PALLENIS MARITIMA (ASTERACEAE) NEW TO CALIFORNIA, WITH NOTES ON RECENT INTRODUCTIONS OF SALT-TOLERANT ORNAMENTAL PLANTS

2012·4.6 MB·English

by  Richard E Riefner

NEW
MARITIMA TO
PALLENIS (ASTERACEAE) CALIFORNIA,
ON
WITH NOTES RECENT INTRODUCTIONS OF
SALT-TOLERANT
ORNAMENTAL
PLANTS
Werner
Richard Greuter
Riefner,
E.
Jr.
Research Associate
Ana
Rancho Santa Botanic Garden c/o Orto Botanico
Avenue
1500 North College Via Lincoln 2/A
Claremont, California 9171 1-3157, U.S.A. 1-90121 Palermo, ITALY
[email protected]
>n, extending to S. Portugal along the
on rocky shores and sea mostly
cliffs
Keywords:
Pflllenis maritima (=Asteriscus maritimus), California, halophytes, non-native plants,
c
lilli
INTRODUCTION
Pallenis maritima (L.) Greuter was not reported for California in The Jepson Manual (Keil 2012a; Jepson Flora
Project 2012), the Consortium of California Herbaria (2012), or other publications that address non-native spe-
&
des
growing Dean
DiTomaso Healy 2008;
outside of cultivation in California (Hrusa 2002; 2007; et al.
et al.
Roberts
2008). Pallenis maritima was not included in the Flora of North America treatment of the Inuleae,
also
“or has USDA, NRCS
been
it reported North America (Preston 2006; 2012).
at outside of cultivation, for
all,
In we and North America,
this paper, provide the documented records of maritima for California
first P.
w
be
fe it grows spontaneously outside of cultivation on rocky shores and sea cliffs in coastal Orange County,
southern We and
California. and review horticultural uses, seashore ecology,
provide voucher documentation,
a
PParent mode We taxonomy and nomenclature of the species
of
^ introduction review the
into the also
State.
,ts Placement within the Asteriscus alliance, in view of the fact that it is still often referred to as Asteriscus
B
621
6(2): -629.2012
622
Cove and Creek Beach, 33° 29'
between Mussel Salt
Dana rocky shore
11-86 (RSA); City of Point,
Riefner
,
seep
m-5 m, subshrubs and seedlings in rock crevices at the edge of an ephemeral calcareous-saline c
2.5
May 12-393 (RSA).
on the coastal strand, 25 2012, Riefner
name has some
known maritimus (L.) Less., a that still usage at
maritima has long been as Asteriscus
Patterns
and
name once taxonomic placement limits are
As any given species can only bear one correct its ac-
present.
may understand the disparity. That background
background be helpful to
an overview of the historical
cepted,
which
which maritima has been placed through time, or to it is currently as-
the genus in P.
is fairly intricate:
Athalmum Kuntze, Bubonium
names: Asteriscus Mill., Hill,
has been given no fewer than eight different
signed,
and Michon. For more
Saulcya
de-
Odontospermum Sch. Patterns Cass.,
Buphthalmum L„ Nauplius Cass., Bip.,
see Greuter (1997).
tails,
and Macaronesian species, together forming a
Mediterranean
maritima belongs to a group of 15
Patterns
by morphology and chromosome number that has been called the Asteriscus alliance (An-
natural unit defined
have been monographed by
These species
Goertzen 2002).
derberg 1994; Francisco-Ortega et al. 1999; et al.
known North and the Atlantic Islands and
endemics of Africa
Most
Wiklund (1983, 1985, 1987). are local, little
m
known even
were botanists pre-lin-
The most widespread, however, to
need not be considered here. three
annual
the to biennial
naean times. They are the upright, annual, non-pungent Asteriscus aquaticus (L.) Less.,
which
by pungent involucral bracts, and the perennial P. maritima, is
characterised
Pallenis spinosa (L.) Cass.,
discussed in this paper.
and named by Toumefort
described
These three species together formed the original genus Asteriscus, first
The name, he explained, means star” and refers to the star-like appearance of the “calyx” (invohi-
(1700). “little
was maintained
Toumefortian concept
The remarkably natural
crum) surrounding the “flower” (capitulum).
when included Asteriscus in
by Miller (1754) when he validated the generic name, but was lost Linnaeus (1737)
own new Buphthalmum name means “ox Linnaeus (1753) redefined Buphthal-
genus (the eye”).
his
artificial
he descnbed
mum but did not succeed in making more natural. Of the 11 Buphthalmum species
considerably,
it
no fewer than seven
placed in
and named over the years only two are today, the remainder are currently
left
different genera (Jarvis 2012).
Most 19th century au-
which
summarized post-Linnaean taxonomy, outlined here.
Greuter (1997) the is
two based on the
but in
thors essentially reverted to the Toumefortian generic concept of Asteriscus, split it
and Asteriscus
maritima
obvious character of pungent vs. unarmed involucral bracts. That is, they left Pallenis
who named the spiny
aquaticus together, but separated Pallenis spinosa. The to do so was Cassini (1822),
first
unarmed ones Nauplius
Athens— and
plants Pallenis (after the borough Pallini E. of Fournier 1934-1940) the
but too
division
(perhaps, by analogy, after the Greek harbour, Nauplion). Subsequent authors accepted the
was that
The problem
up the name Asteriscus for one of the genera—unfortunately not always for the same one.
monograp
A and
the
Asteriscus had not been typified. type was not designated until very late (Jeffrey 1982),
Wiklund
(1985) refused to accept Jeffrey’s choice ofA. aquaticus.
&
differ-
micromorphological
Meanwhile had profound
Briquet (in Briquet CavilUer 1917) discovered
spiny
from their
ences between the two unarmed species, placing them at least as far apart from each other as
went one step
Wiklund 1987)
relative. By consequence, he placed each of the three in a genus of own. (1985,
its
Aste^
mis-typified
further, uniting the superficially dissimilar maritima and spinosa in one genus (her
P. P.
her splitting
named Apart from
iscus”) while leaving Asteriscus aquaticus in the other (which she Nauplius).
acceptance,
one deviating species as a monotypic genus Ighermia (Wiklund which has gained little
1985),
2003,
Greuter >
taxonomic conclusions, with the nomenclature has been generally accepted
rectified, (e.g.,
&
Asteriscus,
Greuter Raab-Straube 2006-2009). That treatment, including the repatriation of Ighermia to
received firm support by molecular studies (Goertzen
et 2002).
al.
._.
correct,
Back then, to the initial question: which name, Pallenis maritima or Asteriscus maritimus, is
^
wis
and
answer, as so often: depends. Accordingly, one follows the concepts of Toumefort (1700),
it if
a sri
correct—
recognize a single genus comprising the whole Asteriscus alhance, then A. maritimus is ^
made during
somewhat outmoded
cally tenable if choice. However, one honors the taxonomic progress
if
3+ and
last centuries maintain two genera, then maritima the preferred choice.
P. is
up 40
grows
woody perennial) that to
spreading
subshrub low,
Pallenis maritima a cushion-forming (a
is
along
Portugal
extending to
on Mediterranean region, S.
tall and about mwide. known from the Western
1 It is
maritima
(Wiklund 1985). Pallenis
and N. France
the Great Britain
Atlantic Coast, with casual occurrences in
occurs within the
and frequently
grows on along the coastline, it
sand dunes, and rocky shores primarily
cliffs, m
found dry grass s
NATURA 201 also
salt-spray zone Mucina 2003; Estrada et al. 1). It is
(Beckett 1993; 1997;
(Wiklund
and sandstone 1985).
marl,
including
ofNorth
Africa and and grows on limestone, clayey soils,
Spain,
cm
up 4
heads
flower to
golden-yellow
abundant
produces
In
spring through summer, maritima
early
P.
The and rnvo-
leaves
than the involucre.
longer
wide, with showy, about as long as or
finely toothed ray florets
and some-
glandular,
densely villous,
hcral P sparsely to
bracts of maritima narrowly obovate-spathulate,
are
four-sided to
and achenes are irregularly
what the
folded. The contain oxalate crystals,
receptacular scales
mantxma
Pallenis is
description).
Wiklund 1985 a detailed
for
subcylindrical and
densely strigose-hispid (see
A
photograph pro-
^own and sea daisy. is
beach daisy,
by common Mediterranean
the names
of gold coin,
ved
as
Figure
1.
Horticultural Use
and
water there-
irrigation
and highly saline
Poflenis maritima severe water stress
exhibits a high tolerance to
Owing diese proper-
to
2005).
I°re gardens (Rodriguez et al.
is useful for revegetation programs and xeroscape
&
&
Vogl
2004).
Vogl-Lukasser
h Yeo 2000;
t*es has been 1996; Walters
widely Europe (Brickell
' cultivated in
a drought-tolerant
by nursery trade as
marketed the
Htlfenis where
maritima
is
is a!so cultivated in California, it
Gardens San Marcos
indy and coastal gardens (Brenzel 2007; California 2012; Growers
soils
2012).
s posted on Dave’s Garden (2012) webpage confirm P. maritima is a hardy, drought-tolerant plant
for
Garden an forum where
1 southern California coastal gardens; Dave’s is on-line people share garden-
and information about growing or purchasing ornamental plants. Casual observations by
the
senior
maritima often planted in street-side gardens in the coastal communities of Laguna
licate P. is Beach
ort Beach, southern California.
dwide, escaped garden plants are one of the primary sources of non-native plant introductions, and
&
is no exception (Mack 2000; Reichard White 2001; Bell et al. 2007). Although the mode of intro-
a species can be difficult to identify, documenting whether or not the plant is intentionally cultivated
Dean
ommercially can be significant (Dehnen-Schmutz et 2005; et 2008). Because maritima
al. al. P.
be our escaped from gardens and commercial
to cultivated in area, likely residential landscape
it
become
to naturalized in seashore habitats along the southern California coast.
and
Salt Tolerance Specialized Seashore Habitats
Halophytes grow naturally in saline environments and have evolved various mechanisms cope with
to salinity
&
where
other plants cannot thrive (Breckle 2002; Parida Das 2005). Tolerance to highly saline exposure
soils,
and
to salt spray, periodic inundation are critical factors affecting vegetation and the establishment of non-na-
& &
many De Kuhn
tive species in habitats of coastal California (Barbour Jong 1977; Zedler 1997; Grewellet
&
al. 2007; Pickart Barbour 2007). Therefore, knowing whether or not a non-native plant is tolerant of saline
conditions can have important implications regarding potential invasive spread and the habitats might
its it
was
Pallenis maritima not identified as a halophyte in the databases of Aronson Menzel and Lieth
(1989),
(2003), or Yensen (2012). Experimental data, however, indicate it is tolerant of high salinity levels, with electri-
cal conductivity measurements ranging from 1.25 to 12.61 dS/m-1 during 150 days of treatment (Rodriguez et
minimum
al. 2005). For Aronson (1989), the criterion required for a plant to be classified as a halophyte is a
with an
salinity level electrical conductivity of 7-8 dS/m-1 during portions or of the
at least significant all
More
plant’s life cycle. recently, however, halophyte has become more demanding, and the
classification
mM
threshold of 200 Nad)
salinity tolerance for a plant to qualify as halophyte has been raised to 20 dS/m-1 (~
&
(Flowers Colmer 2008). For comparison, the salt concentration of seawater approximately equivalent©
is
mM
50 dS/m-1 500
(~ NaCl). See Grigore et al. (2010) for a review of the history and evolving concepts that
d-
Rodriguez
et al. (2005) also reported that salt-treated ulate high Na+ and
P. r
mechanism
and
osmotic adjustment that maintains leaf tugor in response The preferential accu-
to salt stress.
mulation Na+ may
of either and/or Cl- account, in part, for salt tolerance (Nieman et 1988; Gibbs etal. 1989;
al.
&
Boursier Lauchli 1990). Accordingly, maritima a and should be considered forindu-
P. is salt-tolerant species
sion in future revisions of the world-wide halophyte
database.
In Orange County, southern within the
California, P. maritima grows on rocky shores or sea cliffs, often
salt-spray zone and in habitats periodically inundated during storm At one also grows at the
surge. location,
it
edge an
of ephemeral
calcareous-saline The known California is
seep. distribution of maritima in southern
P.
depicted
in Figure
2.
Laboratory analysis using P maritima at
saturated extracts of soil samples taken within the root zone of
each
naturalized population to
indicate the substrate is slightly to moderately alkaline (7.5-8.1 pH), slightly
strongly the*
saline (6.5-34.7 dS/m-1), and of
slightly to highly calcareous. In addition, the micro-habitat
rocky
seashore environments halophyte5
is barren or covered with sparse vegetation, often with documented
such
as Cakile maritima Extripk*
Scop., Carpobrotus chUensis (Molina) N.E. Br, Greene,
Distichlis spicata (L.)
califomica (Moq.) E.H. Zacharias Hubb. There-
(syn., Atriplex califomica Moq.), and Limonium (Stapf)
perezii
fore, P. maritima functions as a halophyte here are
in coastal southern and occupies
California, the habitats
it
similar
to its seashore environment along
the Mediterranean
Sea.
625
and Greuter,
F
Riefrier
There new become estab-
is always some concern about the possible invasiveness of non-native plants that
hed and
lis in natural “casual alien or weed” for Great Britain is listed
ecosystems. maritima cited as a
Pallenis is
** 3 >tential weed known be invasive anywhere around the world
but not to
introduction” in Australia, is
(GCW
2012; ISSG Weeds Australia 2012).
PIER
2012; Randall 2007, 2012;
2012;
Wiklund pointing autoga-
produce viable seed, to
(1985) noted maritima plants kept in isolation
that
P.
®°
Us Orange
0r Huntington Beach, coastal
apomictic reproduction. Additionally, in 2003 a gardener in
Count
may become invasive (Dave’s
maritima slightly
y, California, posted a comment on-line indicating that P.
Garden and
2012). Accordingly, P. maritima possesses reproductive physiological traits that enable
it to escape
We
know
cultivation and become established in seashore environments. do not if it will become a pest
plant
in
North America, but it is expected to expand its range and establish elsewhere in coastal southern Califor
nia
many
Ornamental horticulture promotes invasions in ways, and the tradition of selecting and
distributing
showy
species purely for aesthetic reasons remains an important pathway of dispersal (Dehnen-Schmutz &
Touza 2008). Aesthetically-pleasing and easy-to-grow plants often become popular with
gardeners,
and
thereby frequently become available for sale in large numbers of nurseries and in expanding
horticultural
mar-
Market more
kets. availability drives prices, so the often a species is offered for sale, purchased, and
planted,
the greater chance escape and (Lockwood
its to cultivation find suitable natural habitats et 2005; Dehnen-
al.
&
Schmutz Touza make
2008). In addition, biological characteristics that a plant interesting and easy grow
to
for gardeners, such as climatic suitability, long blooming period, quick propagation, hardiness, and
drought-
may
tolerance, also enhance the successful establishment of escaped plants in new regions and an
start invasion
process (Mack Dehnen-Schmutz We
2000; Anderson
et al. 2005; et al. 2006). believe salt-tolerance in orna-
mentals, especially those selected and marketed
for coastal gardens, contributes to their ability to successfully
escape and become
cultivation established in unintended
habitats.
Pallenis maritima is one of a growing number of salt-tolerant, hardy, and attractive ornamental plants that
have
naturalized recently in southern California. Others include Lagunariapatersonia (Andrews) G. Don (Mal-
&
vaceae), Plecostachys
serpyllifolia (PJ. Bergius) Hilliard B.L. Burtt (Asteraceae), and several species of Umo-
nium
(Plumbaginaceae)
(Hill 2012; Keil 2012b; Preston 2012). Plecostachys and several Limonium
serpyllifolia
&
taxa are highly invasive species (Riefner Nesom BAEDN
2009;
2012).
Halophytes
are receiving substantial attention from botanists and agriculturists as potential fodder crops,
food
plants, biofuel sources,
turf grasses, ornamentals, sand dune stabilizers, and redeemers of salt-affected
among &
land, others (Glenn
et al. 1999; Barrett-Lennard et 2003; Rozema Flowers 2008; Yensen 2008). As
al.
numbers
greater
of become
salt-tolerant plants and meet
cultivated, as the horticultural trade continues to
consumers’ demands
hardy and
for we
attractive plants for coastal gardens, can expect find other non-native
to
halophytes
naturalized along
the California
coast.
ACKNOWLEDGMENTS
Edward
Glenn
P. (University of Arizona) and David
Keil (California Polytechnic State University, San Luis
J.
Obispo) provided comments
helpful We
that greatly improved the manuscript. appreciate the ef-
also greatly
Harvey
forts of Brenneise and
Holiman
Irene
(Library of Rancho Santa Ana Botanic Garden) for assistance
with document Gam
and
retrieval, to Wallace
(Wallace and
Laboratories) for helpful review of saline calcare-
ous
substrate
data.
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