuttallianum Benth. ssp. subsessile (A. Gray) D. M. Thomps., Syst. Bot.
Monogr. 22:95. 1988.

Antirrhinum subsessile A, Gray, Bot. Gaz. 9:54. 1884

148

GUADALUPE ISLAND FLORA

Antirrhinum pusillum Brandegee, Univ. Calif. Publ. Bot. 6:360. 1916.

Antirrhinum nuttallianum Benth. forma pusillum (Brandegee) Munz, Proc. Calif. Acad. Sci. Ser. 4,
1153359. 1926:

Sairocarpus pusillus (Brandegee) D. A. Sutton, A revision of the tribe Antirrhineae, p. 466. 1988.

RANGE. — Coastal southern California to south central Baja California; San Miguel, Santa
Rosa, Santa Cruz, Anacapa, Santa Catalina, Los Coronados, Todos Santos, San Martin,
and Cedros Islands; Arizona.

Palmer found this plant rather rare in deep warm canyons in the “middle” of the island
(Watson 1876:117). It is widespread on the island but rather uncommon, often no more
than one or two seen at a time. The small cleistogamous forma pusillum likewise is
widespread but not common, as often as not the two occurring together. Thompson
(1988:98) found experimentally that the cleistogamous condition of forma pusi/lum may
be induced by suboptimal growing conditions and is easily reversible.

Antirrhinum watsonii Vasey & Rose, Proc. U.S. Natl. Mus. 11:533. 1889.

Antirrhinum kingii S. Watson var. watsonii (Vasey & Rose) Munz, Proc. Calif. Acad. Sci. Ser. 4,
15:367. 1926.

Sairocarpus watsonii (Vasey & Rose) D. A. Sutton, A revision of the tribe Antirrhineae, p. 471.
1988.

RANGE. — Central Baja California to western Sonora and southernmost Arizona; Islas
Cedros and San Benitos.

This plant was first found on the island in 1957 (M5632, M5987, M6157). It is widespread
but not common at low elevations in the southern part, from the lobster camp and West
Anchorage southward and on Outer Islet. Thompson (1988:104) reinstated it as a species,
citing M5987 and M17345 from Guadalupe Island. Dr. Thompson in 1986 annotated six
of my Guadalupe collections.

Castilleja attenuata (A. Gray) T. 1. Chuang & Heckard, Syst. Bot. 16:656. 1991.

Orthocarpus attenuatus A. Gray, Pacific Railr. Rept. 4:121. 1857.

RANGE. — Vancouver Island to NW Baja California; Santa Cruz Island; also central Chile,
where possibly introduced.

This plant has been found only one place on the island, first in 1967: locally common in
seep on northwest slope between the old lookout and Hemizonia Cliff, at 800 m (M//3827/).
I have not found it there in drier years.

+Castilleja exserta (A. Heller) T. 1. Chuang & Heckard, Syst. Bot. 16:657. 1991.

Orthocarpus purpurascens Benth., Scroph. Indicae. p. 13. 1835. Not Castilleja purpurascens
Greenm.

RANGE. — California to Arizona, Sonora, and northern Baja California; San Miguel, Santa
Rosa, Santa Cruz, and Santa Catalina Islands.

Brandegee (1900:22) is the only one to have found this plant on Guadalupe Island, and
Keck (1927:541) cited his specimen as Orthocarpus purpurascens typica. Because it 1s a
west American plant that also grows on the four northern islands in a familiar pattern, it
might be easy to accept as native on Guadalupe. Without explanation, however, Brandegee

149

REID MORAN

called it “introduced” — though his assertion is unconvincing because he said the same
of Lasthenia californica, which seems clearly to be native. The fact that the plant has
not been found before or since suggests that it may indeed be a waif that never became
established. Since there is no way of knowing, I list it as possibly introduced on the island
(Table 2).

Castilleja fruticosa Moran, Trans. San Diego Soc. Nat. Hist. 15:283. 1969.
Castilleja guadalupensis of some references.
RANGE. — Endemic to Guadalupe Island, at the south end and on Outer Islet.

This is a woody perennial 2—S or rarely 9dm high and to 5dm or more wide, with
elandular-puberulent to somewhat villous light green herbage, the bracts and calyx tipped
with Indian orange.

This plant grows on a barren volcanic mesa and in two craters near the south end of
Guadalupe Island, in an area of low rainfall but of frequent fog. In 1948 | found it on
the inner slope of the crater of Outer Islet (Moran 1951:159) but failed to find it on the
main island, even though I collected at the very spot on the mesa where its bright flowers
were conspicuous in later years. Earlier collectors at the south end also had failed to find
it. Until the founding of the weather station in 1946, enough goats evidently came to the
south end to keep certain plants near the point of extinction; but goats have since come
more at their peril, and the plants have gained some slight protection. In 1957 I did find
C. fruticosa on the mesa above the station but saw only herbaceous plants, looking almost
like annuals. In 1967 and later, some plants were small shrubs with woody stems to 2
em thick. In 1957 it extended as far north as the inner slope of the crater of Plaza de
Toros, at 320m (M6479), though in 1970 I failed to find it there. These facts suggest
that in 1957 C. fruticosa might recently have colonized the main island from one of the
nearby islets and that the extent of the colony has since varied with the years depending
on goat damage. The number of goats coming to the south end, and the damage they do,
must vary from year to year, depending on pressures farther north.

Castilleja fruticosa, though still herbaceous at first flowering, is remarkable for the thick
and hard woody stems of older plants. Apparently it is one of the woodiest of the genus.
Perhaps in relation to the maritime habitat, the foliage leaves are quite thick and somewhat
succulent. The capsules are unusually hard and woody and are distinguished by a depres-
sion on the under side, which sometimes becomes a deep transverse groove. Capsules
from past years often persist on dead inflorescence axes, mostly still unopened and still
holding seeds. Such seeds proved viable, and from seedlings (M//3768) Larry Heckard
got a chromosome count of 7 = 12 (Chuang and Heckard 1993).

Castilleja guadalupensis Brandegee, Zoe 5:166. 1903.
Castilleja foliolosa of Guadalupe Island references.
RANGE. — Endemic to Guadalupe Island.

Intricately and divaricately branched shrub 2-6 dm high, the stems hard and woody, closely
cottony tomentose above with very fine dendritic hairs, subglabrescent; internodes to
lcm; axils mostly with shoots, at least a cluster of leaves and often the stem 3—5 cm
long. Leaves spatulate, rounded at apex, mostly 4-6 cm long and 3-10 mm wide above,
+ 1 mm wide at base, not decurrent but clearly jointed to stem, thin, at first closely tomen-

150

GUADALUPE ISLAND FLORA

tose with dendritic hairs, in age subglabrescent, those below and sometimes well above
middle simple and entire, the upper ones lobed at or a little below middle, the lobes
spatulate, 1.5—2.5 cm long; higher leaves with lobes more towards base. Bracts

+ 2.5 cm long, with pair of ascending lobes near base, the upper with another pair near
apex, the lobes spatulate, the middle one 3—5 mm wide, the lateral + 2 mm wide above.
Young spike crowded, 3-6 cm long, with 10-15 or more flowers and buds, the lower
internodes after anthesis becoming as much as 13 mm long. Flowers strictly sessile. Calyx
tomentose, 21—22 mm long, cleft 4-5 mm before and 9-10 mm behind, the double lateral
lobes 3-4 mm wide, retuse to a depth of 0.5—1 mm, the tips rounded. Corolla 19-21 mm
long, densely puberulent along the back with multicellular but unbranched non-glandular
trichomes 0.1—0.2 mm long, glabrous on the thinner margins, the tip or at least part of
the back protruding through posterior sinus of calyx; lip short, the teeth half its length.
Stamens shorter than to slightly exceeding corolla, the filaments glabrous, the anthers
slender, curved, glabrous, + 3 mm long. Style exserted 3—5.5 mm, glabrous, thickened and
hooked at apex, the stigma papillose. Very immature capsules 7 mm long and 5 mm wide.

Palmer found this plant only in the “middle” of the island, rare among fallen branches
(Watson 1876:117). Anthony collected the type specimen (UC 103541) in “an almost
inaccessible spot on the western cliff,’ September 20, 1896. On that date, according to
Gaylord (1897:42), Anthony went to the northern limit of the pine, doubtless far out
towards North Point. A slip in the pocket reads “only one plant of this species could be
reached and that with great difficulty.” Drent collected it again in 1898 (UC), and no one
has found it since. It may possibly survive on the vast inaccessible northwest cliff; but
from Palmer’s statement, it probably was not mainly a cliffdweller, so it is very likely
extinct.

Galvezia speciosa (Nutt.) A. Gray, Proc. Amer. Acad. Arts Sci. 22:31]. 1887.

Gambelia speciosa Nutt., Proc. Acad. Nat. Sci. Philadelphia 4(1):7. 1848.
Antirrhinum speciosum (Nutt.) A. Gray, Proc. Amer. Acad. Arts Sci. 7:376. 1868.

RANGE. — Endemic to Santa Barbara?, Santa Catalina, San Clemente, and Guadalupe
Islands. (Not known from the Coronado Islands or the Baja California peninsula, where
cited by Wiggins [1980:495])

Shrubs prostrate, or in level protected places to 1.5 m high, or hanging to 3 m from cliffs.
Flowers February to August.

Palmer found this shrub frequent in the crevices of high rocks in the “middle” of the
island and called it very ornamental, the bright scarlet flowers continuing all summer
(Watson 1876:117). It is one of the commoner plants of seacliffs in the northeast part of
the island, as well as on shaded cliffs in Oak, Barracks, and Esparsa Canyons. On Islote
Negro and Outer Islet it is not confined to cliffs; and probably the same is true of Inner
Islet, though there only cliffdwellers can be seen from below.

According to Sutton (1988:511), Guadalupe specimens differ from Californian in the rather
broader leaves, with glabrous (not glandular) axils, and in the seeds with less anastomosed
winglike ridges on the ventral face. He thought this material might deserve taxonomic
recognition.

Gray (1887) combined the North American Gambelia Nuttall with the South American
Galvezia Dombey. Sutton (1988) separated Gambelia again on the basis of the thicker-
walled capsules with locules opening by several teeth and the bilaterally symmetrical

151

REID MORAN
seeds. However, David Thompson (1993:1032) kept it in Galvezia.

Linaria canadensis (L.) Dum.-Cours., Bot. cult. ed. 1, 2:96. 1802.

Linaria canadensis (L.) Dum.-Cours. var. texensis (Scheele) Pennell, Proc. Acad. Nat. Sci. Phila-
delphia 73:502. 1921.

RANGE. — British Columbia to SE USA and southern Baja California and Socorro Island:
San Miguel, Santa Rosa, Santa Cruz, Anacapa, Santa Catalina, San Clemente, Los Coro-
nados, Todos Santos, and Cedros Islands.

Palmer found this plant rare on the sides of canyons in the “middle” of the island (Watson
1876:117), and it has seldom been collected since. It grows on grassy slopes of the north
ridge at 600-800 m (M/25394) and in the valley below the Lower Circus at 800m
(M25389), in some years very scarce but in wet years locally common.

Mimulus latifolius A. Gray, Proc. Amer. Acad. Arts Sci. 11:95. 1876.

Eunanus latifolius (A. Gray) Greene, Bull. Calif. Acad. Sci. 1:99. 1885.
Mimulus brandegei Pennell, Proc. Acad. Nat. Sci. Philadelphia 99:170. 1947.

RANGE. — Endemic to Guadalupe and Santa Cruz Islands.

Corolla limb pink or lavender-pink; tube dark purplish red; palate ridges yellow with
purple spots. Capsule light brown, asymmetric, 16 mm long, flattened contrary to partition,
sharply bent 4 mm above base and thence straight; outer cell wider, splitting to bend, the
inner apparently not dehiscent. Seeds light brown, smooth and somewhat shining, ovoid,
apiculate at both ends, 0.5—0.6 mm long, 0.3—0.4 mm thick.

Palmer found this plant only in the “middle” of the island, scattered in warm rather moist
spots (Watson 1876:117). Greene (1885) and Vasey and Rose (1890) listed it without
comment. In April 1958 Ernst and I found five plants scattered along the bed of Arroyo
Melpomene at +450 m (Moran and Ernst 6738) (Plate 2K), but I have not seen it there
since. In April 1970 it was locally common (M/7392) with Polypodium californicum and
Jepsonia malvifolia at 800 m on the west-facing gray cliff above West Anchorage and
1 km south of El Picacho. This population, to the windward of upper Arroyo Melpomene,
is a likely seed source for the plants found in that arroyo in 1958 and is the only known
refuge for the species on the island. On 22 May 1971 I went back, hoping to find seeds,
but the few plants seen were less advanced than those of April the year before. This had
been a very dry year but with a little rain about two weeks before my visit; possibly the
plants had germinated after the rain. The blowing fog of that day perhaps is common,
somewhat moistening the upper west slope. It is noteworthy that the polypodium also
grows here at its southernmost station on the island and the only place I remember it on
any but a northfacing cliff.

Mimulus latifolius belongs to section Oenoe A. Gray, including ten Californian species,
with capsules woody, asymmetric, tardily or not dehiscent. The type of M. Jatifolius is
Palmer 58 (GH; isotypes MO, NY, PH). From Guadalupe Island, Grant (1924:311) further
cited only collections of Bryant in 1885 and Palmer (839) of 1889. She also cited a
collection of Brandegee in 1888 from Santa Cruz Island: Pennell in 1947 separated that
plant as M. brandegei, but Thompson (1993) included it again in M. Jatifolius. It is now
very rare if not extinct on Santa Cruz Island, where it was last collected by Carl Wolf
in 1932 (Junak et al. 1995:264). Another member of Oenoe, M. traskiae A. L. Grant,

GUADALUPE ISLAND FLORA

grew on Santa Catalina Island but apparently has not been found since 1903 and possibly
is extinct.

Scrophularia villosa Pennell, Field Mus. Bot. Ser. 5:223. 1923.
RANGE. — Endemic to Santa Catalina, San Clemente, and Guadalupe Islands.

Shrub I1—1.5m high, the woody stem to 8 dm high, to 3 cm thick at base. Flowers in
April and May; corolla deep maroon shading to almost black above.

On Guadalupe Island this rare shrub first came to light in 1957, surviving on a few
north-facing cliffs in the north and northeast part of the island. All together, I have found
only six: four in 1957 at 250m in Esparsa Canyon (M5970); one in 1958 at 120m in
South Esparsa Canyon (6596); and one in 1971 at 675 m near the middle of the north
slope of the island (M/8390). I visited the site in Esparsa Canyon in four later years, and
by 1970 all four shrubs apparently were gone. Let us pray.

From the other eight species he recognized for western North America, Shaw (1962)
separated S. villosa by its shrubby habit, glandular villous inflorescence, and reduced or
absent sterile filament. He cited M5970 from Guadalupe Island.

Solanaceae

Lycium californicum Nutt. in A. Gray, Bot. Calif. 1:542. 1876.

RANGE. — Southern California to Baja California, Arizona, and Sonora; San Miguel, Santa
Cruz, Anacapa, San Nicolas, Santa Barbara, Santa Catalina, San Clemente, Los Coronados,
Todos Santos, San Martin, San Benito, and Natividad Islands.

Palmer found this shrub on rocky bluffs at the “extreme south end” but not abundant
(Watson 1876:117); his specimen was cited by Hitchcock (1932:329). Howell (1942:152)
reported it as forming thickets that are one of the characteristic features of the south end,
the stems curving to the ground, rooting, and growing up again. Moran (1951:159) found
it common on the floor of the crater on Outer Islet (Fig. 34). It is widespread in the
southern half of the island, from Long Canyon and West Anchorage to Islote Negro and
the south end, especially in the arroyos, from sea level at least to 500 m.

Howell remarked that the flowering parts of his collection were smaller than those de-
scribed by Hitchcock.
Lycium fremontii A. Gray, Proc. Amer. Acad. Arts Sci. 6:46. 1862.

RANGE. — Southern California to Arizona, southern Baja California, and Sonora; Santa
Rosa Island.

This shrub was first found on the island in 1958. So far as known it is very local, on the
rocky walls of Arroyo Melpomene at 400 m (Moran and Ernst 6733, M7848).

+Nicotiana attenuata Torr. in S. Watson, Bot. King Exp. p. 276. 1871.

Nicotiana petuniaeflora Greene, Bull. Calif. Acad. Sci, 1:209. 1885.
Nicotiana bigelovii of Guadalupe Island references.

RANGE. — British Columbia and Montana to Texas and NW Baja California.

153

REID MORAN

“Two or three feet high, stout, viscid-pubescent and somewhat hispid-scabrous: radical
leaves oblong-lanceolate, 3—4 inches long on slender petioles; cauline linear-lanceolate,
longer than the radical, on shorter petioles; calyx-teeth triangular-lanceolate; corolla an
inch and a quarter long, salverform, white changing to bronze-purple; limb three-fourths
of an inch broad, with very shallow, scarcely noticeable, rounded or even retuse lobes.”
(Greene |,c.)

Palmer found this plant (no. 64) in only a few places in the “centre” of the island, in
open spots and good soil; flowers greenish yellow, bronzy below (Watson 1876:117).
Greene (1885:226) added nothing on the distribution. The last collection was in 1898, by
Drent (UC), and the plant undoubtedly 1s extinct on the island.

Goodspeed (1954:491) listed N. petuniaeflora Greene among the insufficiently known
species, as “NV. attenuata Torr.?”; but in treating N. attenuata (p. 429), he did not even
mention it. Under N. attenuata, however, without qualification, he cited Palmer 64 from
Guadalupe Island; and he wrote (p. 47) that the representatives of the species on the
island were identical with those on the mainland and thus gave no indication of a long
period of isolation. Yet presumably Greene’s type collection is of the same species.

*Nicotiana glauca Graham, Edinburgh Phil. Jour. 1828:174. 1828.
RANGE. — Native to South America; naturalized from California to Texas and Mexico.

This noxious weedy shrub first reached the island between 1932 and 1948 (Moran
1951:160): by the time of my first visit it was well established in the mouth of the canyon
at Northeast Anchorage, where the sharp-eyed Tom Howell had failed to find it 16 years
before. It forms thickets there and has spread north and south along the shore and up the
slope. Unfortunately, the goats shun it, and it threatens to take the island. Though botanists
see the invasion as a disaster, ornithologists Howell and Cade (1954) hailed it as cause
for hope, because this large shrub provides extensive cat-resistant habitat for small birds,
with nectar for hummingbirds and seeds for passerines.

*Solanum americanum Mill., Gard. Dict., ed. 8. 1768.

Solanum nodiflorum Jacq., Ic. Pl. Rar. 2:288. 1786-93.
Solanum calvum Bitter, Repert. Sp. Nov. 12:81. 1913.
Solanum nigrum of Guadalupe Island references

RANGE. — Apparently native to tropical America; a widespread weed in North America
and elsewhere.

Palmer found this plant rare in the “middle” of the island and in a canyon near the beach
on the east side, in rich level spots (Watson 1876:117); and on his second trip he found
a single plant in a canyon at the south end (Vasey and Rose 1890:26). Franceschi
(1893c:137) found it more common, a few in crevices of the Lower Circus and more in
the canyon near the landing. I have not seen it in the northern half of the island, but it
grows sparingly farther south, as at the lobster camp and near the mouth of Arroyo
Melpomene.

Bitter based S. calvum on Palmer 60 pro parte (UPS). He described the specimen in great
detail but did not compare it with any other species. It appears not to differ from S.
americanum. Dr. G.L. Stebbins in 1958 verified my identification of one collection
(M6448) with S. nodiflorum.

GUADALUPE ISLAND FLORA
Solanum douglasii Dunal in A. DC., Prod. 131:48. 1852.

Solanum nigrum L. var. douglasii (Dunal) A. Gray, Syn. Fl. N. Amer. ed. 2, 21:228. 1886.
Solanum profundeincisum Bitter, Repert. Sp. Nov. 12:80. 1913.

RANGE. — Central California to Southern Baja California and central Mexico; San Miguel,
Santa Rosa, Santa Cruz, Anacapa, San Nicolas, Santa Barbara, Santa Catalina, and San
Clemente Islands.

Palmer found only two plants, in a canyon near the beach on the east side (Watson
1876:117). Greene (1885:226) also saw two plants in the canyon near the beach, “perhaps
the very same individuals seen by Dr. Palmer.” Franceschi (1893c¢:137) found three plants,
two on the lava dike at the southern side of the landing and one a little way up the
canyon. There are no other reports. The plant is now rare on the island, seen only in the
northeast part. In 1957 two grew on a south-facing cliff at 450m on the north rim of
Barracks Canyon at the mouth (/6//6), but by 1960 one of these was gone. In 1958
several grew on a south-facing talus slope at 200m in South Esparsa Canyon (6602).

Bitter based S. profundeincisum on Palmer 60 pro parte and 6/ (UPS). Again, although
he described the specimens in great detail, he did not compare his proposed species with
S. douglasii or with any other species. I do not see that it differs at all.

Solanum wallacei (A. Gray) Parish ssp. clokeyi (Munz) Thorne, Aliso 7:76. 1979.

Solanum arborescens Clokey, Bull. S. Calif. Acad. Sci. 30:60. 1931. Not Moench.
Solanum clokeyi Munz, Bull. S. Calif. Acad. Sci. 31:69. 1932.

Solanuim wallacei var. clokeyi (Munz) McMinn, Ill. Man. Calif. Shrubs p. 491. 1939.
Solanum wallacei, S. xanti, and S. xanti var. wallacei of Guadalupe references.

RANGE. — Endemic to Santa Rosa, Santa Cruz, and Guadalupe Islands.

Weak shrub to 16 dm high, the stems brittle, | cm thick, to 5 cm at common base. Herbage
viscid, densely soft white pubescent, at first very densely so and yellowish, the trichomes
spreading, mostly unbranched, of unequal lengths intermixed, 0.2—1 (—2) mm long, unis-
eriate, of 2—15 cells tipped with a tiny yellowish globule. Leaf blades ovate to oblong-
ovate, acute or subacute, cordate to subtruncate at base, entire or faintly and irregularly
crenate, 3—6 cm long, 15—37 mm wide; petioles 6-12 mm long. Cymes on peduncles 10—23
mm long, of mostly 2 cincinni of 7-9 flowers each, the axes at length to 23 mm, the
pedicels slender, pubescent, 10-21 mm long. Flowers Jan—May, at times abundant and
showy. Calyx pubescent, in flower campanulate, 3—4 mm long, in fruit shallowly bowl-
shaped, 12 mm wide, the lobes deltoid-ovate, acute or becoming rounded, |.5—2 mm long.
Corolla rotate, shallowly 5-lobed, 2—3 cm wide, blue-violet, pubescent outside on parts
exposed in bud, with paired green spots surrounded by white inside at base opposite each
lobe, the lobes broadly ovate, acute, 7-9 mm long and wide. Filaments adnate 2 mm to
corolla base, equal, 4 mm long and | mm wide at base, puberulent; anthers equal, yellow,
3.54 mm long, 1.5 mm wide, splitting lengthwise. Ovary globular, green, 1.5—2 mm thick;
style 8-10 mm long, papillose in lower third and with a few scattered short trichomes,
glabrous above. Old fruit | cm thick, orange-brown, glabrous and somewhat shiny. Seeds
red-brown, obliquely ovate, 1.92.1 mm long, 1.4-1.6 mm wide, 0.6—0.8 mm thick, the
margins rounded, the surface closely pitted with round-bottomed pits 0.02—0.1 mm wide.
Chromosomes: n = 12 fide Marion Cave.

Palmer found this plant only in the “middle” of the island, in the crevices of rocks (Watson
1876:117). He called it a very showy shrub, in large bunches about two feet high, blooming

]

REID MORAN

all year; flowers numerous, lilac or purple. Greene (1885:226) found it common on the
plateau, in round compact masses three feet or more in height and thickness. Franceschi
(1893c:137) found it on the eastern cliff a little south of the corral, already in flower at
the beginning of January. | have found it (4/7837 etc.) only on the cliff of the Lower
Circus, the place meant by Franceschi and perhaps also by Palmer. Here I saw mainly
one colony of about ten plants, very showy and conspicuous in flower, surviving fortui-
tously high on the cliff in a pocket of rich soil like a window box, suitable for continued
reproduction yet beyond the reach of goats. | saw only one old fruit, from the year before.
In 1988 I saw another plant out of reach farther north on the cliffs.

Palmer described the fruit as small, changing from green to yellow, mottled, and at length
very dark plum color, maturing very slowly. Both Katherine Brandegee (in Franceschi
1893c:137) and Greene identified the Guadalupe Island plants with S. xanti var. wallacei
of Santa Catalina Island, though Gray (Watson 1876:117) had referred Palmer’s collection
to S. xanti and not to the variety. The ssp. clokeyi is a smaller shrub than ssp. wa/llacei,
with shorter, less dense, and less tawny pubescence, smaller leaves, smaller flowers, and
smaller fruit. It needs closer comparison with the variable S. xanti of the mainland.

Umbelliferae

Daucus pusillus Michx, Fl. Bor. Amer. 1:164. 1803.

RANGE. — Vancouver Island to central Baja California, SE USA, and South America;
islands of southern California and Los Coronados and Todos Santos.

Palmer found this plant abundant through the “middle” of the island, among sage brush
on the sides of canyons and in open level places (Watson 1876:115). Greene 1885:223)
noted it without comment. It now seems to be rare, known from only one collection this
century: scarce on grassy slope at 900 m south of the cypress grove (16676).

Lomatium insulare (Eastw.) Munz, Man. S. Calif. Bot. 358. 1935.
RANGE. — Endemic to San Nicolas, San Clemente, and Guadalupe Islands.

Moran (1951:159) found this plant on the inner slope of the crater of Outer Islet and also
near the southwest corner of the south-end mesa. On Outer Islet it grows not only in the
crater but also in the upper valley at 190 m, where one plant measured 7 dm wide. On
the mesa it seemed at first very local in a shallow arroyo at 100 m, just above Campo
Sur (Fig. 77), and so far as I have seen has since spread less than | km to the north.
Plants seen on the mesa in June and August were completely dormant and were dead
above ground.

This plant also grows at 500 to 800m on foggy northwest cliffs from North Point to
Hemizonia Cliff, flowering December to May (M6437, M13813, M17298). Leaf segments
are wider here than in plants at the south end.

Urticaceae

+Hesperocnide tenella Torr., Pacif. Railr. Rept. 4:139. 1857.

RANGE. — Central California to NW Baja California; Santa Cruz, Santa Barbara, Santa
Catalina, San Clemente, Los Coronados, Todos Santos, and San Martin Islands.

GUADALUPE ISLAND FLORA

FIGURE 77. The insular endemic Lomatium insulare (M12069) on the south-end mesa, 5 March
1965. It grows also on Outer Islet and on cliffs at the north end, and on San Clemente and San
Nicolas Islands.

Palmer found this plant in damp shady places, among high rocks, in the “middle” of the
island (Watson 1876:119), Greene (1885:227) listed it without comment, and Franceschi
(1893c:137) found it very common everywhere. Apparently no one has seen it since, and
evidently it is extinct on the island.

Parietaria hespera B. D. Hinton var. hespera.

Parietaria hespera B.D. Hinton, Sida 3:293. 1969.
Parietaria debilis and P. floridana of Guadalupe Island references.

RANGE. — Central California to Utah, New Mexico, and NW Mexico; islands of southern
California and south to Cedros,

Palmer found this plant abundant in damp shady places (Watson 1876:119). It is rather
common among rocks from shore to summit and from the north slope to the south end
mesa and on Islote Negro and Outer Islet.

Viscaceae
+Phoradendron densum Trel., Gen. Phorad. p. 27. 1916.
Phoradendron guadalupense Trel., Gen. Phorad. p. 29. 1916.

Phoradendron bolleanum (Seem.) Eichler ssp. densum (Trel.) Wiens, Brittonia 16:29. 1964.
Phoradendron bolleanum and var. pauciflorum of Guadalupe Island references.

REID MORAN
RANGE. — Oregon to Baja California and Chihuahua.

Palmer found this mistletoe near the “south end”, on Cupressus and more frequently on
Juniperus (Watson 1876:119). Bryant (1887:295) mentioned mistletoe on the cypress at
a height of 20 feet. However, Greene did not see it in 1885, and apparently no one has
found it since. | have examined most of the remaining junipers and could scarcely have
missed the mistletoe had it been there. | have also looked for it several times on some
hundreds of cypress trees and, strange to say, have failed to find it— though there | am
less certain it is absent. If it was originally rare on the cypress, could it have required
the large seed source on the junipers to ensure infection of the less susceptible cypress?
Or is it still there after all?

Fosberg (1941) doubtfully referred P. guadalupense to P. bollenanum var. pauciflorum
(Torr.) Fosberg, saying there was not enough material to decide its status. Wiens (1964:29)
wrote that the Guadalupe plant did not differ in any important way from P. bolleanum
ssp. densum.

Familia ignota

+Planta species
Sanicula menziesii? of Guadalupe Island references.
RANGE. — Presumably endemic to Guadalupe Island.

In the crevices of rocks in the “middle” of the island Palmer found two plants only,
without flowers or fruit, of what Watson (1876:115) doubtfully identified as Sanicula
menziesii Hook. & Arn. No one has since found anything like it, and presumably it is
extinct. Dr. Lincoln Constance examined the specimen and concluded it was not Sanicula
or any other Umbellifer. Since he and Dr. P. A. Munz and other botanists well-versed in
the flora of California failed to recognize even the family of this specimen, I assume that
it was at least an endemic species and so list it with the endemics.

The specimen at GH could not be found recently. The only trace of it is a faded and
blurred copy of a memo from Dr. Munz to Peter Raven (SD), with a sketch of one of
the three leaves. These were “palmately about 5-lobed, with very laciniate and irregular
margins, prominent venation, and distended petiole bases (like an umbellifer). If | were
going to check on it, I'd think Umbelliferae or maybe Jatropha.”

MONOCOTS
Gramineae
Aristida adscensionis L., Sp. Pl. p. 82. 1753.

Aristida bromoides HBK, Nov. Gen. Sp. 1:122. 1815.

RANGE. — California and Texas to Argentina; Santa Rosa, Santa Cruz, Santa Catalina,
San Clemente, and Cedros Islands.

Palmer found this grass in deep canyons among other grasses and plants (Vasey and Rose
1890:27). Hitchcock (1924:544) cited his collections and one by Rose (/695/). It is wide-
spread at low elevations on the island but not common.

158

GUADALUPE ISLAND FLORA
*Avena barbata Pott ex Link, Schrad. Jour. Bot. 2:315. 1799.
RANGE. — Native to Europe; now a widespread weed.

On his second trip Palmer found this grass in canyons and on rough slopes (Vasey and
Rose 1890:27). It is now widespread on the island and in some places fairly common. It
is the only member of the genus collected in recent years.

t+*Avena fatua L., Sp. Pl. p. 79. 1753.
RANGE. — Native to Europe; now a widespread weed.

Palmer found several patches in open places and on the best soil (Watson 1876:120).
Greene (1885:228) saw very little and remarked that apparently it was not established. It
has not been found recently. Gould and Moran (1981:40) tentatively identified Pa/mer
94a (MO) with 4. fatua though it has some characters of A. barbata.

+*Avena sativa L., Sp. Pl. p. 79. 1753.
RANGE. — Native to Europe; widely grown as a crop plant and sometimes escaping.

Gould and Moran (1981:40) identified Palmer 94 (MO) as an atypical form of this species.
Apparently it has not been collected since.

*Bromus berteroanus Colla, Mem. Acc. Turin 39:25. 1835.

Bromus trinit Desy. in Gay, Fl. Chil. 6:441. 1853.
Trisetum barbatum Steudel, Syn. Pl. Glum. 1:229. 1854.
Bromus sterilis of some Guadalupe Island references.

RANGE. — Native to southern South America; a weed in SW USA and NW Mexico.

On his second trip, in a year of frequent rains, Palmer found this grass abundant and
often large at the south end and very common also in the northern part of the island
(Vasey and Rose 1890:26, 27). He added that it made the best forage at the north end,
in the canyons very large and growing so thickly that it looked like grain. It is now
widespread at low elevations but generally scarce. | have never seen it common.

*Bromus diandrus Roth, Bot. Abh. p. 44. 1787.
Bromus maximus Desf., Fl. Atlant. 1:95. 1798.
Bromus rigidus of Guadalupe Island references.
Bromus sterilis of some Guadalupe Island references.

RANGE. — Native to Europe; a weed in SW USA and NW Baja California.

Greene (1885:228) reported this grass as abundant, saying that not even the goats were
fond of it. It is now widespread on the island, from the north end to Arroyo Melpomene,
sometimes locally common.

*Bromus hordeaceus L., Sp. Pl. p. 77. 1753.
Bromus mollis L., Sp. Pl. ed. 2, 1:112. 1762.

RANGE. — Native to Europe; a widespread weed.

159

REID MORAN

Moran (1951:159) found this grass occasional in the canyon mouth at Northeast Anchorage
in 1948. It has since spread up the northeast side of the island to the area of the springs.

*Bromus rubens L., Cent. Pl. 1:5. 1755.

Bromus madritensis ssp. rubens (L.) Husnot, Gram. Fr. Belg. p. 71. 1899.
RANGE. — Native to Europe; a widespread weed.

Howell (1942:146) collected this grass at Northeast Anchorage and reported it not common.
It has now spread upward to the pine ridge and Oak Canyon and south to Long Canyon
and in many places is rather common.

+*Bromus tectorum L., Sp. Pl. p. 77. 1753.
Bromus sterilis of some Guadalupe Island references.

RANGE. — Native to Europe; a frequent weed through much of the USA and in NW
Mexico.

Wallace (1985:87, 98) reported B. tectorum for Guadalupe Island, based on Palmer 99
(NY) in part. Watson (1876:120) had listed this collection as B. sterilis L., with Palmer’s
comment that it was on warm hillsides, sometimes in large patches as if sown, at the
“south end” and “middle”; but according to Wallace it is a mixture of B. fectorum and
B. trinii (= B. berteroanus}. Presumably it was B. berteroanus that was so abundant, for
B. tectorum has not otherwise been reported from the island and probably never became
well established.

+Dissanthelium californicum (Nutt.) Benth., in Hook.f., Icon. Pl. 14:56, pl. 1375.
1881.

Stenochloa californica Nutt., Jour. Acad. Nat. Sci. Philadelphia, Ser. 2, 1:189. 1848.
RANGE. —- Endemic to Santa Catalina, San Clemente, and Guadalupe Islands.

Palmer found this grass on warm rocky slopes in the “middle” of the island but not very
abundant (Watson 1876:120). He said it was very succulent and that the goats were very
fond of it. No one has found it since. Likewise, it has been found only once on Catalina
and once on San Clemente: evidently, it 1s extinct.

*Hordeum murinum |. ssp. glaucum (Steud.) Tzvelev, Novosti Sist. Vyssh. Rast.
8:67. 1971.

Hordeum glaucum Steud., Syn. Pl. Glum. 1:352. 1855.
Hordeum stebbinsui Covas, Madrono 10:17. 1949.

RANGE. — Native to Europe; a widespread weed.

Greene (1885:228) reported “only a few tufts” of H. murinum [ssp. glaucum?] near the
cabins on the plateau, “but the seed is there and it will hardly fail to become abundant.”
The ssp. glaucum is now widespread on the island and certainly one of the most abundant
plants.

160

GUADALUPE ISLAND FLORA
*Hordeum murinum L. ssp. leporinum (Link) Arcang., Comp. FI. Ital. p. 805. 1882.
Hordeum leporinum Link, Linnaea 9:133. 1835.
RANGE. — Native to Europe; a widespread weed.

The ssp. /eporinum is also widespread on the island but is less abundant.

*Lamarckia aurea Moench, Meth. Pl. p. 201. 1794.

RANGE. — Native to southern Europe; a common weed from California to Texas and
northern Mexico.

Meling Lopez (1985) was first to find this weed on the island (Zona Bosque, Meling
Lopez 041, SD). In 1988 it was occasional near the spring and above (3/009).

Melica imperfecta Trin., Mem. Acad. Imp. Sci. St.-Petersbourg, Ser. 6, Sci. Math.
2:59. 1836.

RANGE. — Central California to central Baja California; islands of southern California
and south to Cedros.

Palmer found this grass in crevices of high rocks in the “middle” of the island (Watson
1876:120). Greene (1885:228) found only one tuft in a place inaccessible to goats. It is
now rare, on cliffs in the northeast part of the island, seen in 1967 on the Lower Circus
at 950 m (M/3783) and in Southeast Oak Canyon at 400 m (M1/379/).

Muhlenbergia microsperma (DC.) Kunth, Rev. Gram. 1:64. 1829.
Muhlenbergia debilis (HBK) Kunth, Rev. Gram. 1:63. 1829.

RANGE. — SW USA to Peru; islands of southern California except San Nicolas and of
NW Baja California south to Cedros.

Palmer found this grass abundant on warm slopes in the “middle” of the island and less
common at the “south end” (Watson 1876:120). Palmer’s collections of 1889 were cited
by Hitchcock (1913:294). It is now widespread on the island, from the pine ridge to the
south end mesa, but not very common.

Phalaris caroliniana Walter, Fl. Carol. p. 74. 1788.

Phalaris intermedia Bosc ex Poir., Encycl. Suppl. 1:300. 1810.

RANGE. — Southern USA and northern Mexico; Santa Cruz, San Nicolas, Santa Barbara,
Santa Catalina, and San Clemente Islands.

Brandegee (1900) first reported this grass from Guadalupe Island, but without specific
locality. It appears to be local on the island and rather uncommon, in arroyos below
500 m west to northwest of the lobster camp. This little visited part of the island is where
Cistanthe maritima, Gnaphalium bicolor, Stipa lepida, and Stebbinsoseris heterocarpa
also appear to be local.

Most authors treat this grass as introduced in western North America, often saying it
grows in disturbed places; and there is no clear way to tell. However, its distribution on
the island and in wild parts of northern Baja California suggests to me that it may be

161

REID MORAN

native. Also, Raven (1963:315) quoted Dennis E. Anderson as feeling then that it might
well be native in California, even though later Anderson (1993:1282) has treated it as
introduced.

*Phalaris minor Retz., Obs. Bot. 3:8. 1783.

RANGE. — Native to the Mediterranean region; a widespread weed in North and South
America.

This grass was first found on Guadalupe Island in 1958, scarce at Northeast Anchorage
(M6789, Wiggins & Ernst 216). It persists in that area and also grows in an arroyo bed
near the southwest corner of the island (4/3/0175).

*Poa annua L., Sp. Pl. p. 68. 1753.
RANGE. — Native to Europe but now widespread in North America.

Howell (1942:147) found this grass in the pine grove. It is now abundant on the upper
north slope and about the spring and is scattered elsewhere in the northern part of the
island.

Poa secunda J. Pres] ssp. secunda

Poa secunda J. Presl, Relig. Haenk. 1:271. 1825-30.
Poa scabrella (Thurb.) Vasey, Grasses U.S. p. 42. 1883.

RANGE. — Alaska to northern Baja California; San Miguel, Santa Rosa, Santa Cruz, Ana-
capa, San Nicolas, Santa Catalina, San Clemente, and Todos Santos Islands.

This grass is known on Guadalupe Island from one collection: one clump on face of
Hemizonia cliff at 800 m, 3 May 1967 (M//38/6).

*Polypogon monspeliensis (L.) Dest., Fl. Atlant. 1:67. 1798.
RANGE. — Native to Europe; a widespread weed in North America.

Greene (1885:228) found this grass common about the springs. It is still common there
and occurs more sparingly under pines at the north end and in the streambeds of the
canyons draining the northeast part of the island and south to Juniper Canyon.

Stipa lepida A.S. Hitche., Amer. Jour. Bot. 2:302. 1915.

Nasella lepida (A. S. Hitche.) Barkworth, Taxon 39:610. 1991.

RANGE. — Northern California to southern Baja California; islands of southern California
and Los Coronados and Cedros.

This grass was first found on Guadalupe Island in 1958. It appears to be local on the
east side south of the middle of the island: east slope of El Picacho, 450 m (M6754,
Wiggins & Ernst 199), west of the lobster camp, 70m (Wiggins & Ernst 203).
*Triticum aestivum L., Sp. Pl. p. 85. 1753.

RANGE. — Native to Eurasia; widely cultivated and occasionally escaping but unlikely to

persist out of cultivation.

162

GUADALUPE ISLAND FLORA

Meling Lopez (1985) was first to find this grass on the island (Corrales, zona bosque,
Meling-Lopez 042, SD), and there seems to be no later record.

+*Vulpia bromoides (L.) Gray, Nat. Arr. Brit. Pl. 2:124. 1821.
RANGE. — Native to Europe; widespread in North America.

“Guadaloupe Islands,” April 1889, Dr. E. Palmer 674 (POM/RSA), as checked by Tim
Ross and as noted by Wallace (1985:33, 90). According to McVaugh (1956), this would
be from the northeast part of the island. I have seen no other record.

Vulpia microstachys (Nutt.) Benth. var. pauciflora (Beal) Lonard & Gould, Madrono
22:226. 1974.

Festuca pacifica Piper, Contr. U.S. Natl. Herb. 10:12. 1906.

RANGE. — British Columbia to southern Baja California; San Miguel, Santa Cruz, Ana-
capa, Santa Catalina, San Clemente, San Martin, and Cedros Islands.

Palmer found this grass in bunches on warm slopes and in open places in the “middle”
of the island (Watson 1876:120), and Greene (1885:228) listed it without comment. This
century it seems to have been found only rather scarce near the springs and old ranch
(Moran 6658, Wiggins & Ernst 114).

*Vulpia myuros (L.) C. C. Gmelin var. hirsuta Hack., Cat. Gramin. Portugal, p. 24.
1880.

Festuca megalura Nutt., Jour, Acad. Nat. Sci. Philadelphia Ser. 2, 1:188. 1848.

RANGE. — Bnitish Columbia and Montana to southern Baja California; islands of southern
California and Todos Santos; native to Europe.

On his second trip Palmer found this. grass in canyons, not common (Vasey and Rose
1890:27). It is now very common in the northern part of the island, from sea level to the
top of Mt. Augusta.

Vulpia octoflora (Walter) Rydb. var. hirtella (Piper) Henrard, Blumea 2:320. 1937.

Festuca octoflora Walter var. hirtella Piper, Contr. U.S. Natl. Herb. 10:12. 1906.
Festuca tenella of Guadalupe Island references.

RANGE. — British Columbia to Texas and central Baja California; islands of southern
California and Todos Santos, San Martin, and Cedros.

On his second trip Palmer found this grass in the southern part of the island (Vasey &
Rose 1890:26); his collections were cited by Hitchcock (1913:378) as F. octoflora var.
hirtella. Howell (1942:146) found it not common in the middle part of the island. It
appears to be rather local at widely separated places on the island: South Esparsa Canyon,
50 m (M6595); south flank of El Picacho, 650 m (M6741); Melpomene drainage, 300 m
(M6729A). On the flank of El Picacho it was abundant over a gently sloping area several
hundred meters wide.

163

REID MORAN

Juncaceae

Juncus bufonius L., Sp. Pl. p. 328. 1753.

RANGE. — Widespread on all continents; San Miguel, Santa Rosa, Santa Cruz, Anacapa,
San Nicolas, Santa Catalina, and San Clemente Islands.

Palmer found this rush from the “middle” to the north end of the island, abundant in
springy places (Watson 1876:120). It is still common locally in seasonally wet places
throughout the island, from the spring to tinajas in Arroyo Melpomene.

Liliaceae

Dichelostemma pulchellum (Salisb.) A. Heller, Muhlenbergia 1:132. 1906.

Brodiaea capitata Benth., Pl. Hartw. p. 339. 1857.

Brodiaea insularis Greene, Bull. Calif. Acad. Sci. 2:134. 1886.

Dichelostemma insulare (Greene) Burnham, Muhlenbergia 3:74. 1907.

Dipterostemon insulare (Greene) Rydb., Bull Torrey Bot. Club 39:111, 1912.

Brodiaea capitata Benth. var. insularis (Greene) J. F. Macbr., Contr. Gray Herb. 56:9. 1918.

RANGE. — Southern Oregon to central Baja California; islands of southern California and
south to San Benitos and Cedros.

Greene (1885:216) reported this plant exceedingly common on the tableland about the
spring and on the declivity below. On his second visit, Palmer found it quite plentiful in

eep ravines and on hillsides at the south end (Vasey and Rose 1890:26). It is widespread
on the island, from the north slope to Outer Islet. In the northern part it seldom reaches
flowering size except on cliffs. Farther south, beyond the range of most goats, it is fairly
common in some places and often does flower.

Brodiaea insularis, of the islands from San Miguel to Guadalupe, Greene (1886) separated
mainly on the basis of its larger size: “Closely allied to [B. capitata] but many times

larger... . Scape 3—5 feet high: leaves often a yard long and an inch broad: . . . corms
_ the largest in the genus, often two inches in diameter. . . .” However, the island

plants are by no means always large, and Hoover (1940:471) wrote that in favorable
situations and in favorable seasons plants of the mainland may grow as large as any seen
from the islands. Although he supposed that their often larger size probably had some
genetic basis, he did not separate the island plants taxonomically — nor have others. In
fact, the Guadalupe plant may not belong with the other island plants genetically.

Greene said he could not account for Palmer’s failure to either collect this plant or make
note of its presence on the island. This was remarkable considering that Palmer had
camped for 15 weeks where Greene now found it so common and, in fact, where he
collected three other plants Palmer had not found. The only explanation seems to be that
it was a late arrival not yet established on the island. | therefore list it as a probable
newcomer (Table 2).

Triteleia guadalupensis L. W. Lenz, Aliso 7:145, figs. 1, 2. 1970.
Brodiaea lugens and Triteleia ligens of Guadalupe Island references.

RANGE. — Endemic to Guadalupe Island.

164

GUADALUPE ISLAND FLORA

Corms 2-3.5 cm thick, the thick coat of coarse fibers extending upward to form a neck.
Leaves two, 3—11.5 dm long, 12-32 mm wide. Scapes 3—7.4 dm tall, 2-5 mm thick, smooth
or slightly scabrous; involucral bracts ovate, acuminate, 1—5 cm long. Flowers (Plate 2L)
in March to May, about 20-30, on pedicels 1.5—5 cm long. Perianth golden yellow, 27-32
mm wide, 14-21 mm long (closed), the tube funnelform, 5-9 mm long, the segments
widespreading, 12-14 mm long, 4-8 mm wide, papillose at apex, each with green to dark
purple midvein, the outer ovate, narrowly rounded at apex, the inner elliptic, broadly
rounded and emarginate at apex. Filaments all inserted at mouth of perianth tube, alter-
nately long and short, oblong or subcuneate, subtruncate and mucronate at apex, the mucro
bearing the anther, the longer S—6 mm long, 3 mm wide, the shorter 2—3 mm long; anthers
pale yellow, 2—3 mm long. Ovary twice as long as stipe. Chromosomes: n = 8. (Description
broadened from Lenz 1970).

Brandegee (1900:22) found this plant common along the slopes of Sparmanns Canyon
(perhaps Esparsa Canyon?), and Moran (1951:160) found it high at the north end. It grows
on shaded cliffs in the northern two-thirds of the island, in some places rather common
locally: Hemizonia Cliff, 800 m (M/38// ¢tc.); Oak Canyon 850 m (M//230/); Esparsa
Canyon 250 m (M5969); head of canyon west of the lobster camp (Weber & McCoy
12016) {near the following]; east slope of El Picacho, 350 m (4/2063, type collection).

In a careful revision of 7riteleia, Hoover (1941) placed Brandegee’s specimen under 7.
lugens Greene, a rare species which, in his sense, had a remarkable disjunct distribution
in the inner North Coast Ranges of California, the San Gabriel Mountains, and Guadalupe
Island. On the basis of living plants, Lenz (1970) separated 7. guadalupensis, with Moran
12063 (SD59967) as holotype; and he concluded that the plant of the San Gabriel Moun-
tains also was specifically distinct. Thus he restricted typical 7. /ugens to central California,
where it is now known from Monterey and San Benito Counties as well as the North
Coast Ranges. He distinguished 7. guadalupensis from T. lugens by its generally larger
size, its larger corms whose thick coat of coarse fibers extends above the top to form a
distinct neck, and its golden yellow perianth with a longer and broader tube. He said that
in flower color and shape it approached the plants of Monterey and San Benito Counties
more closely than it did the more northern ones. From field-collected buds of the type
collection, Dr. Marion Cave got a meiotic chromosome count of n= 8.

Palmae

Brahea edulis H. Wendl. in S. Watson, Proc. Amer. Acad. Arts Sci. 11:146. 1876.
Erythea edulis (H. Wendl.) S. Watson, Bot. Calif. 2:212. 1880.
RANGE. — Endemic to Guadalupe Island.

This is a palm to 10 m tall, with a clean columnar trunk 24 dm thick above the enlarged
base, with green costapalmate leaves 2—3.5 m long (Fig. 78), and with a finally pendent
inflorescence 1—3 m long bearing edible black drupes 2.5—3.5 cm thick (Fig. 79).

Palmer reported the palm as frequent in deep warm ravines, from the northern end to
Jacks Bay [West Anchorage] (Watson 1876:120). Franceschi (1893a) wrote that the only
considerable remaining grove was on the northwest slope, covering an area not less than
one mile and a half long, and one-half to one mile wide, though formerly extending
farther north as shown by burned trunks. He said that palms used to grow in the canyon
coming down to the landing [Barracks Canyon], “where man has destroyed them all for
thatching the cabins with their leaves and eating the central Cabbage, which is indeed

165

REID MORAN

FIGURE 78. The endemic Guadalupe Island palm (Brahea edulis) on the north slope of the island,
13 April 1970. This handsome palm in commonly grown in southern California and elsewhere.

delicate and sweet.” And he told of what appeared to be the scattered remains of a
formerly considerable grove on the two sides of Esparsa Canyon. Dudley (1899:283)
reported 50 or more palms in each of three canyons opening into the central valley [Arroyo
Melpomene] above the south anchorage. Brown mentioned a small grove on the west
side near Steamer Point (Thayer and Bangs 1908:102). Hanna (1925:237, pl. 19, fig. 2)
reported about a thousand palms remaining in Esparsa Canyon.

Much of the fine grove described by Franceschi still stands on the northwest slope (Fig.
24), extending about 3 km in a northeast-southwest direction and from the top of the
seacliffs at about 450 m up to 800 m and having probably several thousand trees. A few
grow among the pines to the northeast (Fig. 20). No palms remain in Barracks Canyon,
as Franceschi said, though a few old palm roots were still to be seen in 1960. Several
hundred palms remained in Esparsa Canyon in 1970, at about 250 to 400 m, but scarcely
the thousand seen by Hanna 50 years before. Palms remain in three other shallower canyons
on the east side, south as far as El Picacho. In the one emptying at the lobster camp I
counted 69 palms in 1957, at 250 to 450 m. On the west side there are still palms in the
canyon above West Anchorage; and in 1957 about 30 grew in a hanging valley at
350 m some 7 km south of El Picacho (Lindsay 1966:8, photo). I saw none in the tributaries
of Arroyo Melpomene.

All these palm groves are of old trees: no seedling escapes the goats. I have seen only
two palm seedlings on the island. One was in Arroyo Melpomene, but it has since dis-
appeared. Possibly this was from a seed dropped by someone carrying a palm inflorescence
to the village to feed a domestic goat once kept there. The other was in the arroyo not

166

GUADALUPE ISLAND FLORA

FIGURE 79. Fruit of the endemic Guadalupe Island palm (Brahea edulis)(M12066) on the north
slope of the island, 4 March 1965. Palmer remarked that the fruit was eaten by man, goats, birds,
and mice.

far above the lobster camp, where it was not likely to survive the goats for long.

Palmer said that the clusters of fruit (Fig. 79) weighed 40 or 50 pounds and that the fruit
was eaten by man, goats, birds, and mice. Franceschi wrote that the fruits were similar
in size and taste to prunes, and he evidently considered them an additional dividend to
be got from growing an excellent palm. At just the right stage between greenness and
decay, the fruits are indeed quite edible; but to me they are less like prunes than like
some inferior kind of date.

Bailey (1937) had six species of Erythea, native in northwest Mexico; Uhl and Dransfield
(1987) recognized about 16 species of Brahea, including Ervthea, native to Mexico and
Guatemala, though saying that a new taxonomic treatment was much needed. The nearest
species geographically is B. armata, of northern and central Baja California. An interesting
question is how the original palm seed got to the island (Moran 1978). The fruits and

167

REID MORAN

seeds of B. edulis are too dense to float in sea water, and it would take an enterprising
bird to carry so heavy an object so long a distance. The inflorescence might float; but it
is most unlikely either to become detached or to be carried down a turbulent rain-swollen
arroyo to the sea while still bearing fruit.

Guadalupe Island palm is well established in cultivation in southern California and else-
where and makes a very handsome tree. Its merits greatly impressed Dr. F. Franceschi,
a noted horticulturist, later of Santa Barbara but in 1892 newly arrived in California (cf.
Tucker 1945); and in the winter of 1892—93 he visited Guadalupe Island largely to gather
fuller information about it. Writing later about the palm in its island home and in culti-
vation, Franceschi (1893a) noted that fresh seed germinate in a few weeks, though older
ones take several months; and that afterwards this palm grows nearly as quickly as Wash-
ingtonia filifera and Phoenix canariensis, two well-known fast growers. And he wrote
that horticulturists “‘are sure to find it to be the most elegant of all Fan Palms from the
North American continent.”

Zosteraceae

Phyllospadix torreyi S. Watson, Proc. Amer. Acad. Arts Sci. 14:303. 1879.
RANGE. — Oregon to central Baja California; islands of southern California.

Dudley (1899:283) reported this surf-grass as scarce, near the west and south anchorages.
It is common enough on the intertidal reef at West Anchorage and off the landing at
Campo Sur. Also, it was collected for me by diver Wes Andrew at the south end and by
him and by divers Jim Stewart and Jay Quast at Northeast Anchorage.

Zostera marina L., Sp. Pl. p. 968. 1753.

RANGE. — Alaska to Baja California and Atlantic coast; Santa Rosa, Santa Cruz, Anacapa,
Santa Catalina, Los Coronados, San Martin, San Geronimo, and Cedros Islands.

This eel-grass was collected for me in January 1960 by divers Jim Stewart and Jay Quast
at 140 feet at Northeast Anchorage (/7844). Anne Mower annotated the sterile specimen
in 1969, saying it looked like the shallow-water form.

DOUBTFUL AND EXCLUDED SPECIES

Chamaesyce melanadenia (Torr.) Millsp., Field Mus. Nat. Hist. Bot. Ser. 2:410. 1916.

Euphorbia melanadenia Torr., Pacit. Railr. Rept. 4:135. 1857.

Wheeler (1934, 1936:439) reported this species from Guadalupe Island on the basis of
specimens (F, GH) collected by Palmer in 1889. On two specimens the labels — one
handwritten, the other typed — say “Lagoon Head” but not “Guadalupe Island.” A third
label has printed on it “Lagoon Head” and “March 9, 1889”; but “Guadalupe Island,
Lower Calif.” is written below. The field number (783) falls in the series for Lagoon
Head and inland rather than in that for Guadalupe Island (McVaugh 1956:239, 230), and
the date agrees. Furthermore, Vasey and Rose (1890:12) cited this collection as from hills
40 miles back from the ocean, Lagoon Head, Baja California, showing they had fuller
information than appeared on these labels. Thus the report for Guadalupe Island seems
clearly mistaken.

168

GUADALUPE ISLAND FLORA

Pachycereus pringlei (S. Watson) Britton & Rose, Contr. U.S. Natl. Herb. 12:422.
1909.

When in a letter to Spencer Baird, Xantus improvised Cereus giganieus on Guadatupe
Island, Zwinger (1986:88, note 13) wrote: “Cereus giganteus, Saguaro, does noi reach
this far west and south; what Xantus saw was Pachycereus pringlei, Cardon, 2 then-un-
collected species.” It is still uncollected, however, and there is no reason to believe thai
Xantus saw any giant cactus on the island; see under Xantus, p. 45.

Solanum tenuilobatum Parish, Proc. Calif. Acad. Sci. Ser. 3, 2:165. 1901.

A specimen of this species (SD119427) is labeled as from Guadalupe Island: Arroyo,
Zona Sur, 3km N Morro Sur, Alf Enrique Meling L., 30 March 1982. This rare plant ts
otherwise known from a few places in chaparral at 200-1100 m in southwesiem San
Diego County, California (Beauchamp 1986:229), and northwest Baja California. | ques-
tion whether this plant would grow in such a dry place as the south end of the island
and reserve judgement about its occurrence on the island. See Enrique Meling, p. 52.

Stillingia linearifolia S. Watson, Proc. Amer. Acad. Arts Sci. 14:297. 1879.

The circumstances for this species closely paraliel those aescribed above for Chamaesyce
melanadenia. Rogers (1951, fig. 21) showed a dot for Guadalupe {sland on his map for
this species; and in the list of specimens (p. 248) he cited E. Palmer 785 (F, GH, US)
from “lagoon head, Guadaloupe Island.” At least one of these specimens (F) has the
printed “Lagoon Head” label with “Guadaloupe Island” wrtien below. However, number
785 again falls in the series for Lagoon Head and inland rather than in that for Guadalupe
Island (McVaugh 1956:239, 230); and Vasey and Rose (1890:12) cited this collection as
from Lagoon Head. Thus this report for Guadalupe !sland !ikewise seems clearly mistaken.

In tabulating plant distributions on the Channel islands o: California plus Guadalupe
Island, Dunkle (1950), on the pages shown, marked for Guadalupe Island the following
plants for which I have found no other record: Cryptaniha clevelandii Greene var. hispidis-
sima (Greene) I. M. Johnst. (pp. 306, 330), Fceveria greenci (Rose) A. Berger [= Dudleva
greenei Rose] (p. 304), Lycium richii A. Gray var. houssei (Greene) I. M. Johnst. [= L.
brevipes Benth. var. hassei (Greene) C. L. Hitche. |(o. 306), and Senecio lyonii A. Gray
(p. 307). Assuming these to be clerical errors, | exciude them pending further evidence.
Apparently Dunkle never visited the island.

In a list of San Clemente Island plants also growing on one or more of the other Channel
Islands and the adjacent mainland, Raven (1963:2°8) starred the name of Cressa truxi!-
lensis var. vallicola to show it was also on Guadalupe Island. I have seen no other record
and think this probably a clerical error.

Wiggins (1980) cited for Guadalupe Island the following previously unreported species:
Amsinckia spectabilis Fisch. & C. A. Mey. (p. 217), Galvezia juncea (Benth.) Ball! var.
juncea (p. 495), Harfordia macroptera (Benth.) Greene & Parry (p. 168), Hemizonia
streetsii A. Gray (p. 37 but not p. 362), Navarretia hamata Greene ssp. hamata (p. 405),
Polyearpon depressum Nutt. (p. 622), and Ruppia maritima L. (p. 953). Dr. Wiggins
visited the island only once, in April 1958, with Sherwin Carlquist, W. R. Ernst, and me:
and we collected pretty much together. The list of Wiggins and Ernst collections includes
none of these species, and I believe he collected none of them. Wallace (1985) showed
some of these species for Guadalupe Island in his distribution table, but for each he
referred to Wiggins (1980), thus saying that he neither accepted nor denied the report.

169

REID MORAN

| have seen no specimens and no other record of any of these species from Guadalupe
Island, and | exclude them all from the list of Guadalupe Island plants for want of evidence.

Acknowledgements

During the half century of my interest in Guadalupe Island, so many people have helped
in so many ways that I can scarcely even list them. First, I am grateful to the Mexican
officials who granted permission to visit the island and to collect plants, to all those who
helped me reach the island, including many captains and crews, and to my various com-
panions ashore. | think first of George Lindsay and our great island adventure of 1948,
with Lewie and Marcho Cavanagh on their 36-foot ketch Marviento, and of his continuing
help and support. Then I am especially indebted to Professor Carl Hubbs of the Scripps
Institution of Oceanography, who kindly included me on many of his trips and who took
a great interest in my work with the plants. Others who took me on valuable and memorable
trips to the island, include the Pacific Project of the Smithsonian Institution, Harold Pringle
on Pastime, Bill Libby and his pinecone crew, Margie Stinson and Capt. Eddie McEwan
on Pacific Queen, and Dr. Martin Gonzalez and the Mexican Navy.

For the use of their specimens and for many other courtesies I thank the curators of the
following herbaria: CAS, GH, NCC, ND, NY, RSA, SD, UC, US. Particularly helpful
with references from library and herbarium were Annetta Carter, Judy Gibson, and Billie
Meeder. Martin Gonzalez kindly lent me his copy of his valuable report to the Mexican
government on the natural resources of the island.

Among the botanists who have given information or advice about particular plants, often
of their favorite families, are Marion Cave, Lincoln Constance, Alva Day, Lauramay
Dempster, Wally Ernst, Paul Fryxell, C. L. Hitchcock, Fran Hommersand, Tom Howell,
Ivan Johnston, Bill Libby, George Lindsay, C. H. Muller, Bob Ornduff, Peter Ray, Peter
Raven, José Rico C., Reed Rollins, Tim Ross, Dale Smith, John Strother, Charles Uhl,
and William A. Weber.

Walter and Irja Knight gave advice and help with the photographs and other matters,
Colleen Sudekum drew the maps, José Delgadillo made the climatic diagram, and Dr.
Alejandro Rodriguez kindly translated the abstract into Spanish.

Those who have reviewed the manuscript or read large parts of it, catching small and
large errors and suggesting improvements, include Frank Almeda, Claire Brey, Tony Bur-
gess, Sherwin Carlquist, Tom Daniel, Steve Junak, Chuck Quibell, Peter Raven, and Bob
Thorne.

To all these good people I am grateful for their invaluable help at various stages of this
work.

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184

GUADALUPE ISLAND FLORA

Index to Plant Names

Accepted names are in roman type, synonyms and misapplied names in italic. Page
numbers in bold are for principal references only.

Agoseris heterophylla 21, 23, 40, 46, 78
Alchemilla occidentalis 144
Allocarya acanthocarpa 69
Allophyllum gilioides 40, 134
Amblyopappus pusillus 34, 36, 38, 79
Ambrosia camphorata 26, 31, 34, 43, 77,
79, 90
Amsinckia intermedia 67
menziesii 31, 67
spectabilis 67, 169
vernicosa 67
Anagallis arvensis 20, 141
Antirrhinum kingii watsonii 149
nuttallianum pusillum 149
nuttallianum subsessile 148
pusillum 149
speciosum 151
subsessile 149
watsonil 22, 36, 43, 50, 149
Aphanes occidentalis 34, 144
Aphanisma blitoides 34, 36, 38, 47, 76
Arctostaphylos sp. 40, 46, 107
Aristida adscensionis 19, 158
bromoides 158
Artemisia californica 31, 34, 40, 77, 79
Aspidium munitum 56
Atriplex barclayana 34, 36, 38, 43, 77, 90
barclayana dilatata 77
barclayana palmeri 77
californica 23, 38, 41, 49, 77
dilatata 77
palmeri 77
rosei 77
Avena barbata 20, 30, 159
fatua 20, 159
sativa 20, 159
Baeria chrysostoma palmeri 87
coronaria 21, 88
gracilis 21, 87
palmeri 87
Baeriopsis guadalupensis 34, 36, 38, 39,
42, 49, 80
Bahia lanata 81
Brahea edulis 30, 42, 165
Brassica campestris 102
nigra 20, 101

rapa 20, 46, 102
Brodiaea capitata 164
capitata insularis 164
insularis 164
lugens 164
Bromus berteroanus 20, 159, 160
diandrus 20, 30, 159
hordeaceus 20, 50, 159
madritensis rubens 160
maximus 159
mollis 159
rigidus 159
rubens 20, 30, 49, 160
sterilis 159, 160
tectorum 20, 160
triniti 159, 160
Calandrinia ciliata 31, 137
ciliata menziesii 137
discolor 140
grandiflora 140
maritima 34, 140, 141
menziesii 137
Calamintha palmeri 115
Calystegia macrostegia macrostegia 30,
34, 38, 44, 96
Camissonia guadalupensis 128, 130
guadalupensis clementina 130
guadalupensis guadalupensis 41, 42,
44, 128
robusta 22, 23, 50, 51, 130
Capsella bursa-pastoris 20, 34, 47, 102
procumbens 103
Castilleja attenuata 41, 50, 149
exserta 23, 40, 47, 149
foliolosa 150
fruticosa 26, 34, 38, 39, 42, 50, 150
guadalupensis 40, 42, 48, 150
Caulanthus lasiophyllus 103
Ceanothus crassifolius 30, 40, 143
crassifolius planus 143
cuneatus 40, 46, 143
perplexans 143
Centaurea melitensis 20, 46, 80
Cerastium glomeratum 20, 34, 71
Cereus giganteus 45, 168
Chamaesyce melanadenia 107, 168, 169

185

REID MORAN

pondii 41, 43, 46, 107
Chamomilla suaveolens 89
Cheilanthes newberryi 34, 53
Chenopodium album 78
ambrosiodes 78
murale 20, 36, 46, 78
Cistanthe guadalupensis 36, 38, 39, 42,
67, 137, 140
maritima 41, 47, 141, 161
Claytonia perfoliata mexicana 31, 141
Clinopodium palmeri 115
Collomia gilioides glutinosa 134
Convolvulus macrostegius 96
occidentalis 96
occidentalis macrostegius 96
Coreopsis gigantea 30, 38, 39, 80
Corethrogyne cana 83
Coryneum cardinale 61
Cotvledon sp. 100
Crassula connata 31, 34, 38, 97
connata erectoides 98
connata eremica 98
Cressa truxillensis vallicola 96, 169
Crossosoma californicum 30, 31, 38-40,
44, 48, 101
Cryptantha clevelandii hispidissima 169
foliosa 31, 34, 38, 42, 68
maritima genuina 68
maritima maritima 34, 36, 68
Cupressus forbesii 58, 60, 61
goveniana 58
guadalupensis 56, 60, 61
guadalupensis forbesii 60, 61
guadalupensis guadalupensis 30, 42, 56
macrocarpa 56, 61
macrocarpa guadalupensis 56
sargentii 58
Cuscuta corymbosa 22, 41, 43, 50, 96, 97
corymbosa grandiflora 97
corymbosa stylosa 97
Daucus pusillus 19, 30, 41, 156
Descurainia pinnata menziesii 38, 41, 102
pinnata halictorum 102
Dichelostemma insulare 164
pulchellum 21, 23, 30, 38, 46, 164
Diplostephium canum 83
Dipterostemon insulare 164
Dissanthelium californicum 40, 44, 46,
160
Dodecatheon clevelandii insularis 41, 142
media 142

Dudleya greenei 100, 169
guadalupensis 36, 38, 39, 43, 50, 98
hassei 100
virens 47, 100, 101
virens extima 34, 43, 44, 50, 100
virens hassei 101
virens insularis 101
virens virens 101

Echeveria greenei 169

Echinocystis guadalupensis 105

Ellisia chrysanthemifolia 112

Emmenanthe penduliflora 112

Endocronartium harknessi 66

Epilobium foliosum 40, 130
minutum 130
minutum foliosum 130

Eriogonum molle 135
giganteum 136
giganteum formosum 136
zapatoense 30, 38, 39, 43, 44, 50, 135

Eniophyllum caespitosum 81
lanatum grandiflorum 40, 81, 131

Eritrichium angustifolium 68
muriculatum 68
ramosissimum 68

Erodium brachycarpum 20, 50, 111
cicutarium 20, 30, 34, 111
moschatum 20, 30, 34, 111

Erysimum insulare 102
morani 38, 39, 43, 50, 102
suffrutescens 102
suffrutescens grandifolium 103

Erythea edulis 165

Eschscholzia californica californica 19,

21, 23, 40, 46, 131
californica hypecoides 131
crocea 131
elegans 43, 131, 133, 134
elegans ramosa 132, 133
frutescens 132, 133
gigas 131
palmeri 34, 36, 38, 43, 46, 132, 133
ramosa 44, 133, 134
rigida 131
Eucrypta chrysanthemifolia
chrysanthemifolia 112
chrysanthemifolia bipinnatifida 112

Eunanus latifolius 152

Euphorbia guadalupensis 107
melanadenia 168
misera 30, 31, 38, 39, 45, 48, 108

pondii 107
Festuca megalura 163
octoflora hirtella 163
pacifica 163
tenella 163
Filago arizonica 81
californica 21, 23, 31, 46, 81
Frankenia grandiflora 111
salina 48, 111
Franseria bipinnatifida 79
camphorata 79
Galium angulosum 31, 40, 43, 146
aparine 20, 34, 52, 147
Galvezia juncea juncea 148, 169
speciosa 30, 31, 36, 38, 39, 44, 151
Gambelia speciosa 151
Gasoul crystallinum 66
nodiflorum 66
Gilia divaricata 134
guadalupensis 135
multicaulis 135
multicaulis millefoliata 135
multicaulis nevinii 135
nevinii 31, 44, 135
pusilla 135
pusilla californica 135
pyvgmaea 135
Githopsis diffusa candida 71
diffusa guadalupensis 34, 41, 43, 71
specularioides 71
Gnaphalium bicolor 41, 50, 83, 161
chilense 83
sprengelli 83
stramineum 41, 83
Guillenia lasiophylla 103
Gymnogramma triangularis 54
Haplopappus canus 30, 31, 40, 44, 83
detonsus 84
Harfordia macroptera 135, 169
Harpagonella palmeri 41, 68, 69
Hazardia cana 83
detonsa 83
Hedeomoides tenuiflora 114
Hemizonia clementina 84, 87
frutescens 34, 43, 84, 85

greeneana greeneana 34, 36, 38, 39,

43, 46, 49, 77, 84, 85, 87
greeneana peninsularis 85
minthorni 84, 87
palmeri 34, 43, 46, 80, 87
streetsil 84, 87, 169

187

GUADALUPE ISLAND FLORA

Herniaria cinerea 20, 50, 71
Hesperelaea palmeri 28, 40, 43, 46, 128
Hesperocnide tenella 40, 156
Hesperonia heimerlii 127, 128
Heteromeles arbutifolia macrocarpa 28,
40, 50, 144, 145
Heuchera ? 147
Hordeum glaucum 160
leporinum 160
murinum glaucum 20, 160
murinum leporinum 20, 160
stebbinsii 160
Hosackia argophylla 115
grandiflora 115, 116
ornithopus 115
Hutchinsia procumbens 38, 41, 49, 103
Hypochoeris glabra 20, 34, 49, 87
Jepsonia malvifolia 34, 44, 147, 152
parryi 147
Juncus bufonius 19, 163
Juniperus californica 30, 62
Krynitzkia foliosa 68
maritina 68
ramosissima 68
Lamarckia aurea 20, 52, 161
Lasthenia californica 21, 87, 150
chrysostoma 87
coronaria 20, 23, 47, 88
Lavatera insularis 121, 123
lindsayi 39, 43, 44, 50, 121

occidentalis 26, 30, 31, 39, 40, 44, 121,

123
Layia elegans 88
platyglossa 21, 22, 40, 47, 88
Lepidium bipinnatifidum 103
lasiocarpum 103
lasiocarpum latifolium 31, 43, 103
menziesii 103
nitidum 23, 49, 103
oblongum 103
oblongum insulare 31, 103
Lepigonum macrothecum 74
Leptosyne gigantea 80
Limnia guadalupensis 141
Linanthus pygmaeus continentalis 135
pygmaeus pygmaeus 31, 44, 135
Linaria canadensis 19, 41, 52, 152
canadensis texensis 152
Lomatium insulare 34, 39, 44, 50, 156
Lotus argophyllus ornithopus 30, 39, 115
grandiflorus 26, 41, 52, 115, 116

REID MORAN

grandiflorus mutabilis 115 californica cedrosensis 128
guadalupensis 115, 116 heimerlii 127, 128
leucophaeus 116 laevis 127
Lupinus affinis 117 Montia perfoliata 141
aliclementinus 117 Muhlenbergia debilis 161
bicolor 23, 48, 116 microsperma 19, 161
bicolor microphyllus 116 Myosurus minimus filiformis 34, 41, 46,
clementinus 117 142
guadalupensis 41, 44, 46, 116 Nasella lepida 162
moranit 117 Navarretia hamata hamata 134, 169
nanus 117 Nemophila aurita 113
niveus 34, 43, 117 racemosa 113
Lycium brevipes hassei 169 Neomammillaria goodridgii 70
californicum 34, 36, 38, 39, 45,67, 153 = Nesothamnus incanus 89
fremonti 41, 50, 153 Nicotiana attenuata 40, 48, 153, 154
richii hassei 169 bigelovii 153
Malacothrix cleveland 40, 89 glauca 20, 26, 50, 154
Malosma laurina 30, 31, 40, 67 petuniaeflora 40, 153, 154
Malva borealis 123 Notholaena californica 54
parviflora 20, 30, 34, 123 newberryi 53
Mammillaria blossfeldiana shurliana 36, Oenanthe guadalupensis 129, 130
38, 39, 43, 46, 70 Oenothera guadalupensis 128, 129
goodridgii 70 micrantha 20, 130
Marah guadalupensis 43, 104, 105, 107 Oligomeris glaucescens 143
macrocarpus 105 linifolia 19, 34, 36, 39, 143
macrocarpus major 105 subulata 143
major 105, 107 Opuntia prolifera 46, 70
oreganus 107 Orthocarpus attenuatus 149
Matricaria discoidea 89 purpurascens 21, 149
matricarioides 41, 89 Pachycereus pringlei 70, 168, 169
Medicago hispida 117 Parietaria debilis 157
polymorpha 20, 49, 117 floridana 19, 157
polymorpha brevispina 117 hespera hespera 31, 34, 36, 38, 39, 157
Megarrhiza guadalupensis 104 Pectocarya linearis ferocula 23, 50, 69
Melica imperfecta 41, 161 penicillata 69
Melilotus indica 20, 46, 117 recurvata 69
Mentzelia dispersa 120, 121 Pellaea mucronata mucronata 54
micrantha 20, 120, 121 ornithopus 54
Mesembryanthemum crystallinum 20, 34, Pentagramma triangularis 34, 54, 55
36, 38, 39, 46, 66 triangularis maxon 54
nodiflorum 20, 34, 36, 49, 66 triangularis triangularis 54
Micrampelis guadalupensis 105 triangularis viscosa 55
Micropus californicus 40, 46, 89 Pericome 91
Microseris heterocarpa 93 Perityle californica 89
lindleyi 93, 95 emoryi 34, 36, 39, 89
linearifolia 95 grayi 89
Mimulus brandegei 152 incana 25, 30, 31, 36, 38, 39, 42, 43,
latifolius 41, 44, 152 89, 91, 96
traskiae 152 Petromecon frutescens 42, 132
Mirabilis californica 34, 36, 38, 39, 127, palmeri 132, 133
128 Phacelia floribunda 36, 39, 44, 113

188

phyllomanica 30, 31, 41, 43, 48, 113
phyllomanica interrupta 113
Phalaris caroliniana 47, 161
intermedia 19, 161
minor 20, 50, 162
Pholistoma racemosum 31, 39, 113
Phoradendron bolleanum densum 157,
158
bolleanum pauciflorum 157, 158
densum 40, 46, 157
guadalupense 157, 158
Photinia arbutifolia macrocarpa 144
Phyllospadix torreyi 48, 168
Pinus insignis binata 62
muricata 62, 64
muricata cedrosensis 64, 65
radiata binata 28, 43, 62, 63, 64, 66
radiata guadalupensis 62, 64
remorata 62, 64
Pityrogramma triangularis 54
Plagiobothrys acanthocarpus 21, 23, 41,
50, 69, 70
californicus 70
collinus californicus 23, 49, 70
collinus gracilis 70
Planta sp. 40, 42, 43, 46, 158
Plantago fastigiata 134
insularis 134
ovata 19, 34, 38, 134
patagonica 134
Platystemon californicus 21, 23, 40, 48,
134
Poa annua 20, 49, 162
scabrella 162
secunda secunda 34, 41, 50, 162
Pogogyne tenuiflora 40, 43, 46, 114
serpylloides 115
Polycarpon depressum 71, 169
Polypodium californicum 34, 55, 152
scouleri 34, 55, 131
Polypogon monspeliensis 20, 46, 162
Polystichum munitum 34, 41, 56
munitum solitarium 56
solitarium 56
Pterostegia drymarioides 31, 34, 137
Quercus chrysolepis 108, 110
crassipocula 110
tomentella 30, 44, 108, 110, 111
Ranunculus hebecarpus 34, 142
Rhamnus crocea 143
crocea pirifolia 143

189

GUADALUPE ISLAND FLORA

pirifolia 30, 31, 40, 44, 143
Raphanus sativus 20, 52, 104
Rhus integrifolia 22, 30, 39, 50, 67
laurina 67
Ribes sanguineum 40, 46, 112, 131
Ruppia maritima 169
Ruta chalepensis 20, 50, 147
graveolens 147
Sairocarpus pusillus 149
watsonii 149
Sanicula menziesii 158
Satureja palmeri 40, 43, 115
Scrophularia villosa 30, 31, 40, 44, 45,
50, 153
Senecio lyon 169
palmeri 23, 30, 31, 41, 43, 91
Sida hederacea 126
leprosa hederacea 126
Silene antirrhina 19, 40, 46, 74
gallica 20, 30, 34, 74
Sisymbrium canescens 102
irio 20, 49, 52, 104
orientale 20, 50, 104
reflexum 103
Solanum americanum 20, 154
arborescens 155
calvum 154
clokeyi 155
douglasii 41, 155
nigrum 155
nigrum douglasii 155
nodiflorum 154
profundeincisum 155
tenuilobatum 52, 169
wallacei 155, 156
wallacei clokeyi 30, 41, 44, 48, 155,
156
xanti 155, 156
xanti wallacei 155, 156
Sonchus oleraceus 20. 30, 34, 38, 92
tenerrimus 20, 46, 92
Specularia biflora 71
Spergularia macrotheca 38, 39, 46, 74, 76
macrotheca talinum 74
marina 23, 49, 76
Sphaeralcea palmeri 34, 38, 39, 43, 46,
80, 125
sulphurea 30, 34, 43, 125, 126
Stellaria nitens 34, 76
Stebbinsoseris heterocarpa 41, 48, 93, 161]
Stenochloa californica 160

REID MORAN

Stephanomeria diegensis 21, 22, 40, 48, 93
exigua deanei 93
guadalupensis 30, 34, 39, 43, 45, 48, 93
Stillingia linearifolia 107, 169
Stipa lepida 41, 50, 161, 162
Stylophorum 132
Suaeda californica 78
californica pubescens 78
fruticosa 20, 78
taxifolia 46, 78
torreyana 78
Syrmatium ornithopum iW5
Talinum guadalupense 48, 137, 140
Thelypodium lasiophyllum 103
Thysanocarpus erectus 41, 43, 104
Tillaea erecta 97
leptopetala 97
minima 97
Tissa macrotheca 74
pallida 74
talinum 74, 76
Trifolium amplectens 118
depauperatum truncatum 118
gracilentum 23, 49, 119

190

gracilentum palmeri 119
hydrophilum 119
microcephalum 119
minutiflorum 118
palmeri 31, 34, 38, 44, 119
truncatum 119
Triodanis biflora 71
Trisetum barbatum 159
Triteleia guadalupensis 34, 43, 45, 48,
164, 165
lugens 164, 165
Triticum aestivum 20, 52, 162
Troximon heterophyvilum 78
Uropappus lindleyi 52, 95
Vicia exigua 120
exigua hassei 120
hasse1 120
ludoviciana ludoviciana 120
Volvulus macrostegius 96
Vulpia bromoides 20, 163
microstachys pauciflora 163
myuros hirsuta 20, 30, 163
octoflora hirtella 163
Zostera marina 50, 168

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