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VOLUME 49, NUMBER 1 JANUARY-MARCH 2002 


EFFECTS OF FIRE ON THE REPRODUCTIVE BIOLOGY OF AGAVE PALMERI (AGAVACEAE) 


[ELE SSS OTIS DB resco SB Ee PE EE SI Re 1 
ADULT SEX RATIO OF ARCEUTHOBIUM GILLII (VISCACEAE) 
Robert L. Mathiasen and Carolyn M. Daugherty .............cccccccceeeeeeeveneeees 12 


ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE (POLYGONACEAE), A NEW VARIETY 
FROM THE SOUTHERN SIERRA NEVADA, CALIFORNIA 


| ROSE OING NOTE OO oN I RR eR TI Te a 16 
HESPEROYUCCA WHIPPLEI AND YUCCA WHIPPLEI (AGAVACEAE) 
Jeffrey A. Greenhouse and JON, StrOMOCB ecceccccccess-c00ceese0s.04000000nenaneees- 20 


SYMPATRY BETWEEN DESERT MALLOw, EREMALCHE EXILIS, AND KERN MALLow, 

E. KERNENSIS (MALVACEAE): MOLECULAR AND MORPHOLOGICAL PERSPECTIVES 

Katarina Andreasen, Ellen A. Cypher and Bruce G. Baldwin................. 22 
POLLINATION OF CYTISUS SCOPARIUS (FABACEAE) AND GENISTA MONSPESSULANA 

(FABACEAE), Two INVASIVE SHRUBS IN CALIFORNIA 

Ingrid M. Parker, Alexandra Engel, Karen A. Haubensak and 

Karen Goodell ....... ARE A PTS Bi os en scwecececcsecee DS 
FOxTAIL PINE IMPORTANCE AND CONIFER DIVERSITY IN THE KLAMATH MOUNTAINS 

AND SOUTHERN SIERRA NEVADA, CALIFORNIA 


Andrew J-Eekert and JOGA sO: SAWCT ie FB Ss ISP ne ono 00 ET wae 33 
NOTEWORTHY BRYOPHYTE RECORDS FROM THE MOJAVE DESERT 

Lloyd R. Stark, Alan T. Whittemore and Brent D. Mishler................00000+ 49 
COPNTETEGRNA, oo ss ccs Ee re cn scene ceccecettocss 54 
© TERN yea ses I sat soc oi Ue hc oa wadevounaneedcbboveSes 54 
IB YATS1 0) Sornse acer On cl ere ome mi 7A 220: SEEN ARR nEet12)\ 54 
AN TONGTONINVN es eso ce oe ook Ne Og ee ee ed ey ee ccateeu sche epee 55 
INTER RY DED ICC) ck yang tee ee Sen err eg 1061) 0 Ae Oe an 54 
CO) EE Oe en ate inc Sn BS NRE SM Fld Sec anuys aia uc tica seuunb suneentceuies 58 


THE MANZANITAS OF CALIFORNIA, ALSO OF MEXICO AND THE WORLD, BY 
Putte V. WELLS 
MuachaelC. Vasey Gnd WV ThOMGS POLK «.s50502.03ssccsssesesnecssssancteeestespunsiee 46 
ILLUSTRATED FIELD GUIDE TO SELECTED RARE PLANTS OF NORTHERN CALIFORNIA, 
EDITED BY GARY NAKAMUA AND JULIE KIERSTEAND NELSON 


TEATRO) eS CNN Cre 5 ee ee arate Des SEN CR co Sistah AS caidas iedtsis donut as Wee Uwaus 48 
Monocots III/Grasses IV .. RECEIVED BY: re ES fo ee 15 
FERRVATIUINI elo oeSiceressiccdeonton. estan: leartsias a 59 


INDEXING (2 4) 


PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY 


Maprono (ISSN 0024-9637) is published quarterly by the California Botanical Society, Inc., and is issued from the 
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Class of: 


2002—NorMAN ELLSTRAND, University of California, Riverside, CA 
Cara M. D’ Antonio, University of California, Berkeley, CA 
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Jon E. Kee.ey, U.S. Geological Service, Biological Resources Division, 
Three Rivers, CA 
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INGRID PARKER, University of California, Santa Cruz, CA 
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OFFICERS FOR 2001—2002 


President: Bruce BALDwin, Jepson Herbarium and Dept. of Integrative Biology, 1001 Valley Life Sciences Bldg. 
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The Council of the California Botanical Society comprises the officers listed above plus the immediate Past President, 
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Editor of Maprono; three elected Council Members: BiAN Tan, Strybing Arboretum, Golden Gate Park, San Fran- 
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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 


MADRONO, Vol. 49, No. 1, pp. 1-11, 2002 


EFFECTS OF FIRE ON THE REPRODUCTIVE BIOLOGY OF 
AGAVE PALMERI (AGAVACEAEB) 


Liz A. SLAUSON 
Scottsdale Community College, 9000 E. Chaparral Rd., 
Scottsdale, AZ 85256-2626 


ABSTRACT 


Fire is an important management tool that is used increasingly to restore natural composition, structure, 
and processes in semi-arid grasslands, woodlands, and forests of the southwestern U.S. I investigated the 
effects of fire on floral resources, fruit and seed set, and survivorship of Agave palmeri, an important 
food source of the endangered lesser long-nosed bat. Nectar production and sugar concentration, pollen 
and nectar standing crops, and fruit and seed production were measured in flowering plants with leaves 
damaged by fire and compared with unburned plants at three sites in two different human-ignited fires. 
In general, no significant differences were found in total nectar production, nectar sugar concentration, 
standing pollen crops, or fruit and seed set between burned and unburned plants. Standing nectar crops 
were slightly smaller than total nectar production amounts, but large amounts of nectar and pollen re- 
mained available at dawn in both burned and unburned plants. Initial mortality measured across all size 
classes at one site was only 3.3%. Although levels of burn damage relative to plant size were quite 
variable, plants with greater damage (61—100%) tended to be <0.6 m in height and diameter. These results 
indicate that fire did not appreciably decrease food resources of the lesser long-nosed bat or the repro- 
ductive resources and survivorship of A. palmeri. The rocky, low fuel habitats preferred by A. palmeri 
combined with certain morphological and physiological adaptations of the plant, such as a rosette shape 
and storage of accumulated carbohydrates in the center of the rosette, appear to protect the majority of 


stored resources within the plant’s center during fire events. 


Key words: Agave palmeri, fire, Leptonycteris curasoae yurbabuenae, lesser long-nosed bat. 


INTRODUCTION 


Agave palmeri Engelm. is a perennial, rosette- 
shaped leaf succulent, and is widespread in desert 
scrub, oak savanna, and oak woodland communities 
of the Southwest Borderlands: the international 
four-corners area of Arizona, New Mexico, Sonora, 
and Chihuahua. Prescribed fire is one management 
tool that is being tested to control the conversion 
of grasslands into shrubland communities, to re- 
duce high fuel loads in woodland communities, and 
to return communities of the Southwestern Border- 
lands to pre-settlement states (Allen 1996; Edmins- 
ter 1996). Although the pollen and nectar of A. pal- 
meri are major food sources of the endangered less- 
er long-nosed bat (Leptonycteris curasoae yurba- 
buenae) (Hayward and Cockrum 1971; Howell 
1972; Howell and Roth 1981; Slauson 2000), little 
is known about the effects of fire on A. palmeri, 
and in particular, on its production of food resourc- 
es for floral visitors. 

Agave palmeri has a prolonged juvenile period 
that may last 20—40 years, during which time water 
and carbohydrates accumulate in the leaves. Once 
maturity is reached, plants are monocarpic, flow- 
ering only once and then dying. Mature plants 
range in size from 0.5—1.6 m tall and 0.5—2.5 m 
wide with 56-124 leaves (Noble and Smith 1983; 
Slauson 2000). The inflorescence, or flowering 
Stalk, is a large, open panicle that varies from 2-8 
m in height with 10—30 umbels (flower cluster on 
side branch of inflorescence) (Slauson 1996; Hodg- 


son 1999), and produces on average 1500-2200 
flowers over the flowering season (Slauson 2000). 
Anthers are sizeable (8.7—20 mm in length) and 
produce large amounts of pollen if not removed on 
the night of dehiscence by bats, moths, or rain; pol- 
len is usually harvested the following morning by 
various bees (Slauson 2000). Flowers secrete nectar 
nocturnally over a 6-day flowering period and pro- 
duce an average of 2.5 ml of total nectar, with peak 
production on the second day (Slauson 1999, 
2000). These large quantities of pollen and nectar 
attract many animals (Slauson 2000), including the 
lesser long-nosed bat. Nocturnal visitors have the 
‘‘first crack’? at pollen and newly secreted nectar. 
The lesser long-nosed bat was federally listed as 
endangered in 1988 due to apparent low and de- 
clining numbers, disturbance of roosts, and an in- 
terdependence with its food resources (Shull 1988), 
but its status as endangered is controversial (see 
Cockrum and Petryszyn 1991). Several species of 
columnar cacti and agaves provide a “‘nectar cor- 
ridor’’ for the lesser long-nosed bats, from spring 
as they migrate north from Central America and 
Mexico, through fall when they return to southern 
roosts (Gentry 1982; Fleming et al. 1993). In late 
July and early August, substantial numbers of lesser 
long-nosed bats migrate to higher elevations in 
southeastern Arizona (Cockrum 1991) where their 
primary food source from July through September 
is A. palmeri (Howell 1972; Howell and Roth 
1981). In describing this mutualistic relationship, 


p) MADRONO 


Howell and Roth (1981) suggested that A. palmeri 
was also “‘strongly dependent’? upon the lesser 
long-nosed bat for pollination. More recently, Slau- 
son (2000) has shown that this mutualistic relation- 
ship is asymmetrical; bats depend on this agave 
species part of the year for food, but A. palmeri 
does not require bats for adequate sexual reproduc- 
tion. 

Episodic fires were common throughout most 
ecosystems of the Southwest Borderlands for at 
least 300—400 years prior to ca. 1900 (Baisan and 
Swetnam 1990; Swetnam and Baisan 1996a, b), in- 
cluding those occupied by A. palmeri. Kaib (1998) 
has shown desert grasslands burned approximately 
every 5—10 years, and ranged between 10 to hun- 
dreds of km? in size, while canyon pine-oak forests 
burned every 5—9 years and covered areas of at 
least 50 km’. A drastic reduction of these wide- 
spread, episodic surface fires occurred north of the 
border between ca. 1870—1900, initially as a result 
of intensive grazing that decreased fuels and the 
ability of fire to spread across large landscapes 
(Bahre 1991; McPherson et al. 1993; Allen 1996; 
Swetnam and Baisin 1996a, b). Continued grazing, 
land use, and fire suppression practices by govern- 
ment agencies throughout the 20th century resulted 
in the virtual extinction of natural disturbance fires 
in southwest ecosystems (Bahre 1991; Allen 1996; 
Swetnam and Baisan 1996a). These practices have 
contributed to many changes in both the structure 
and function of these communities, ranging from 
shrub invasion of desert grasslands due to a lack of 
fine fuels to carry low intensity fires (Hastings and 
Turner 1965; Humphrey 1987; McPherson 1995), 
to severe risk of catastrophic and stand-replacing 
wildfires in forests due to great fuel accumulations 
(Cooper 1960; Covington and Moore 1994). 

Many succulents tolerate fire to some degree 
(Thomas and Goodson 1992), but desert grassland 
fires have also been reported to kill succulent plants 
(Niering and Lowe 1984; Nobel 1988), particularly 
the smaller size classes. McLaughlin and Bowers 
(1982) reported that large succulent plants that ini- 
tially survived fires had increased mortality in later 
years. Several other factors besides plant size may 
affect the responses of succulents to fire including 
the size and patchiness of the fire, the time of year 
fire occurs, fuel conditions, weather conditions, to- 
pography, and plant morphology and physiology 
(McPherson 1995). Little data are available regard- 
ing the effects of fire on agaves. In simulated grass- 
land fire experiments, A. gigantensis had 0% mor- 
tality after 14 months with fuel densities up to 1600 
g m ~~? and temperatures of 400—600°C (these tem- 
peratures are normally expected in intense grass- 
land fires) (Thomas and Goodson 1992). Although 
the leaf tips collapsed at fuel densities of 400 g m ~~ 
and damage increased as fire intensity increased, 
the center of the rosettes remained unburned. In 
field surveys of semi-desert grassland sites in 
southern Arizona that had burned in wildfires in the 


[Vol. 49 


prior 18 months, mortality from fire was 18% in A. 
palmeri (Thomas and Goodson 1992). Of the re- 
maining living plants, 83% exhibited regrowth from 
the apical meristem, whereas 17% survived un- 
burned in refugia. These refugia were created by 
either the patchy nature of the fire, which skipped 
Over areas with adequate fuels, or by rocky areas 
with little flammable material. 

The prolonged juvenile period of agaves results 
in large amounts of stored resources that are used 
for flower (including nectar and pollen), fruit, and 
seed production. Once flowering is initiated, a large 
and irreversible translocation of stored resources 
occurs from the rosette to the developing inflores- 
cence. Death results presumably because resources 
normally reserved for growth and maintenance are 
mostly allocated to reproduction. Nobel (1977) ob- 
served that A. deserti Engelm. diverted over 68% 
of its stored biomass to a developing inflorescence. 
No data are available regarding how the loss of 
stored resources from fire damage in reproductive 
agaves affects nectar and pollen production or sub- 
sequent fruit and seed set. 


Research questions. 1 studied aspects of the floral 
biology of Agave palmeri relevant to nectar bat vis- 
itation immediately following two summer fires. 
These fires, one prescribed and one accidental, 
burned patchily and caused little immediate mor- 
tality of mature agave plants. Plants that were re- 
productive the year of each fire had partially or ful- 
ly emerged inflorescences at the time of the fires, 
and either were flowering or flowered soon after. I 
compared reproductive burned plants with un- 
burned plants and asked (1) did nectar production 
rate or concentration differ between the two treat- 
ment classes? (2) What was the standing crop of 
nectar and pollen at dawn, after possible bat visi- 
tation? (If floral rewards of populations were close- 
ly cropped by bats, then an adverse effect of burn- 
ing on nectar production would be important. A 
surplus in both nectar production and standing 
crops would indicate that floral foods were not in 
limited supply for nocturnal visitors. Differences 
between burned and unburned plants could indicate 
production differences or discrimination by noctur- 
nal visitors.) (3) Did fruit and seed set differ be- 
tween burned and unburned plants? (A significant 
decrease in fruit or seed set could have considerable 
impacts on future floral resources for bats.) (4) Was 
burn damage of plants related to their size, specif- 
ically the height and diameter of the rosette? 


METHODS 


Study sites. Fieldwork was conducted at two dif- 
ferent fire sites, a prescribed burn and an accidental 
human-ignited fire. The prescribed burn (known as 
the Maverick Burn) was conducted on June 24—25, 
1997 by the U.S. Forest Service on the southern 
edge of the Peloncillo Mountains (Fig. 1). Approx- 
imately 8000 acres burned in a mosaic pattern with- 


2002] 


Arizona 


New Mexico 


Peloncillo Mtns. 
Animas Mts. 


Chihuahua 


Sonora 


Fic. 1. Locations of CE GT (Maverick Burn), and MF 
(Gray Ranch) study sites. 


in the 17,000 acres designated as the primary burn 
area (Encinas 1997). Two study sites were chosen 
within the primary burn area that represented typ- 
ical habitats for A. palmeri (Gentry 1982). The first 
study site was located in the southern portion of the 
burn area on Cowboy Flats (CF site, lat 31°26'N, 
long 109°2'W, elevation ~1585 m) near a small 
lesser long-nosed bat roost. This site was on a 
rocky, south-southwestern facing hillside and mesa 
top in an oak savanna community. Plant species 
composition included Quercus emoryi, Q. oblon- 
gifolia, Juniperus monosperma, Prosopis velutina, 
Yucca schotti, Nolina microcarpa, Dasylirion 
wheeleri, Fouquieria splendens, Calliandra sp., 
Gutierrezia sarothrae, Bouteloua gracilis, B. cur- 
tipendula, B. hirsuta, B. radicosa, Hilaria belan- 
geri, Erogrostis intermedia, Muhlenbergia sp., Ly- 
curus phleiodes, Aristida spp., Schizachrium cir- 
ratum, and Heteropogon contortus. The second 
study area was located along the northern border of 
the burn area along Geronimo Trail (GT site, lat 
31°32’N, long 109°2'W, elevation ~1675 m), and 
was characterized by flat to steep (0O—45°) slopes 
with south-southwestern exposures and rocky soils 
in an oak savanna/oak woodland community. Plant 
species included Q. emoryi, Q. oblongifolia, J. 
monosperma, J. deppeana, P. velutina, Arctostaph- 
ylos pungens, Y. schotti, G. sarothrae, B. gracilis, 
B. curtipendula, B. hirsuta, B. radicosa, H. belan- 
geri, E. intermedia, Aristida spp., S. cirratum and 
M. emersleyi. 

The accidental human-ignited fire started in 
Mexico in May 1999, crossed the international bor- 
der, and burned approximately 22,000 acres in a 
mosaic pattern over a 2-day period on the Gray 


SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 3 


Ranch in southeastern New Mexico (Fig. 1). The 
third study site was located on the Gray Ranch near 
McKinney Flats (MF site, lat 31°23’N, long 
108°42’'W, elevation ~1570 m) on a south-facing, 
rocky slope in a semi-arid grassland/oak savanna 
community. Plant species included Q. emoryi, Q. 
oblongifolia, P. velutina, N. microcarpa, Juniperus 
sp., Cylindropuntia spinosior, G. sarothrae, B. 
gracilis, B. curtipendula, B. hirsuta, B. radicosa, 
H. belangeri, Aristida spp., and S. cirratum. Agave 
populations (all size classes excluding seedlings) at 
study sites ranged from approximately 1000—3000 
plants per ha. 

Although several plots were identified as poten- 
tial study sites prior to the Maverick fire, ignitions 
were initiated across broad landscape areas rather 
than igniting individual study plots, and none of the 
potential study plots burned. Therefore, thorough 
pre-treatment characterization of plots was not pos- 
sible. Due to the large areas of steep and inacces- 
sible terrain, the mosaic pattern of the fire, the 
patchy distribution of agaves, and limitations of 
sampling nectar and pollen with 12-foot ladders, 
only two suitable sites that actually burned could 
be located within the primary burn area. At these 
sites, fire behavior was characterized by partial to 
complete understory consumption (Clark 2000). 
The Gray Ranch fire was accidental, and only one 
accessible agave population burned. 

Research was conducted at both fire sites during 
late July and August during the peak flowering pe- 
riod of A. palmeri and when migrating lesser long- 
nosed bats are normally present in southeastern Ar- 
izona. Work was conducted August 4—9, 1997 at 
the CF site, August 11—15, 1997 at the GT site, and 
July 20—22 and August 2—4, 1999 at the MF site. 


Nectar and pollen studies. 1 studied nocturnal 
nectar production in bagged, or exclosed, flowers 
to determine whether this important floral reward 
was affected by fire. I also measured standing crops 
of nectar and pollen at dawn in unbagged, or open, 
flowers. Standing crops reflect both the effects of 
harvesting by animals and production of floral re- 
wards by flowers. Plant selection was limited to 
plants whose flowers could be reached with 12-foot 
ladders in burned and unburned areas at each site. 
A plant was classified as a “‘burned plant” if >1% 
of the rosette had tissue death due to fire. Rosette 
height, width, and number of umbels were mea- 
sured for each study plant. For study plants in 
burned areas, burn damage (% of rosette damaged 
by fire) was scored from 0 to 6: 0 = no burn dam- 
age, 1 = 1-20%, 2 = 21-40%, 3 = 41-60%, 4 = 
61-80%, 5 = 81-—100%, and 6 = entirely dead. 

Sample sizes for nectar and pollen studies at all 
sites were restricted due to the mosaic pattern of 
the fire, the patchy distribution of agaves on pri- 
marily inaccessible and rocky terrain, and the num- 
ber of plants in flower that could be reached with 
12-foot ladders. In general, all plants that could be 


4 MADRONO 


reached at a site were sampled. Total nectar pro- 
duction, nectar sugar percentage, and standing nec- 
tar and pollen crops were measured on six plants 
each in burned and unburned plots at the CF site 
and nine plants at the unburned GT site. Flowers 
on most plants in the burned GT plot were too high 
to reach with ladders, and only 2—3 plants could be 
sampled, depending on the day and appropriate 
flower stage availability. At the MF site, 20 and 10 
plants were measured in the burned and unburned 
plots, respectively. To decrease any positional ef- 
fects of nectar volume, nectar sugar concentration 
or standing crops, experimental flowers were locat- 
ed on umbels positioned in the middle section of 
the inflorescence, whenever possible. 

For studies of total nectar production and nectar 
sugar concentration at the CF and GT sites, 5-10 
predehiscent (day 1 of 6-day flowering period) 
flowers on one umbel were numbered, enclosed 
with a nylon mesh bag prior to dusk, and nectar 
was allowed to accumulate until dawn (nectar is 
produced only during the night). Anthers of pre- 
dehiscent flowers generally split open and dehisced 
pollen shortly after dusk, entering the dehiscent 
(day 2) floral stage. Dehiscent stage flowers were 
used in nectar and pollen experiments as nectar pro- 
duction is greatest in dehiscent flowers, and pollen 
is only available during the dehiscent stage (Slau- 
son 2000). Plants were sampled for three successive 
days for total nectar production studies (different 
flowers sampled each day) and one day for nectar 
sugar concentration studies. Nectar production was 
measured by withdrawing the nectar present in the 
floral tube at dawn with a tuberculin syringe and 
blunt end needle. Nectar sugar concentration (per- 
cent sucrose equivalents on a weight/weight basis) 
was measured in the field at dawn with a hand-held 
refractometer. Data collection methods for total 
nectar production and nectar sugar concentration 
were modified for the MF site in an attempt to in- 
crease the number of plants sampled for compari- 
son to standing nectar crops (see below). Plants 
were sampled one day only and data collection was 
spread over a 3-day period (4—7 plants per treat- 
ment were sampled each morning at dawn). To de- 
termine if nectar resources decreased significantly 
over time, total nectar production and nectar sugar 
concentration measurements were repeated on the 
same study plants two weeks later (August 2—4, 
1999). Due to the difference in sampling method- 
ology, fire sites were analyzed separately. 

To indirectly determine the degree of nectar use 
by nocturnal floral visitors (bats and moths), stand- 
ing nectar crops (amount of nectar present at dawn 
in flowers left available to visitors) were measured 
and compared to total nectar production (exclosed 
flowers) at all three study sites. In order to most 
accurately measure dawn standing crops, sampling 
was begun as close to dawn as possible (<one 
hour), and was completed before dawn when an 
abundance of bees and other diurnal animals ap- 


[Vol. 49 


peared that could substantially decrease standing 
nectar amounts. Standing crops were measured on 
umbels adjacent to those used for studies of total 
nectar production and nectar sugar concentration. 
Nectar volume was sampled on three successive 
days and nectar concentrations were sampled one 
day on 5—10 dehiscent flowers per plant at the CF 
and GT sites. Methodology differed at the MF site; 
standing nectar crops and nectar sugar concentra- 
tions were measured on 4—7 plants (five dehiscent 
flowers per plant) each morning at dawn during the 
first sampling period (July 20—22). Nectar and nec- 
tar sugar concentrations were measured as previ- 
ously described. 

Standing pollen crops (amount of pollen present 
at dawn in flowers left available to visitors) were 
measured using a qualitative 5-class index at all 3 
sites to estimate pollen resource use by nocturnal 
floral visitors. Standing pollen crops were measured 
by evaluating the amount of pollen present at dawn 
and scoring pollen from 0 to 5 (O = no pollen pre- 
sent, ..., 5 = all or large amounts of pollen pre- 
sent). At the CF and GT sites, standing pollen crops 
were measured on umbels adjacent to those used 
for total nectar production and nectar sugar con- 
centration studies. Five to ten dehiscent flowers per 
plant were sampled for three successive days (dif- 
ferent flowers sampled each day). At the burned GT 
site, low numbers of available flowers and inability 
to reach flowers with ladders allowed sampling on 
only two plants (one plant sampled for one day and 
one plant sampled for two days). Methodology dif- 
fered at the MEF site; plants were sampled one day 
only and data collection was spread over a 3-day 
period (4—7 plants per treatment were sampled each 
morning at dawn). 


Fruit and seed set studies. To determine whether 
fire significantly effected fruit and seed set of aga- 
ves, 12 plants each from burned and unburned sites 
at the CF and GT sites, and 20 plants from the 
burned and 22 plants from the unburned MF site 
were randomly selected by a coin toss after the fire. 
Plants were allowed to be open-pollinated, stalks 
were cut down, fruits were collected in October and 
November, and mean percent fruit and seed set for 
each inflorescence were calculated. To determine 
fruit set, the total number of mature fruits and 
aborted flower scars on each inflorescence were 
counted, and percent fruit set was calculated (num- 
ber of fruits/(number of fruits + aborted flower 
scars)). Mean seed set per plant was determined by 
placing all fruits of an inflorescence in a paper bag, 
randomly selecting 20 fruits, and calculating seed 
set for each fruit (number of black, fertilized seeds/ 
(number of black, fertilized seeds + number of 
white, unfertilized ovules)). Capsules had dehisced 
in several inflorescences prior to fruit collection, so 
seed set could only be measured on only six plants 
each in burned and unburned plots at the CF site, 
six plants in the burned GT plot, eight plants in the 


2002] 


TABLE 1. 


SLAUSON: FIRE EFFECTS ON AGAVE PALMERI = 


MEAN PLANT SIZE AND BURN DAMAGE +1 SE OF REPRODUCTIVE A. PALMERI USED IN NECTAR AND POLLEN 


STUDIES AT CE GT, AND MEF Sires. RH = rosette height (m), RW = rosette width (m), BD = burn damage estimate 
(O = no burn damage, | = 1—20% of rosette burned, 2 = 21-40%, 3 = 41-60%, 4 = 61-80%, 5 = 81-100%, 6 = 
dead). Values with different superscripts were significantly different (P < 0.02) within study sites. 


Site RH (m) 
Burned CF 0.93 + 0.04 
(eo) 
GT 1.13 + 0.092 
Gi) 
MF 0.79 + 0.04 
(a = 14) 
Unburned CF 0.88 + 0.05 
(n = 6) 
GT 0.82 + 0.05? 
(ee) 
MF OMT OWS 
(n = 10) 


unburned GT plot, 15 plants in the burned MF plot, 
and 16 plants in the unburned MF plot. 


Mortality and demography studies. To examine 
agave demography and mortality from fire, four 20 
x 50 m plots were established within the burned 
area at one fire site (MF site) after the fire. Fire 
severity appeared relatively uniform within the 
burned area, and was not considered in plot loca- 
tion. Azimuth (degrees) and direction (m) to the 
plots from the road that divided the burned and un- 
burned areas were selected from a random numbers 
table. All plants within plots were tagged, rosette 
height and diameter were measured, survivorship 
and reproductive status noted, and degree of burn 
damage (as previously described) recorded. 


Data analysis. Total nectar production, standing 
nectar crop, and exclosed nectar sugar concentra- 
tion data had normal distributions, variances, and 
residuals at all three study sites, and were analyzed 
by one-way repeated measures ANOVA (plants = 
repeated measure). Sample size in burned plants for 
total nectar production and nectar sugar concentra- 
tion studies was reduced to 14 as six plants were 
either finished flowering or flowers could not be 
reached during the second sampling period (August 
2—4). Standing nectar crop data for the burned GT 
plot were excluded from the analysis as no flowers 
were available for measurement on several plants 
on various days. Due to the small number of plants 
with paired data values for standing nectar sugar 
concentrations (n < 4 plants per treatment) for the 
CF and GT sites, data were analyzed by the Proc 
Mixed procedure of SAS (SAS 1999) which uses 
maximum likelihood estimation procedures to han- 
dle missing data in repeated measures analysis. 
Fruit and seed set data had normal variances, dis- 
tributions, and residuals, and were analyzed by one- 
way ANOVA. All statistical analyses were per- 
formed using SYSTAT 9.01 (SPSS Inc. 1999) un- 
less otherwise noted. 


RW (m) BD 
1.48 + 0.10 2.6 + 0.6 
1.46 + 0.09 333) =) 1.2 
1.19 = 0:08 oy B= 10 EY 
1.32 SOM — 
125 S=0109 = 
1.19 + 0.08 — 

RESULTS 


Burn damage and size of nectar plants. Damage 
to burned agaves used for nectar studies ranged 
from a mean score of 1.6 (1—20% of rosette burned) 
at the MEF site to 3.3 (41-60% of rosette burned) 
at the GT site (Table 1). Differences between sites 
were not significant (Kruskal-Wallis test, K/W = 
3.14, P = 0.206), but sample size was low for the 
GT site (n = 3). In general, plants from the MF 
(Gray Ranch) site were smaller than those from the 
CF and GT sites. Burned plants were significantly 
taller than unburned plants at the GT site (one-way 
ANOVA with Bonferroni adjustment, F, ,) = 7.614, 
P = 0.02). Number of umbels per inflorescence 
ranged from 16—23 among sites. 


Nectar, nectar sugar concentration, and pollen 
production. Mean nectar production did not differ 
greatly between burned and unburned plants, but 
results were variable between sites, and occasion- 
ally between plants. Total nectar production was 
significantly higher in burned plants at the CF site 
(one-way repeated measures ANOVA, F; 4 
6.366, P = 0.03) (Fig. 2a), and a significant linear 
decrease in nectar production was observed over 
the three-day period (one-way repeated measures 
ANOVA, F; 19 = 7.740, P = 0.019). At the GT site 
where burn damage was greater (3.3, or 41—60% 
of rosette burned), mean nectar production was sig- 
nificantly lower in burned plants (one-way repeated 
measures ANOVA, F,, = 19.184, P = 0.005) (Fig. 
2b), however, sample sizes were very small for the 
burned site (n = 3 plants). Nectar production was 
also observed to significantly decrease linearly over 
the three-day period (one-way repeated measures 
ANOVA, F,, = 11.040, P = 0.016). At the MF 
site, no significant differences in total nectar pro- 
duction were found between burned and unburned 
plants during either sample period (July 20—22 and 
Aug 2-4, Fig. 2c) (one-way repeated measures AN- 
OVA, F,» = 0.662, P = 0.425), although nectar 


Total nectar production (ml) Total nectar production (ml) 


Total nectar production (ml) 


a) CF site 


Burn treatment 


@ burned 
4 unburned 


Jul 20-22 
Date 


Aug 2-4 


MADRONO [Vol. 49 


production did decrease slightly over time, but not 
significantly (one-way repeated measures ANOVA, 
ley = Dy IP = ODS). 

Dawn nectar sugar concentrations of exclosed 
flowers were significantly higher at the CF site 
(mean + SE = 20.6 + 0.9%, range 17—27%) than 
the GT site (mean + SE = 17.5 + 0.4%, range 
15.5-19%) (one-way ANOVA, F,, = 7.02, P = 
0.029). Although burned plants (mean + SE = 19.8 
+ 1.3%) tended to have slightly higher sugar con- 
centrations than unburned plants (mean + SE = 
18.7 + 0.7%), differences were not significant 
(one-way ANOVA, F,, = 0.56, P = 0.475). At the 
MEF site, dawn nectar sugar concentrations of ex- 
closed flowers ranged from 12.7—18.8% (Jul 20—22 
mean + SE = 16.1 = 1.2%, Aug 2—4 mean = SE 
= 14.6 + 1.7%). Sugar concentrations were not sig- 
nificantly different between burn treatments (one- 
way repeated measures ANOVA, F,,, = 0.585, P 
= 0.452), but decreased significantly on the second 
sampling date (Aug 2—4) (one-way repeated mea- 
sures ANOVA, F,,, = 13.580, P = 0.001). 

Pollen levels were not observed to vary between 
sites, burn treatment, or over time at all three study 
sites. Only one burned plant at the GT site and one 
unburned plant at the CF site were observed to dif- 
fer from a pollen score of 5, and then only for one 
sampling time. 


Standing nectar and nectar sugar concentration 
crops vs. total nectar production. Standing nectar 
crops were lower than total nectar production, but 
significant amounts of standing nectar (>0.54 ml) 
were available at dawn at all sites. A significant 
interaction was present in nectar production be- 
tween the burn treatment and standing nectar crop 
vs. total nectar production at the CF site (one-way 
repeated measures ANOVA, F,,) = 5.765, P = 
0.037), and as nectar production was averaged 
across the burn treatment, it was not advisable to 
interpret the standing crop vs. total nectar produc- 
tion test. Data were subsequently analyzed by run- 
ning paired t-tests on standing nectar crop and total 
nectar production data by burn treatment. Standing 
nectar crop (mean + SE = 0.565 + 0.033 ml) was 
significantly lower than total nectar production 
(mean + SE = 0.662 + 0.027 ml) in burned plants 
(t-test, ¢ = —1.767, P = 0.017, alpha/2 (0.025) to 
maintain Type I error), but no significant difference 
was found in nectar production between standing 
nectar crop (mean + SE = 0.546 + 0.029 ml) and 
total nectar production (mean + SE = 0.558 = 
0.028 ml) in unburned plants (t-test, t = —0.244, P 
= 0.646, alpha/2 (0.025) to maintain Type I error). 
At the GT site, no significant differences in nectar 


< 


Fic. 2. Mean total nectar production (ml) of dehiscent 
A. palmeri flowers in burned and unburned plots, (a) CF 
site, (b) GT site, and (c) MF site. Vertical lines = 1 SE. 


2002] 


production were found between standing nectar 
crop (mean + SE = 0.612 + 0.031 ml) and total 
nectar production (mean + SE = 0.711 = 0.018 
ml) in unburned plants (one-way repeated measures 
ANOVA, F,, = 1.101, P = 0.371) or in nectar 
production between days (one-way repeated mea- 
sures ANOVA, F,, = 3.927, P = 0.081). Data for 
standing nectar crop and total nectar production in 
burned plants were not analyzed due to low sample 
size (n = 2) and missing paired data values (flowers 
were not available for both open and exclosed nec- 
tar production on all three nights for the three plants 
with flowers that could be reached). At the MEF site, 
no significant differences in nectar production were 
found between burned and unburned plants (one- 
way repeated measures ANOVA, F)5, = 2.764, P 
= (0.108) or standing nectar crop and total nectar 
production (one-way repeated measures ANOVA, 
Fi53 = 0.791, P = 0.381) (standing crop mean + 
SE = 0.515 + 0.038 ml, total nectar production 
mean + SE = 0.557 + 0.042 ml). 

Nectar sugar concentrations of standing nectar 
crops ranged from 16.7—29.6% at the CF site and 
16.2—21.9% at the GT site. Nectar sugar concentra- 
tions were significantly higher at the CF site (Proc 
Mixed procedure, F,5, = 7.11, P = 0.015) and in 
standing crop flowers at both sites (Proc Mixed pro- 
cedure, F',, = 8.70, P = 0.021) (CF site: standing 
crop mean + SE = 21.8 + 1.1%, total nectar pro- 
duction mean + SE = 20.6 + 0.9%; GT site: stand- 
ing crop mean + SE = 19.2 + 0.6%, total nectar 
production mean + SE = 17.5 + 0.4%). At the MF 
site, standing nectar sugar concentrations ranged 
from 12.6-19.5%. No significant differences in 
nectar sugar concentrations were found between 
burned (mean + SE = 15.8 + 0.3%) and unburned 
plants (15.0 + 0.5%) (one-way repeated measures 
ANOVA, F453, = 2.220, P = 0.147) or between 
standing nectar crop (mean + SE = 15.6 + 0.3%) 
and total nectar production (mean + SE = 15.5 + 
0.2%) (one-way repeated measures ANOVA, F, 5. 
= 1.167, P = 0.289). 


Fruit and seed set. Burned and unburned plants 
had similar levels of fruit (17-22%) and seed set 
(19-23%) (Fig. 3). No significant differences were 
found in fruit set between study sites (one-way AN- 
OVA, Fy., = 1.915, P = 0.154). Fruit set was 
somewhat lower in burned plots, but not signifi- 
cantly (one-way ANOVA, F,,, = 0.294, P = 
0.589). Seed set was slightly higher in two of the 
three burned plots, but differences were not signif- 
icant (one-way ANOVA, F\ 45 = 0.641, P = 0.427). 
Seed set at the CF site was significantly lower than 
the GT and MEF sites (one-way ANOVA, Bonfer- 
roni test, P < 0.05). 


Mortality and demography. Overall mortality of 
plants in demography plots measured two months 
after the Gray Ranch fire (MF site) was 3.3% (Plot 
1 = 4.6%, n = 194, Plot 2 = 2.5%, n = 355, Plot 
3 = 3.9%, n = 276, Plot 4 = 3.0%, n = 459). Mean 


SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 7 


50 


45 Population 


40 & CF 
35 &@ GT 
O MF 


30 
25 
20 
15 
10 


Fruit set (%) 


burned unburned 
Treatment 


a- 4 
| 


burned unburned 
Treatment 


Seed set (%) 
AAS 
(eo) 


Fic. 3. Mean fruit and seed set (%) of A. palmeri in 
burned and unburned plots, CE GT, and MEF sites. Vertical 
lines = 1 SE. 


burn damage + SE was 3.1 + 0.03 (41-60% of 
rosette damaged) (range of plots = 3.4 + 0.06 to 
2.7 + 0.05) with approximately 37% of plants fall- 
ing into this damage class (Fig. 4). Burn damage 
classes 4 (22% of plants) and 2 (16% of plants) 
were the next most frequent damage classes. A 
multinomial logit model (Agresti 1990) relating 
burn class damage to rosette height and width cor- 
rectly predicted damage class for only 25% of ob- 
servations. The variability in this relationship is 
shown in Figure 5. Although median plant size 
across all levels of burn damage did not differ 
greatly, the range was quite variable. Rosette height 
and width values associated with burn damage clas- 
ses 23 were more concentrated than those of burn 
damage classes 1-2. 

Rosette height and width size class distributions 
illustrate the relatively young age of the population 
(Fig. 6). Greater than 90% of the population was 
<0.3 m in height and width. 


8 MADRONO [Vol. 49 
600 1000 
900 
0.4 0.7 
500 
800 
0.6 
2 a 20 ou 
= 400 age & 05% 
row = co 600 2 
° 300 S ° 500 0.4 8 
(<b) oO 
2 eae ee ao 03 2 
2 200 > z= 300 o> 
01 = Ons 
100 aut 
100 0.1 
0 0 0.0 
0 1 20 40 60 80 dead 0.0 0.1 02 03 04 05 06 0.7 08 09 1.0 


Burn damage (%) 


Fic. 4. Burn damage (% of rosette damaged by fire) of 
A. palmeri at the MEF site. 0 = no burn damage, 1 = 
1-20%, 2 = 21-40%, 3 = 41-60%, 4 = 61-80%, 5 = 
81—100%, 6 = dead. 


DISCUSSION 


Burning did not appear to negatively impact the 
production of nectar in A. palmeri. Mean total nec- 
tar production for all sites and times sampled (range 
+ SE = 0.46—-0.71 + 0.5 ml) was similar to or 
greater than previous reports of nectar production 


Rosette height 
Rosette width 


Burn damage 


0 0.5 1.0 1.5 
Rosette height and width (m) 


Fic. 5. Relationship between plant size (m) and burn 
damage (% of rosette damage by fire) of A. palmeri, MF 
site. 0 = no burn damage, 1 = 1—20%, 2 = 21-40%, 3 
= 41-60%, 4 = 61-80%, 5 = 81-100%, 6 = dead. Lower 
box plot of each pair indicates rosette height, upper box 
plot indicates rosette width. Length of box plot = range 
within 50% of values fall, center notch = median, box 
edges = first and third quartiles, whiskers = values within 
the first quartile — (1.5 X midrange) and the third quartile 
+ (1.5 X midrange), asterisks = values between first quar- 
tile — (1.5 to 3.0 X midrange) and third quartile + (1.5 
to 3.0 X midrange), and open circles = values <first quar- 
tile — (3.0 X midrange) and >third quartile + (3.0 X 
midrange). 


Rosette height (m) 


Number of plants 
Jeg Jed uoimodold 


OH) 0:1 0 OL OF Oks OLS (0.7 O13 O.2 710 


Rosette width (m) 


Fic. 6. Size class distribution of A. palmeri, MF site. n 
= 1298 plants. 


in dehiscent flowers of unburned plants (Howell 
1979; Slauson 1999, 2000). The significantly lower 
nectar production of burned plants at the GT site 
may be a result of sampling error due to the low 
sample size (n = 3 plants). Nectar sugar concentra- 
tions were not significantly different between 
burned and unburned plants at any site, and values 
were well within the range reported in previous 
studies of unburned plants (Howell 1979; Slauson 
1999, 2000). Nectar production and nectar sugar 
concentrations decreased over time in both burned 
and unburned plants, and this trend suggests that 
nectar resources may naturally decrease during the 
flowering period as stored resources are depleted 
and fruit and seed production increase. Pollen pro- 
duction did not appear to be affected adversely as 
large amounts of pollen were present on anthers at 
dawn at all sites and sampling times. 

Standing nectar crops at dawn were smaller than 
total nectar production in all treatments at all sites 


2002] 


(only significantly lower at the burned CF site), pre- 
sumably as a result of nectar use by moths and bats. 
However, at all study sites large amounts of nectar 
(>0.54 ml) and pollen (score = 5) were available 
in standing crop flowers at dawn, and indicate that 
food resource availability was not a limiting factor 
for either nocturnal (bats or moths) or diurnal vis- 
itors at any study sites during the times sampled. 
Standing crop results were similar to previous re- 
ports from other populations (Slauson 2000). 

The study sites examined did not appear to con- 
tain the fuel loads required to support a fire of the 
intensity needed to kill mature (reproductive) aga- 
ves or cause significant initial mortality when all 
size classes were considered. Although the majority 
of the population was quite small in size at the MF 
site (Fig. 6), mortality was <4%. Mean burn dam- 
age of mature plants was moderate at all sites (ap- 
proximately 10—43% of rosette burned), and in no 
case did mean damage exceed class 3 (41-60% of 
rosette burned). Relatively few large, non-repro- 
ductive plants (>0.6 m in height or diameter) had 
burn damage >60% (Fig. 5), and mean burn dam- 
age of all size classes at the CF site was approxi- 
mately 43%, despite the fact that over 90% of the 
population was <0.3 m in height and diameter. 

The low to moderate burn damage and high ini- 
tial survivorship of agaves may be due in part to 
their preferred habitats. Agave populations of mod- 
erate to high densities are characteristically found 
in xeric sites with rocky soil surfaces where com- 
petition with other plants is low (Gentry 1982). 
These rocky, low fuel habitats may serve as a re- 
fugia from fire for some agaves, especially smaller 
agaves which may be protected near the bases of 
surface rocks and cobbles. Plants located in less 
rocky, denser grassland or woodland habitats with 
increased fuel loads would presumably sustain 
greater damage. The greater burn damage observed 
in plants at the GT site (41-60%) may have been 
due to larger amounts of available fuels present in 
the surrounding oak savanna and oak woodland 
community. Robinett and Barker (1996) noted that 
frequent seedling establishment of A. palmeri oc- 
curs around dried, dead adult plants, and fuel loads 
created by dead adults can produce intense heat in 
a fire that kills surrounding seedlings. The vari- 
ability in the fuel loads of individual agave micro- 
habitats most likely explains the unpredictability of 
burn damage as a function of plant size observed 
in this study, especially in the smaller size classes 
(Fig. 5). 

Certain morphological and physiological adap- 
tations may also lessen the effect of fire on agaves. 
A number of agave species may resprout from rhi- 
zomes after fire (Gentry 1972), although most pop- 
ulations of A. palmeri are not rhizomatous (Slauson 
personal observation). Agaves may also benefit 
from the release of nutrients after a fire due to their 
Shallow root system (Gentry 1982). The rosette 
form of agaves to some degree protects the apical 


SLAUSON: FIRE EFFECTS ON AGAVE PALMERI 9 


meristem and the majority of stored resources lo- 
cated in the center of the plant. Reproductive aga- 
ves may have an even greater advantage: beginning 
in late winter and early spring, stored water and 
carbohydrates in reproductive agaves are translo- 
cated from the leaves to the center of the rosette 
where they are utilized for the development of the 
inflorescence (Nobel 1977; Tissue and Nobel 
1990). Concentrated in the center of the plant, the 
accumulated resources have maximum protection 
from fire, while the outer leaves, which are most 
susceptible to fire damage, can burn with little 
overall loss of stored carbohydrates. The timing of 
inflorescence emergence may also be important in 
reducing fire damage in reproductive agaves. Most 
natural lightning strikes that result in fires occur 
from May—August (Sellers and Hill 1974; Swetnam 
and Baisan 1996b). By this time, flower stalks are 
generally taller than the surrounding vegetation in 
grassland communities and above the fire zone 
(Slauson, personal observation). Spring fires or 
fires in denser oak woodland communities where 
the fire zone reaches into tree canopies could po- 
tentially damage emerging inflorescences more se- 
verely. 

Low fruit set is common in outcrossing, her- 
maphroditic plants such as agave (Sutherland and 
Delph 1984). Sutherland (1982) found that mean 
fruit set per inflorescence in paniculate agaves is 
consistently around 20% despite hand or open pol- 
lination treatments, suggesting that fruit set is pri- 
marily resource limited. Slauson (2000) studied 
fruit and seed set in three different open-pollinated 
populations of A. palmeri in Arizona, and also 
found fruit set to average around 20% while seed 
set varied from 26—33%. Fire did not appear to af- 
fect reproductive output in this study as no signif- 
icant differences in fruit and seed set were found 
between burned and unburned plants, and results 
(Fig. 3) were very similar to previous reports of 
unburned plants. The rosette shape and transloca- 
tion of resources in reproductive agaves appear to 
be important adaptations to fire that protect stored 
resources critical for reproduction. The significantly 
lower seed set at the CF site was observed in both 
burned and unburned plants, and suggests the cause 
is most likely a result of pollinator and/or resource 
availability at the site and not a result of burn treat- 
ment. 

Periodic fires may be important in promoting 
germination and establishment events of A. palmeri 
by reducing competition and opening up germina- 
tion sites, especially in dense grassland/woodland 
habitats. Germination of A. palmeri generally oc- 
curs in mid-late summer after monsoon rains have 
begun. During July 1999 (two months after the fire) 
approximately four inches of precipitation fell at 
the ME site, and during establishment of demog- 
raphy plots in burned areas in late July—early Au- 
gust, 16 seedlings were observed. These plants 
were assumed to be seedlings as they were <40 X 


10 MADRONO 


20 mm in height and diameter, and had no burn 
damage despite being located near burned plants. 
A cursory search for seedlings was conducted in 
several unburned areas adjacent to the burned site, 
but no seedlings were found. 


CONCLUSIONS 


Prescribed natural and human-ignited fires are 
important ecosystem management tools used to al- 
ter community composition and forage conditions, 
decrease fuel accumulation, and reduce the poten- 
tial of catastrophic, stand-replacing wildfires. Al- 
though historic changes in the southwestern U.S. 
landscape have been significant due to fire exclu- 
sion, the return of fire in the sites studied did not 
appreciably impact nectar and pollen production, 
fruit and seed set, or initial mortality of A. palmeri. 
Standing crop results indicated that at least through 
mid-August in the sites examined, food resources 
of lesser long-nosed bats were not limited as a re- 
sult of fire. Habitat preferences and several mor- 
phological and physiological adaptations appear to 
adequately protect the majority of stored resources 
of A. palmeri during fire events, although fuel loads 
within individual microhabitats of agaves can result 
in variable fire damage to rosettes. Further study is 
needed to more clearly understand how fire affects 
population dynamics and mortality of A. palmeri 
beyond the initial post-fire period. 


ACKNOWLEDGEMENTS 


The author thanks J. McAuliffe, P. Scott, C. Edminster, 
and two anonymous reviewers for helpful guidance and 
comments on the manuscript; J. Borgmeyer, M. Johnson, 
K. Mueller, K. Rice, S. Ahearn, D. Hansen, N. Grant, S. 
Garrison, T. Omar, K. Smith, and volunteers and staff of 
the Desert Botanical Garden for field and laboratory as- 
sistance; P. Sundt for assistance with plant species data, 
R. King, Rocky Mountain Research Station, for statistical 
advice and consultation; B. McDonald of the Sycamore 
Ranch and Ben Brown and the Animas Foundation of the 
Gray Ranch for logistical and study site location assis- 
tance; and Ginny and Dave Dalton for assistance in bat 
identification and observations. This research was sup- 
ported in part by funds provided by the Rocky Mountain 
Research Station, U.S. Department of Agriculture, Forest 
Service, and the Desert Botanical Garden. 


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AND B. S. ROTH. 1981. Sexual reproduction in aga- 
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McPHERSON, G. R. 1995. The role of fire in the desert 
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MADRONO, Vol. 49, No. 1, pp. 12—15, 2002 


ADULT SEX RATIO OF ARCEUTHOBIUM GILLIT (VISCACEAE) 


ROBERT L. MATHIASEN 
School of Forestry, Northern Arizona University, Flagstaff, AZ 86011 


CAROLYN M. DAUGHERTY 
Geography and Public Planning, Northern Arizona University, 
Flagstaff, AZ 86011 


ABSTRACT 


The adult sex ratio of Arceuthobium gillii was determined from five populations in southern Arizona 
and one population in Chihuahua, Mexico. A total of 6154 mistletoe plants were sexed on a total of 38 
host trees; 3096 of these were male plants (50.3%) and 3058 were female (49.7%). A chi-square analysis 
indicated that these numbers were not significantly different from an essentially 1:1 sex ratio. Furthermore, 
although there was variation between trees (more male or female plants) the sex ratio on individual trees, 


and for each population, was also essentially 1:1. 


Key Words: Chihuahua pine dwarf mistletoe, Chihuahua pine, sex ratio 


Dwarf mistletoes (Arceuthobium spp., Visca- 
ceae) are dioecious, parasitic flowering plants that 
commonly occur on members of the Pinaceae in 
western North America. In general, the sex ratio 
(female plants : male plants) of dwarf mistletoes has 
been reported to be 1:1 (Hawksworth and Wiens 
1996; Mathiasen et al. 1998). However, other in- 
vestigators have reported female-biased sex ratios 
for several dwarf mistletoes: Arceuthobium ameri- 
canum Engelm. in Canada (Muir 1966), A. globos- 
um Hawksw. & Wiens subsp. globosum and A. 
strictum Hawksw. & Wiens in Durango, Mexico 
(Hawksworth and Wiens 1996), A. pusillum Peck 
in Minnestoa (Baker et al. 1981), and most notably, 
A. tsugense (Rosendahl) G. N. Jones subsp. tsu- 
gense and subsp. mertensianae Hawksw. & Nick- 
rent from the Pacific Northwest and Alaska (Wiens 
et al. 1996). The majority of these reports of fe- 
male-biased sex ratios for dwarf mistletoes were 
based on small sample sizes (approximately 50— 
500 total plants). However, the latter study used 
over 3000 plants from 16 populations distributed 
from southern Oregon to southeast Alaska and re- 
ported a significant female-biased sex ratio for 10 
of the populations and for the pooled data from all 
16 areas (59% females: 41% males). 

Because of the female-biased sex ratio reported 
for Arceuthobium tsugense (Wiens et al. 1996), 
Hawksworth and Wiens (1996) recommended that 
the sex ratios of other dwarf mistletoes be exam- 
ined. Therefore, this study was initiated to provide 
additional information on the adult sex ratios of 
dwarf mistletoes. Arceuthobium gillii Hawksw. & 
Wiens was chosen for this study because of the 
strong sexual dimorphism exhibited by this species 
(Hawksworth and Wiens 1996, see page 147). This 
strong sexual dimorphism allowed male and female 
plants to be easily distinguished. Other dwarf mis- 
tletoes also exhibit extreme sexual dimorphism and 


a study of this characteristic would be a valuable 
contribution to the systematics of the genus (Hawk- 
sworth and Wiens 1996, Mathiasen et al. 1998). 


MATERIALS AND METHODS 


The adult sex ratio for Arceuthobium gillii was 
determined for five populations in southern Arizona 
and for one population in Chihuahua, Mexico (Ta- 
ble 1, Fig. 1). At each of the locations sampled, at 
least four, severely infected Chihuahua pines (Pinus 
chihuahana Englem.) were selected, their diameters 
at breast height (1.3 m above the ground) measured 
to the nearest 0.5 cm, and a dwarf mistletoe rating 
(DMR) assigned to each tree using the 6-class sys- 
tem (Hawksworth 1977). For reasons of safety and 
efficiency sampling was restricted to trees <30 cm 
in diameter at breast height and a DMR >4. 

Selected trees were flagged and after diameters 
and dwarf mistletoe ratings were recorded, each 
tree was cut as close to the ground as possible. 
After each tree was cut, its total height to the near- 
est 0.1 m, was measured. Live branches that broke 
from the tree when it was cut were examined for 
mistletoe plants first. Then each live branch still 
attached to the tree, starting at the bottom of the 
crown, was removed and examined for mistletoe 
plants. The sex of each observed plant on a branch 
was recorded and the branch discarded well away 
from the tree so the branch would not be re-sam- 
pled. This process was repeated until all the live 
branches on each tree had been examined. Only 
plants that could be accurately sexed were tallied. 
One person examined branches for mistletoe plants 
and one person recorded data. This was to insure 
that the person examining mistletoe plants did not 
know how many males or females had been tallied 
as sampling proceeded. The same person recorded 
data until a tree had been completed. Data were not 
summarized until all the selected trees had been 


2002] MATHIASEN AND DAUGHERTY: DWARF MISTLETOW SEX RATIO 13 


TABLE 1. POPULATION LOCATIONS, NUMBER OF TREES SAMPLED, MEAN DIAMETERS, MEAN DWARF MISTLETOE RATINGS, 
MEAN TREE HEIGHTS, NUMBER OF PLANTS SAMPLED, AND ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM 
GILLI. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). DBH = diameter breast height; 
DMR = dwarf mistletoe rating (Hawksworth 1977). 


Mean Mean 
Trees DBH Mean height Plants Percent 

Population location sampled (cm) DMR (m) sampled female P 
Bear Canyon 

Santa Catalina 

Mountains, AZ 7 16.0 Deo} 8.5 1094 47.9 0.164 
Gardner Canyon 

Santa Rita 

Mountains, AZ 4 15.0 5.8) 7.0 Syl 50.5 0.825 
Carr Canyon 

Huachuca 

Mountains, AZ 9 16.0 5.6 8.1 1348 50.6 0.663 
Pinery Canyon 

Chiricahua 

Mountains, AZ 4 18.0 5.6 8.3 1936 50.7 0.525 
Upper Cave Creek 

Chiricahua 

Mountains, AZ 7 20.0 5.6 10.6 568 49.3 OW3m 
Chihuahua, Mexico 

(Sierra Madre 

Occidental) 7 16.0 Sis) Woe 697 48.2 0.344 
Total/mean 38 17.0 5.4 8.4 6154 49.7 0.702 

120° ey 110° 1053 


JOD 


Arizona 
35° 


New Mexico 


30° 


ie TOE 105° 


Fic. 1. Approximate locations of study sites. 1—Bear Canyon, Santa Catalina Mountains, Arizona; 2—Gardner Can- 
yon, Santa Rita Mountains, Arizona; 3—-Carr Canyon, Huachuca Mountains, Arizona; 4—Pinery Canyon, Chiricahua 
Mountains, Arizona; 5—Upper Cave Creek, Chiricahua Mountains, Arizona; and 6—Sierra Madre Occidental, Chi- 
huahua, Mexico. 


14 MADRONO [Vol. 49 


TABLE 2. ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM GILLII FOR EACH TREE SAMPLED IN CARR CANYON, 
HuAcHUCA MOUNTAINS, ARIZONA. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). DBH 
= diameter breast height; DMR = dwarf mistletoe rating (Hawksworth 1977). 


dire DBH Height Plants Percent 

number (cm) DMR (m) sampled female IP 

1 16.0 5 8.1 93 54.8 0.351 
yy 10.5 6 5.6 153 53.6 0.374 
3 19.5 6 10.3 196 49.0 0.775 
4 16.0 5 7.8 76 44.7 0.359 
5) 18.0 5 8.7 141 S32 0.449 
6 14.0 6 Wood) 83 48.2 0.742 
7 20.5 6 8.6 268 49.3 0.807 
8 16.5 6 8.1 164 47.6 L532 
9 16.0 5 8.0 174 54.0 0.289 
Total/mean 16.0 5.6 8.1 1348 50.6 0.663 


sampled. After summarizing the total number of 
plants sexed for the selected trees, additional trees 
were sampled to bring the total number of plants 
sexed to a minimum of 500, if necessary. 

Data were collected in May of 1998—2000 and 
in June 2001. This was either during anthesis of 
male plants or shortly after it (June). A chi-square 
analysis was used to determine if the ratio of male 
to female mistletoe plants exhibited a sex bias. We 
used a P value of =0.05 to determine the existence 
of statistically significant differences (Zar 1999). 


RESULTS 


The general location of study sites, the number 
of trees sampled, tree mean diameters, mean dwarf 
mistletoe ratings, mean heights, the number of mis- 
tletoe plants sexed, and the percentage of female 
plants for each population sampled are presented in 
Table 1. We sampled a total of 6154 mistletoe 
plants that could be accurately sexed on 38 trees. 
Of the plants we sexed, 3096 (50.3%) were males 
and 3058 (49.7%) were females. The difference in 
the number of male and female plants was not sig- 
nificantly different from the number expected for a 
1:1 sex ratio (P = 0.628); therefore, the adult sex 
ratio of Arceuthobium gillii on the 38 trees was 
essentially 1:1 (Table 1). 

There was a large amount of variation in the sex 
ratio for A. gillii between individual trees at each 
of the study sites. An example of this variation is 
illustrated by the trees sampled in the Huachuca 


(Table 2) and Santa Rita Mountains, Arizona (Table 
3). Some trees had more female plants than males 
and vice versa, but no trees exhibited significantly 
different sex ratios from a 1:1 ratio. In addition, 
when the sex ratio was determined using all of the 
trees in each population, no significant differences 
from a 1:1 sex ratio were detected for any of the 
populations (Tables 1-3). 


DISCUSSION 


Although several investigators have reported sig- 
nificant female-biased adult sex ratios for several 
dwarf mistletoes, we found that the adult sex ratio 
for Arceuthobium gillii is essentially 1:1. These re- 
sults were expected because many other dwarf mis- 
tletoes also have 1:1 adult sex ratios (Hawksworth 
and Wiens 1996; Mathiasen et al. 1998). Because 
many of the reports of female-biased adult sex ra- 
tios for dwarf mistletoes are based on relatively 
small sample sizes (<500 plants), they may not rep- 
resent an accurate estimate of the sex ratios for 
these mistletoes (Mathiasen and Shaw 1998; 
Daugherty and Mathiasen 1999). 

Our results demonstrate the variation in sex ratio 
that can occur among individual trees. This tree-to- 
tree variation has been demonstrated in other stud- 
ies of mistletoe sex ratio (Nixon and Todzia 1985; 
Mathiasen and Shaw 1998; Daugherty and Mathi- 
asen 1999). Because of this tree-to-tree variation, a 
large sample of mistletoe plants should be sampled 
for dioecious mistletoe sex ratio studies and data 


TABLE 3. ADULT SEX RATIOS (PERCENT FEMALE) FOR ARCEUTHOBIUM GILLII FOR EACH TREE SAMPLED IN GARDNER CAN- 
YON, SANTA RITA MOUNTAINS, ARIZONA. No sex ratios exhibited a significant sex bias. Chi-square statistics (P = 0.05). 
DBH = diameter breast height; DMR = dwarf mistletoe rating (Hawksworth 1977). 


Tree DBH Height Plants Percent 

number (cm) DMR (m) sampled female iE 

1 14.5 5 al ae) 50.5 0.925 
2 10.0 6 Se 123 54.5 0.321 
3 20.0 5 V8 206 49.0 0.781 
4 16.0 5 7.8 69 49.3 0.904 
Total/mean 15.0 53) 7.0 Sill 50.5 0.825 


2002] MATHIASEN AND DAUGHERTY 
should be analyzed using the results for the entire 
population and not on an individual tree basis (Ma- 
thiasen and Shaw 1998). We sampled over 6000 
adult plants from several separate populations of A. 
gillii and found that this species exhibits a 1:1 adult 
sex ratio in each population sampled and when the 
data were pooled for all populations. Therefore, we 
contend that A. gillii will exhibit a consistent 1:1 
adult sex ratio whenever large numbers of plants 
(>500) are sexed for populations of this dwarf mis- 
tletoe. 

We plan to determine the adult sex ratios for oth- 
er dwarf mistletoes in the Southwest and for those 
already reported to have female-biased sex ratios 
based on small samples, such as Arceuthobium 
americanun, A. globosum, and A. strictum (Hawk- 
sworth and Wiens 1996). However, we now hy- 
pothesize that when a large number of dwarf mis- 
tletoe plants are sampled for each of these dwarf 
mistletoes, the overall adult sex ratio will be essen- 
tially 1:1 as it was for the populations of A. gillii 
we sampled in this study. 


ACKNOWLEDGMENTS 


We would like to extend our appreciation to Del Wiens 
for suggesting this study be conducted. The field assis- 
tance of Dave Russell in the Chiricahua Mountains is sin- 
cerely appreciated also. 


LITERATURE CITED 


BAKER, FE A., D. W. FRENCH, AND G. W. HUDLER. 1981. 
Development of Arceuthobium pusillum on black 
spruce. Forest Science 27:203-—205. 


MADRONO, Vol. 49, No. 1, p. 15, 2002 


: DWARF MISTLETOW SEX RATIO 15 


DAUGHERTY, C. M. AND R. L. MATHIASEN. 1999. Adult sex 
ratio of Phoradendron juniperinum in ten severely 
infected Juniperus monosperma in northern Arizona. 
Madrono 46:169—176. 

HAWKSWORTH, E G. 1977. The 6-class dwarf mistletoe rat- 
ing system. USDA Forest Service Research Note 
RM-48. 

AND D. WIENS. 1996. Dwarf mistletoes: biology, 
pathology, and systematics. USDA Forest Service 
Agric. Handb. 709. 

MATHIASEN, R. L., C. G. PARKS, B. W. GEILS, AND J. S. 
BEATTY. 1998. Notes on the distribution, host range, 
plant size, phenology, and sex ratio of two rare dwarf 
mistletoes from Central America: Arceuthobium 
hawksworthii and A. hondurense. Phytologia 84:154— 
164. 

AND D. C. SHAw. 1998. Adult sex ratio of Arceu- 
thobium tsugense in six severely infected Tsuga het- 
erophylla. Madrono 45:210—214. 

Murr, J. A. 1968. Biology of dwarf mistletoe (Arceuthob- 
ium americanum) in Alberta. Internal Report A15, 
Canada Department of Fisheries and Forestry, pp. 1— 
29, Calgary, Alberta. 

NIXoNn, K. C. AND C. A. TobziA. 1985. Within-population, 
within-host species, and within-host tree sex ratios in 
mistletoe (Phoradendron tomentosum) in central Tex- 
as. American Midland Naturalist 114:304—310. 

WIENS, D., D. L. NICKRENT, C. G. SHAW, E G. HAwK- 
SWORTH, P. E. HENNON, AND E. J. KING. 1996. Embry- 
onic and host-associated skewed adult sex ratios in 
dwarf mistletoe. Heredity 77:55—63. 

ZAR, J. H. 1999. Biostatistical analysis. Prentice Hall, Up- 
per Saddle River, NJ. 


ANNOUNCEMENT 


First ANNOUNCEMENT AND CALL FOR SESSIONS 


FOURTH INTERNATIONAL SYMPOSIUM ON GRASS 
SYSTEMATICS AND EVOLUTION AND 


THIRD INTERNATIONAL CONFERENCE ON THE 
COMPARATIVE BIOLOGY OF THE MONOCOTYLEDONS 


The Third International Conference on the Com- 
parative Biology of the Monocotyledons and Fourth 
International Symposium on Grass Systematics and 
Evolution will be hosted by Rancho Santa Ana Bo- 
tanic Garden (Claremont, CA, USA) on 30 March— 
5 April 2003. Topics will include morphology, 
anatomy, development, reproductive biology, mo- 
lecular biology, cytology, genomics, genetics, bio- 
chemistry, paleobotany, phylogenetics, classifica- 
tion, biogeography, ecology, and data integration. 
Sessions will be devoted to particular groups within 


monocots such as grasses and orchids. Monocots 
III will provide a rare opportunity for researchers 
in diverse fields to interact, share ideas, and form 
collaborations. We invite proposals from those who 
wish to organize sessions. A call for contributed 
papers and posters will follow. The conference pro- 
ceedings will be published. Springtime marks the 
flowering peak of the diverse California flora, and 
field trips are planned. Visit www.monocots3.org 
for conference details; or write Monocots III, Ran- 
cho Santa Ana Botanic Garden, 1500 North College 
Avenue, Claremont, CA 91711-3157 USA; E-mail 
info@monocots3.org; fax 1.909.626.7670; tele- 
phone 1.909.625.8767 ext. 333. Co-sponsors in- 
clude the American Society of Plant Taxonomists, 
Botanical Society of America, and the International 
Association for Plant Taxonomy. 


MaproNno, Vol. 49, No. 1, pp. 16-19, 2002 


ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE (POLYGONACEAB), 
A NEW VARIETY FROM THE SOUTHERN SIERRA NEVADA, 
CALIFORNIA 


DANA A. York! 
Death Valley National Park, PO. Box 579, Death Valley, CA 92328 


ABSTRACT 


Eriogonum ovalifolium var. monarchense is a new variety discovered on a limestone formation in the 
southern Sierra Nevada. It is only known from one population in the Kings River drainage basin. It is 
morphologically similar to E. ovalifolium var. vineum; an endangered species found on limestone outcrops 
in the San Bernardino Mountains. The habit of the two varieties is different primarily in the angle that 


the flowering stems arise from the base. 


Key words: Eriogonum ovalifolium var. monarchense, Kings River, Sierra Nevada, Monarch buckwheat, 


limestone 


In 1995, botanical explorations in the Kings Riv- 
er canyon of the southern Sierra Nevada yielded 
three previously unknown vascular plant taxa from 
the limestone (marble) outcrops around Boyden 
Cavern, near Kings Canyon National Park. Heter- 
otheca monarchensis Semple, Shevock, & York 
and Gilia yorkii Shevock & A. G. Day were de- 
scribed within a few years of their discovery. A 
new variety of buckwheat from the Boyden Cavern 
limestone required years of research and follow-up 
collecting to verify its taxonomic status. The first 
collections of the three new taxa were made on the 
same day (31 July 1995) in Monarch Wilderness 
(Sierra and Sequoia National Forests). 


Eriogonum ovalifolium Nutt. var. monarchense 
D. A. York, var. nov. (Fig. 1)—Type: USA, CA, 
Fresno Co, 86 km E of Fresno, Sierra National 
Forest, Monarch Wilderness, 2.4 km NW of Boy- 
den Cave on N side of the Kings River canyon, 
36°50'08"N, 118°49'19"W (NAD 83), 1815 m, 31 
July 1995, York 111 & Shevock (holotype CAS; 
isotypes JEPS, NY). Paratype: USA, CA, Fresno 
Co, 86 km E of Fresno, Sierra National Forest, 
Monarch Wilderness, 2.4 km NW of Boyden 
Cave on N side of Kings River canyon, 
36°50'08"N, 118°49'19"W (NAD 83), 1815 m, 19 
July 1996, York 1250 (RSA). 


Eriogonum ovalifolium var. vineum accedentes 
sed caules floriferentes decumbenti ad ascendenti 
sunt. 

Pulvinate perennials forming mats up to 30 cm 
across (Fig. 1A); leaves basal, petiolate, tomentose, 
5—22 mm long, the margins flat to slightly crisped, 
petioles 2-10 mm long, blades elliptic to orbicular, 
3-12 mm long, 3-12 mm wide; flowering stems 
scapose, l|—many per matted clump, decumbent to 
ascending, 2—6(9) cm long, tomentose to floccose; 


'R-mail: Dana_York @nps.gov 


inflorescences capitate, the head 1.5—4 cm across; 
bracts scale-like, 3, 1-5 mm long; involucres clus- 
tered 4—6 per head, sessile, tomentose, turbinate, 
5—8 mm long, with 5 rounded or acute teeth up to 
2 mm long; flowers white to cream with green (ag- 
ing red) midribs, 4-6 mm long, glabrous, the peri- 
anth lobes dimorphic, the perianth lobes of the out- 
er whorl mostly twice the width of the inner whorl 
(Fig. 1B); stamens mostly exserted, 1-3 mm long, 
the anthers 0.4—0.6 mm long; achenes brown, 2-3 
mm long, glabrous. 


DISTRIBUTION, HABITAT, AND PHENOLOGY 


Eriogonum ovalifolium var. monarchense (Mon- 
arch buckwheat) is a rare neoendemic found in 
eastern Fresno County, in the southern Sierra Ne- 
vada. The only known population (type locality) 
grows on the north side of a limestone formation 
in the Kings River canyon above 1800 m (5900 
feet), in the vicinity of Boyden Cavern. The plants 
are located in Monarch Wilderness, just below the 
Monarch Divide, in the Sierra National Forest. 
Monarch Divide rises over 880 m above the canyon 
floor and is the boundary between the middle and 
south forks of the Kings River and the Sierra and 
Sequoia National Forests. The divide is rugged and 
varied with a mix of metamorphic and igneous 
rocks. The limestone component is typified by steep 
slopes and sheer cliffs. 

Eriogonum ovalifolium var. monarchense forms 
dense mats on ledges and crevices in sandy soils 
developed from decomposed limestone. The popu- 
lation consists of approximately 30 plants scattered 
over a few thousand square meters. Flowers are 
present from June to August. Associates include 
Argyrochosma jonesii (Maxon) Windham, Bromus 
madritensis L. ssp. rubens (L.) Husn., Cercocarpus 
intricatus Wats., Erigeron aequifolius Hall, Erysi- 
mum capitatum (Dougl.) Greene ssp. capitatum, 
Garrya flavescens Wats., Gilia yorkii, Heuchera ru- 
bescens Torr. var. rydbergiana Rosend., Butt. & 


2002] 


= 


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SS 


ae 


6 
D> 


Pk 
ae te 
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4.0. ede 
sexe 


Fic. 1. 


ex 2 
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YORK: ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE 1) 


1 cm 


Habit of Eriogonum ovalifolium Nutt. var. monarchense D. A. York, and a detailed illustration of a flower 


from holotype collection and photographs. A. Mature plant in flower. B. Flower with detail of perianth lobes in an 


erect position. Drawings by Laura Cunningham. 


Lak., Pinus monophylla Torr. & Frém., Selaginella 
asprella Maxon, Streptanthus fenestratus (Greene) 
J.T. Howell, and Yucca whipplei Torr. 


RELATIONSHIPS 


The new taxon differs in several respects from 
the other California varieties of Eriogonum ovali- 
folium (Table 1). Eriogonum ovalifolium var. mon- 
archense is morphologically similar to E. ovalifol- 
tum Nutt. var. vineum (Small) Jepson; an endan- 
gered species known only from limestone outcrops 
in the San Bernardino Mountains. Eriogonum oval- 
iYfolium var. vineum has flowering stems that are 
generally erect in contrast to the decumbent to as- 
cending flowering stems of E. ovalifolium var. mon- 
archense. These two varieties occur in disjunct 
mountain ranges and are approximately 325 km 
apart from each other. 

Eriogonum ovalifolium Nutt. var. purpureum (A. 
Nelson) Durand is a variety that occurs in the east- 


ern Sierra Nevada and other mountain ranges in 
western North America on various substrates (Re- 
veal 1989). It differs from E. ovalifolium var. mon- 
archense by having longer leaf blades and shorter 
involucres. 

Eriogonum ovalifolium Nutt. var. nivale (Canby) 
M.E. Jones is geographically close to, but not sym- 
patric with, E. ovalifolium var. monarchense. In 
California, it occurs mostly on granites in subalpine 
and alpine habitats throughout the Sierra Nevada, 
Cascades, and White Mountains (Hickman 1993). 
It differs from E. ovalifolium var. monarchense by 
having generally smaller leaves and shorter flow- 
ering stems and perianth lobes. 


KEY TO THE CALIFORNIA VARIETIES OF 
ERIOGONUM OVALIFOLIUM 


las-Blowers yellow. sm << 24a. . 2.2 var. ovalifolium 
1b. Flowers white, cream, red, or purplish 


2a. Flowers 2—3 mm long var. nivale 


18 


COMPARISON OF ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE WITH THE OTHER E. OVALIFOLIUM VARIETIES KNOWN FROM CALIFORNIA. 


TABLE 1. 


E. o. vineum 


E. o. purpureum 


E. o. ovalifolium 


E. o. nivale 


E. o. eximium 


E. o. monarchense 


Characters 


erect erect erect erect 


erect 


decumbent to ascend- 


Flowering stems 


ing 
2-5 cm long 
unmargined 


3-6 cm long 
unmargined 
orbicular 


5—20 cm long 
+ unmargined 


obovate 


4—20 cm long 
+ unmargined 


obovate 


0.3—5 cm long 


5—10 cm long 


+ unmargined 


orbicular 


brown-margined 


Leaf blades 


elliptic to spatulate 
0.5—2 cm long 


4—6.5 mm 


elliptic to orbicular 
0.3—1.2 cm long 


5-8 mm 


0.6—1.2 cm long 


5—7 mm 


0.5—2 cm long 


4—6.5 mm 


1-6 cm long 
4—6.5 mm 
yellow 


0.2—0.8 cm long 


2—4 mm 


Involucre lengths 


Flowers 


white to cream 


purple or white to 


white to cream white to cream 


white to cream 


cream 
4—6 mm long 


3-5 mm long 


rocky 


2-3 mm long 4—6 mm long 


4—6 mm long 


sandy 
granites 


4—6 mm long 


rocky 


sandy or gravelly 
various substrates 


W. North America 


sandy or gravelly 
various substrates 


W. North America 


sandy or gravelly 


granites 


Habitat 


limestone 


limestone 


San Bernardino Mtns. 


W. North America 


N. Sierra Nevada, 


S. Sierra Nevada, 


Range 


Nevada 
1800—3400 m elev. 


California 
1800 m elev. 


MADRONO 


1200—2800 m elev. 1500—2100 m elev. 


1200—2900 m elev. 


1500—4100 m elev. 


[Vol. 49 


2b. Flowers >3 mm long 
3a. Leaves obviously brown-margined 
Ce nr Cae. Meet, AS a Sic var. eximium 
3b. Leaves not distinctly brown-margined 
4a. Flowering stems 4—20 mm long; in- 
volucres 4—6.5 mm long; various 
SUDSERateSs eee es ee var. purpureum 
4b. Flowering stems 2—6 (9) mm long; 
involucres 5—8 mm long; carbonate 
substrates 
5a. Flowering stems decumbent to 
ascending; southern Sierra Ne- 
vada Range var. monarchense 
5b. Flowering stems mostly erect; 
San Bernardino Mountains .. . 
A AE rte S25 5 var. vineum 


DISCUSSION 


Eriogonum ovalifolium var. monarchense is al- 
lopatric from the other varieties of E. ovalifolium. 
The calcareous habitat and associated species 
where Eriogonum ovalifolium var. monarchense is 
found are more typical of pinyon pine communities 
in the eastern Sierra Nevada and the desert ranges 
of California and Nevada. It is possible that Erio- 
gonum ovalifolium var. monarchense is allied with 
the varieties from the desert and southern Califor- 
nia mountains and not with the Sierran Eriogonum 
ovalifolium var. nivale. There are four other calci- 
cole vascular plants endemic to the King River ba- 
sin (York 1999). These include Eriogonum nudum 
Benth. var. regirivum Reveal & J. Stebbins, Gilia 
yorkil, Heterotheca monarchensis, and Streptanthus 
fenestratus. Heterotheca monarchensis (occurs on 
south-facing slopes with a population very near Er- 
togonum ovalifolium var. monarchense) and Gilia 
yorkii are allied with their respective desert con- 
geners (York 1999). They evolved from desert taxa 
that spread into the California Floristic “Province 
during Xerothermic periods of the Quaternary 
(York 1999). Other evidence of this desert link in- 
clude taxa found on the limestone outcrops around 
Boyden Cavern, such as Achnatherum hymenoides 
(Roem. & Schultes) Barkworth, Argyrochosma jo- 
nesii, Melica frutescens Scribn., and Petrophyton 
caespitosum (Nutt.) Rydb., that are mostly rare in 
the Sierra Nevada and common in the desert ranges 
(York 1999). 


RARITY 


Eriogonum ovalifolium var. monarchense, pre- 
viously unknown and uncollected, is a rare taxon 
due to its lithophytic nature on limestone, a rel- 
atively uncommon substrate in the southern Si- 
erra Nevada. Because of the remoteness and rug- 
ged physiography of the Kings River canyon 
limestones, it is unlikely that Eriogonum ovali- 
folium var. monarchense occurs outside the river 
basin. It is a rare and localized endemic worthy 
of conservation efforts. Anthropogenic impacts 
are not likely because the population is remote 
and within a designated wilderness. If the type 


2002] 


locality is indeed the only population, then this 
taxon is vulnerable to extinction due to stochastic 
events or genetic drift. 


ACKNOWLEDGMENTS 


I thank Laura Cunningham for her carefully detailed 
work on the illustrations, and Johanna Arnegger (my Aus- 
trian friend) for correcting the Latin diagnosis. I also thank 
Jim Shevock for following me through poison oak, dense 
brush, loose rock, and up steep slopes during an uncom- 
fortably hot day to assist me in collecting the type mate- 


YORK: ERIOGONUM OVALIFOLIUM VAR. MONARCHENSE 19 


rial. James Reveal and an anonymous reviewer provided 
useful comments on my drafts for which I am grateful. 


LITERATURE CITED 


HICKMAN, J. C. 1993. Polygonaceae in J. Hickman (ed.), 
The Jepson manual: Higher plants of California. Uni- 
versity of California Press, Berkeley. 

REVEAL, J. L. 1989. The eriogonoid flora of California 
(Polygonaceae: Eriogonoideae). Phytologia 66:295— 
414. 

York, D. A. 1999. A phytogeographic analysis of the 
Kings River Basin, California. M.S. thesis. California 
State University, Fresno, CA. 


MADRONO, Vol. 49, No. 1, pp. 20—21, 2002 


HESPEROYUCCA WHIPPLEI AND YUCCA WHIPPLEI (AGAVACEAE) 


JEFFREY A. GREENHOUSE 
Jepson Herbarium, 1001 Valley Life Sciences Building #2465, 
University of California, Berkeley, CA 94720-2465 


JOHN L. STROTHER 
University Herbarium, 1001 Valley Life Sciences Building #2465, 
University of California, Berkeley, CA 94720-2465 


ABSTRACT 


The name Yucca whipplei dates from 1861, not 1859. The names Hesperoyucca, Hesperoyucca whip- 
plei, and Hesperoyucca whipplei var. graminifolia date from 1893. 


Chaparral yucca has been widely known for 140 
years or so as Yucca whipplei. The plants have long 
been recognized as markedly unlike other yuccas 
(e.g., Engelmann 1871). Baker (1892) suggested 
the plants are generically distinct from other yuc- 
cas. But, chaparral yucca has continued to reside 
within Yucca in most taxonomic and floristic treat- 
ments. Primarily on molecular similarities and dif- 
ferences, Bogler and Simpson (1995), Clary and 
Simpson (1995), and others cited by them, have 
revived the suggestion that chaparral yucca should 
be placed in a separate genus, Hesperoyucca. Here, 
we review the histories of the botanical names. 

When the name Yucca whipplei was first used in 
print, Torrey (1859, p. 222) described (or at least 
diagnosed) the taxon and wrote, “If it prove to be 
a distinct species it may be called Y. Whipplei.”’ 
We consider that use of Yucca whipplei by Torrey 
to have been as a provisional name and, therefore, 
we believe that the name was not validly published 
in 1859. The next use of Yucca whipplei that we 
have found was also by Torrey (1861). Conditions 
for valid publication of Yucca whipplei were met 
by Torrey in 1861. We believe the correct citation 
to be: 


Yucca whipplei Torrey in J. C. Ives, Rep. Colorado 
R. 4(Botany):29. 1861. 


Plants of Yucca whipplei have also been called 
Hesperoyucca whipplei. There is no entry for Hes- 
peroyucca in the published version of ING (Farr et 
al. 1979). In June 2001, requests to the website for 
Index Nominum Genericorum (currently, http:// 
rathbun.si.edu/botany/ing/) for Hesperoyucca elic- 
ited the response, ““No records found with: Search 
for Name “‘Hesperoyucca.’ ”’ 

The earliest use of Hesperoyucca of which we 
are aware was by Engelmann (1871), who parti- 
tioned Yucca into: “§ EU-YUCCA,”’ with three 
subordinate groups (Sarcocarpa, Clistocarpa, and 
Chaenocarpa) with one species each, and *Ԥ HES- 
PERO-YUCCA,” which included Yucca whipplei 


as the only species. In 1873, Engelmann provided 
a summary “SYSTEMATIC ARRANGEMENT” 
for Yucca in which he named: I. Euyucca (which 
included: A. Sarcoyucca, B. Clistoyucca, and C. 
Chaenoyucca) and II. Hesperoyucca. In both clas- 
sifications, the rankless Hesperoyucca included 
only Yucca whipplei. 

In a third paper (“‘corrections and additions” to 
the 1873 paper), Engelmann (1875) provided yet 
another “‘synopsis”’ for Yucca and that time parti- 
tioned Yucca into four codrdinate elements named 
Sarcoyucca, Clistoyucca, Chaenoyucca, and Hes- 
peroyucca. Once again, Engelmann did not indicate 
rank for his subdivisions of Yucca. 

Baker (1876), in notes on Yucca whipplei, wrote, 
**... Dr. Engelmann, who has considered it [i.e., 
Yucca whipplei|] as the type of a new subgenus, 
which, alluding to its western locality, he has called 
Hesperoyucca. At present this subgenus is only 
known to contain a single species.’’ We believe that 
Baker’s remarks were sufficient to establish sub- 
generic rank and that the correct citation is: 


Yucca L. subg. Hesperoyucca (Engelmann) Baker, 
Gard. Chron. n.s. 6:196. 1876. Based on Yucca 
[unranked] Hesperoyucca Engelmann in S. 
Watson et al., Botany (Fortieth Parallel), 497. 
1871, as ““§ 2. HESPERO-YUCCA.” 


In 1892, Baker wrote of Y. whipplei, “‘I now 


think this had better be kept as a genus distinct 


from Yucca, under Engelmann’s name Hesperoyuc- 
ca.”’ Baker, nevertheless, treated whipplei as be- 
longing to Yucca in 1892. We suggest that Baker’s 
statement in 1892 may have been a “‘note-added- 
in-proof”’ and that Baker’s statement was not suf- 
ficient to establish generic rank for Hesperoyucca. 

Trelease (1893) accepted Baker’s suggestion of 
generic rank for Hesperoyucca, made a specific 
combination in Hesperoyucca, and coined a varietal 
nomen novum in Hesperoyucca. We believe correct 
citations for the names are: 


2002] 


Hesperoyucca (Engelmann) Trelease, Rep. (An- 
nual) Missouri Bot. Gard. 4:208. 1893. Based 
on: Yucca [unranked] Hesperoyucca Engel- 
mann in S. Watson et al., Botany (Fortieth Par- 
allel) 497. 1871. —Type: Hesperoyucca whip- 
plei (Torrey) Trelease = Yucca whipplei Tor- 
rey. 

Hesperoyucca whipplei (Torrey) Trelease, Rep. 
(Annual) Missouri Bot. Gard. 4:208. 1893, as 
‘Y. Whipplei’ under ‘HESPEROYUCCA’. 
Given the use of Hesperoyucca whipplei else- 
where in the article by Trelease (figure cap- 
tions, in a name at varietal rank), we consider 
the ““Y”’ at the comb. nov. to be a typographic 
error and the attribution of the combination to 
Baker to have been superfluous. Basionym: 
Yucca whipplei Torrey. 


Hesperoyucca whipplei (Torrey) Trelease var. gra- 
minifolia Trelease, Rep. (Annual) Missouri 
Bot. Gard. 4:215. 1893. We maintain that Tre- 
lease coined a nom. nov. based on: Yucca gra- 
minifolia Wood, 1868 [Proc. Acad. Nat. Sci. 
Philadelphia 20:167], non Zucc., 1837. 


After submitting this paper to Madrono for pub- 
lication (June 2001), drafts of the manuscript (or 
relevant excerpts) were sent to colleagues with es- 
pecial interest in botanical nomenclature. A flurry 
of e-mails ensued. Some nomenclaturists held that 
Baker validated Hesperoyucca at generic rank in 
1892. Others opined that Baker did not validly pub- 
lish Hesperoyucca at generic rank and that Hespe- 
royucca was validly published at generic rank by 
Trelease in 1893. To us, Baker explicitly treated 
Hesperoyucca at subgeneric rank in 1876 and did 
not explicitly treat Hesperoyucca at generic rank in 
1892. 

As a result of having the matter of valid publi- 
cation of Hesperoyucca at generic rank bandied 
about, the website for ING posted this entry in No- 
vember 2001: 


Hesperoyucca (G. Engelmann) J. G. Baker, Bull. 
Misc. Inform. Kew 1892: 8. Jan 1892. 

T.: H. whipplei (J. Torrey) W. Trelease (‘Y. whip- 
ple?) (Rep. (Annual) Missouri Bot. Gard. 4: 
208. 1893) (Yucca whipplei J. Torrey) 

Yucca [unranked] Hesperoyucca G. Engelmann in 
S. Watson, U.S. Geol. Surv. 40th Parallel, Bot. 
497. Sep—Dec 1871. 

PHAN.-AGAVACEAE (104) 06 Nov 2001 


GREENHOUSE AND STROTHER: HESPEROYUCCA 21 


In December 2001, after further bandying, the 
ING website entry was changed to: 


Hesperoyucca (G. Engelmann) W. Trelease, Rep. 
(Annual) Missouri Bot. Gard. 4: 208. 9 Mar 
1893. 

T.: H. whipplei (J. Torrey) W. Trelease (‘Y. whip- 
pler) (Yucca whipplei J. Torrey) 

Yucca [unranked] Hesperoyucca G. Engelmann in 
S. Watson, U.S. Geol. Surv. 40th Parallel, Bot. 
497. Sep-Dec 1871._For the publication date, 
see I.c. (Rep.) 5: 3. 1894. 

PHAN.-AGAVACEAE (104) 14 Dec 2001 


ACKNOWLEDGMENTS 


This study resulted from work on the Jepson Flora Pro- 
ject and was supported in part by a grant from the William 
R. Hewlett Revocable Trust. We appreciate comments and 
opinions sent to us by e-mail from farflung colleagues. 
We thank D. J. Bogler and S. Verhoek for formal reviews 
of the manuscript and K. H. Clary for informal comments 
on an early version of this paper. 


LITERATURE CITED 


BAKER, J. G. 1876. New garden plants. Gard. Chron. n.s. 
6:196—197. 

. 1892. CCXXIII-Agaves and arborescent Lili- 
aceae on the Riviera. Bull. Misc. Inform. [Kew] 61: 
5-10. 

BOoGLER, D. J. AND B. B. Simpson. 1995. A chloroplast 
DNA study of the Agavaceae. Syst. Bot. 20:191—205. 

CLARY, K. H. AND B. B. Stmpson. 1995. Systematics and 
character evolution of the genus Yucca L. (Agava- 
ceae): Evidence from morphology and molecular 
analyses. Bol. Soc. Bot. México 56:77-88. 

ENGELMANN, G. 1871. Appendix (pp. 496—497) to S. Wat- 
son, List [Catalogue] of plants collected in Nevada 
and Utah, in S. Watson et al., Botany (Fortieth Par- 
allel). 

. 1873. Notes on the genus Yucca. Trans. St. Louis 

Acad. Sci. 3:17—54. 

. 1875. Notes on the genus Yucca. No. 2. Trans. 
St. Louis Acad. Sci. 3:210—214. 

FarR, E. R., J. A. LEUSSINK, AND E A. STAFLEU. 1979. 
Index nominum genericorum (plantarum), Vol. 2, 
Eprolithus—Peersia. Regnum Veg. 101:631—1276. 

TORREY, J. 1859 [1858]. Jn W. H. Emory, Report on the 
United States and Mexican boundary survey, Vol. 2, 
pp. [27]—270, Botany of the boundary. 

. 1861. In J. C. Ives, Report upon the Colorado 
River, part 4, pp. [1]—30, Catalogue of the plants col- 
lected upon the expedition. 

TRELEASE, W. 1893. Further studies of yuccas and their 
pollination. Rep. (Annual) Missouri Bot. Gard. 4: 
181-226. 


MApRONO, Vol. 49, No. 1, pp. 22-24, 2002 


SYMPATRY BETWEEN DESERT MALLOW, EREMALCHE EXILIS AND 
KERN MALLOW, E. KERNENSIS (MALVACEAE): MOLECULAR 
AND MORPHOLOGICAL PERSPECTIVES 


KATARINA ANDREASEN 
Laboratory of Molecular Systematics, Swedish Museum of Natural History, 
Stockholm, Sweden; 
Jepson Herbarium and Department of Integrative Biology, University of 
California, Berkeley CA 94720-2465 


ELLEN A. CYPHER 
Endangered Species Recovery Program, 1900 North Gateway Boulevard, 
Suite 101, Fresno, CA 93727 


BRUCE G. BALDWIN 
Jepson Herbarium and Department of Integrative Biology, 
University of California, Berkeley CA 94720-2465 


ABSTRACT 


Molecular and morphological data support an extended distribution of Eremalche exilis into the south- 
ern San Joaquin Valley and southern Inner South Coast Ranges of California, within the range of the 
morphologically similar, rare, and endangered E. kernensis. Nuclear rDNA sequences for plants from 
Kern County that were morphologically indistinguishable from E. exilis were identical at all non-poly- 
morphic sites to sequences of undisputed EF. exilis from the Mojave Desert and Sonoran Desert, but 
differed from sequences of E. kernensis by seven sequence mutations. Morphologically, E. exilis can be 
distinguished from E. kernensis by anther position—in flowers of EF. exilis, the anthers are held at the 
same level as the stigmas; in bisexual flowers of E. kernensis, the anthers are situated well below the 
stigmas. Based on limited rDNA data, we found no evidence of gene flow between sympatric populations 


of E. exilis and E. kernensis. 


Key words: Eremalche, Malvaceae, California floristics, ITS, ETS, rare and endangered plants. 


Botanists have disagreed about whether the de- 
sert mallow, Eremalche exilis (A. Gray) Greene, 
occurs within the California Floristic Province 
(Kearney 1956; Twisselmann 1956, 1967; Hoover 
1970; Bates 1992). In The Jepson Manual, Bates 
(1993) indicated that the distribution of E. exilis in 
California is restricted to the Mojave and Sonoran 
Deserts, whereas Twisselmann (1956, 1967) and 
Hoover (1970) indicated that £. exilis occurs in the 
San Joaquin Valley and Inner South Coast Ranges 
of western Kern County and eastern San Luis Obis- 
po County, California. If E. exilis is present in the 
southern San Joaquin Valley and southern Inner 
South Coast Ranges of California, then it may be 
of importance for evaluating the origin and status 
of the rare and endangered Kern mallow, E. ker- 
nensis C. B. Wolfe [=E. parryi subsp. kernensis (C. 
B. Wolfe) D. M. Bates pro parte; Bates (1992) ex- 
panded the circumscription of Kern mallow], which 
is endemic to the region. Eremalche kernensis has 
been suggested to be of hybrid origin between E. 
exilis and the widespread central Californian taxon 
E. parryi (Greene) Greene (Kearney 1956). A co- 
occurrence of these taxa would be relevant to that 
hypothesis. Here we present molecular and mor- 
phological data that support an extended distribu- 


tion of E. exilis into the southern San Joaquin Val- 
ley and southern Inner South Coast Ranges of Cal- 
ifornia, within the range of E. kernensis. 


MOLECULAR DATA 


Plants from Kern County that we could not dis- 
tinguish morphologically from E. exilis were in- 
cluded in an ongoing project that aims to clarify 
whether E. kernensis is evolutionarily distinct from 
E. parryi (i.e., that the two taxa are independent, 
non-interbreeding, evolutionary lineages). Individ- 
uals from two populations of putative E. exilis from 
western Kern County were sampled for the molec- 
ular analysis. One specimen (Cypher 99-1A) was 
collected in the Lokern Natural Area (T29S, R22E, 
sect. 29 SE% of NW% of SE%, MDM), and was 
growing intermixed with the rare and endangered 
Kern mallow, E. kernensis. The other specimen 
(Vanherweg 99-14) was collected on a hillside west 
of Derby Acres (T31S, R22E, sect. 10 E% of NW%, 
MDM). In addition, one sample of E. exilis from 
the Mojave Desert (Heckard 4508) as well as sey- 
eral samples of E. kernensis were included. The 
DNA was extracted from pressed and dried leaf 
material of individual plants and the Internal and 
3'-External Transcribed Spacers (ITS and ETS) in 


2002] 


ANDREASEN ET AL.: SYMPATRY OF EREMALCHE EXILIS AND E. KERNENSIS 23 


TABLE 1 WARIABLE NUCLEOTIDE POSITIONS IN THE INTERNAL (ITS) AND EXTERNAL (ETS) TRANSCRIBED SPACER REGIONS 
OF E. EXILIS AND E. KERNENSIS. Vouchers are deposited at JEPS and sequences at EMBL. EC = E. Cypher; BV = B. 
Vanherweg; SB = San Bernardino Co.; * = non-identical nucleotide positions between E. exilis and E. kernensis; Each 


number corresponds to one DNA position and polymorphic positions are abbreviated: s = cg; y = 


ct; w = at; k = gt; 


r = ag; m = ac. ? = not sequenced. — = gap; NN = ac/-. 

Taxon Collection data Accession numbers ITS (1-646) and ETS (710-1152) positions 
ITS/ETS hes 

a3 a tah cat eee lL al alicia Galea aL 
17111111711222444667778889999900001111 
QL Da sy7/ 23S SSO IL 3 213} 7) 7/ QOL AG 10S} 335) 
91205885908487260955567839462153782 
E. exilis Kern EC 99-1A AJ416060/AJ4 16065 eget CGECagebacgeqegqgcEgtecEEGcrtacs 
E. exilis Kern BV 99-]4 AJ416062/AJ4 16067 cgtcctcagctacgceg?gggctgtccttycwtacs 
E. exilis SB Heckard 4508 AJ416061/AJ416066 catcctyagctacgmgmgkrctrtcmttyctyacg 
E. kernensis Kern EC 99-3 AJ416063/AJ416064 yoyyyycmrmeryrcc ?ngg--gctcwgcgttNNg 


the nuclear ribosomal DNA were amplified and se- 
quenced as described by Andreasen and Baldwin 
(2001). 

ITS and ETS sequences for plants identified on 
the basis of morphology as E. exilis from western 
Kern County were identical to the ITS and ETS 
sequences for indisputed E. exilis from the Mojave 
Desert, except at twelve nucleotide positions (Table 
1). At eleven of these positions one or two of the 
E. exilis samples were polymorphic, with one ad- 
ditional nucleotide besides the one present in the 
non-polymorphic sample(s). At one position, E. ex- 
ilis from the Mojave Desert had a unique state (po- 
sition 111). ETS and ITS sequences of Eremalche 
kernensis (one sample sequence is shown in Table 
1) were polymorphic for many positions but not for 
the same positions that were polymorphic in the 
samples of E. exilis. In addition to the polymorphic 
positions, FE. exilis samples and E. kernensis dif- 
fered at eight nucleotide positions in the ETS and 
ITS sequences. In the ITS region, the two taxa dif- 
fered by two point mutations (positions 268 and 
646). In the ETS, E. exilis and E. kernensis differed 
by four point mutations (positions 938, 973, 1006, 
and 1021) and E. exilis was marked by a two base 
pair insertion (at positions 825 and 826). This in- 
Sertion was unique for E. exilis and was not present 
in E. rotundifolia or any other sequenced taxon 
from tribe Malveae (Andreasen and Baldwin 2001, 
unpublished data). In addition, ETS sequences of 
E. kernensis were polymorphic for a two base-pair 
indel (ac/-; NN in Table 1) at positions 1137 and 
1138. All other Malveae sequences had ‘“‘ac”’ at 
these positions. Phylogenetic analysis of ETS and 
ITS sequences for each of the taxa of Eremalche 
and various outgroup taxa in tribe Malveae af- 
firmed that samples of E. exilis from the Mojave 
Desert and western Kern County constitute a line- 
age (data not shown). 


MORPHOLOGICAL AND DISTRIBUTIONAL DATA 


Overlapping variation in flower color, flower 
size, and growth form has contributed to uncertain- 


ty about the identity of Eremalche populations in 
the southern San Joaquin Valley and southern Inner 
South Coast Ranges. Both E. exilis and E. kernensis 
most commonly have white petals, although petals 
of E. exilis may be tinged with pink and petals of 
E. kernensis occasionally are pale lavender (Wolf 
1938; Kearney 1956; Cypher unpublished data). 
Keys to species of Eremalche typically have used 
petal and calyx lengths to distinguish E. exilis from 
E. kernensis and E. parryi (e.g., Kearney 1956; 
Munz 1968; Bates 1993). However, these keys 
failed to differentiate petal length by gender. Ere- 
malche exilis has only bisexual flowers, whereas E. 
kernensis is gynodioecious, 1.e., with bisexual-flow- 
ered plants and pistillate-flowered plants (Bates 
1992, 1993). In E. kernensis, petals of pistillate 
plants are shorter (3 to 7 mm long) than petals of 
bisexual plants (4.5 to 10 mm long) (Cypher un- 
published data). Pistillate plants of E. kernensis 
have petals that often are similar in size to those of 
E. exilis, which has petals ranging in length from 
4 to 5.5 mm (Bates 1993). 

For the most part, pistillate flowers of E. kernen- 
sis have only styles and no stamens, but in a small 
percentage of flowers, there may be a few (less than 
five, Cypher unpublished data) vestigial stamens 
that are reduced in size and do not shed pollen. The 
absence of stamens (or, more infrequently, the pres- 
ence of few, vestigial, non-functional stamens) in 
small-flowered (pistillate) plants of E. kernensis 
should be sufficient to distinguish morphologically 
E. kernensis from E. exilis, but the differing posi- 
tion of the anthers relative to the stigmas in the two 
taxa has caused confusion. In £. exilis, the anthers 
are held at the same level as the stigmas, whereas 
in bisexual flowers of E. kernensis the anthers are 
situated well below the stigmas, near the base of 
the corolla. Thus, bisexual flowers of E. exilis have 
been mistaken for pistillate flowers of E. kernensis, 
when anthers at the same level as the stigmas have 
been interpreted as stigmas. The growth form of E. 
exilis has been described as prostrate or decumbent 
(Munz 1968; Bates 1993), whereas Eremalche ker- 


24 MADRONO 


nensis stems may be erect or prostrate (Wolf 1938; 
Munz 1968). We have observed erect, slender- 
stemmed plants of E. exilis in western Kern Coun- 
ty, as well as individuals of both E. exilis and E. 
kernensis that have a stout, erect central stem and 
several prostrate lateral stems. 

Investigations in the Lokern area of western 
Kern County (west of the community of Button- 
willow) from 1996 through 2001 revealed that E. 
kernensis does not range more than 3.2 km south 
of Lokern Road. The southernmost occurrences of 
E. kernensis that we have confirmed are in T29S, 
R22E, sect. 27, 28, and 29, MDM. Based on motr- 
phological characters, we have confirmed the pres- 
ence of E. exilis in T29S, R22E, sect. 14, 19, 20, 
a), Pe, Dy 2S, 2, SZ, BG! BS, IMIDE im WIOS, 
R22E, sect. 4; and in T31S, R22E, sect. 10, MDM. 
Eremalche exilis and E. kernensis are sympatric in 
at least T29S, R22E, sect. 14, 20, 23, 27, 28, and 
29, MDM, sometimes growing in mixed colonies. 

In addition to the above mentioned localities we 
have identified four collections (at UC) from the 
southern Inner South Coast Ranges and southern 
San Joaquin Valley of California that match the de- 
scription of E. exilis: (1) Hoover 9350, collected in 
1965 in San Luis Obispo County (“Hill just east of 
San Juan River, La Panza District, in sandy calcar- 
eous soil’’); (2) Hoover 441, collected in 1935 in 
Fresno County (“10 mi e. of Coalinga’’); (3) Alice 
Eastwood s. n., from “Huron, Calif.’’ in Fresno 
County (this collection is from 1893, before the 
Huron vicinity was converted to farmland); and (4) 
Dean Wm. Taylor 10171, collected in 1989 in Kern 
County (“‘just west of California Aqueduct ca. 1.3 
miles SE of Highway 58’’). The fourth collection 
(Dean Wm. Taylor 10171) yielded ETS and ITS 
sequences that were identical to ETS and ITS se- 
quences of one of the FE. exilis specimens from 
Kern County described above (Cypher 99-1A) ex- 
cept at ETS position 1152, where DWT 1/0171] had 
a “g’ and Cypher 99-1A had a “cg.” 


CONCLUSIONS 


The morphological and molecular data presented 
here lead us to suggest that Eremalche exilis has a 
wider distribution in California than the distribution 
reported in The Jepson Manual (Bates 1993). The 
distribution of E. exilis should be extended beyond 
the Mojave Desert and Sonoran Desert to include 
the California Floristic Province (southern San Joa- 
quin Valley and Inner South Coast Ranges) in Kern 
and San Luis Obispo counties. Since there are no 
known recent collections of E. exilis from Fresno 
County it is uncertain if it still occurs there. The 


[Vol. 49 


habitat destruction in the San Joaquin Valley sug- 
gests that E. exilis is likely to have disappeared 
from the Huron area. In contrast to the extended 
distribution of EF. exilis, the range of the rare and 
endangered FE. kernensis may be even narrower 
than previously believed because of earlier mis- 
identifications of E. exilis as E. kernensis. 

The molecular data demonstrate that E. exilis and 
E. kernensis are genetically and evolutionarily dis- 
tinct entities. Seven sequence differences between 
them were found, including one insertion unique 
for E. exilis. These positions are not polymorphic 
in either taxon, contrary to the expectation for hy- 
bridizing taxa. We found no evidence of gene flow 
between E. exilis and E. kernensis in rDNA se- 
quences of plants collected from the one area of 
sympatry sampled in this study. The previously 
proposed hybrid origin of E. kernensis and its ge- 
netic distinctness from EF. parryi is under continu- 
ing investigation and will be addressed in a later 


paper. 


ACKNOWLEDGMENTS 


We thank B. Vanherweg for collecting one of the Ere- 
malche exilis specimens included in this investigation. We 
also thank B. Wessa for DNA sequencing of one of the 
E. exilis samples (DWT 10171). This study was financed 
by a grant from the US Fish and Wildlife Service for a 
biosystematic study of Kern mallow. 


LITERATURE CITED 


ANDREASEN, K. and B. G. BALDWIN. 2001. Unequal evo- 
lutionary rates between annual and perennial lineages 
of checker mallows (Sidalcea, Malvaceae): Evidence 
from 18S—26S rDNA internal and external tran- 
scribed spacers. Molecular Biology and Evolution 18: 
936-944. 

Bates, D. M. 1992. Gynodioecy, endangerment, and sta- 
tus of Eremalche kernensis (Malvaceae). Phytologia 
72:48—54. 

. 1993. Eremalche. P. 748 in J. C. Hickman (ed.), 
The Jepson manual: Higher plants of California. Uni- 
versity of California Press, Berkeley. 

Hoover, R. FE 1970. The vascular plants of San Luis Obis- 
po County, California. University of California Press, 
Berkeley. 

KEARNEY, T. H. 1956. Notes on Malvaceae. VIII. Ere- 
malche. Madrofio 13:241—243. 

Munz, P. A. 1968. A California flora and supplement. Uni- 
versity of California Press, Berkeley. 

TWISSELMANN, E. C. 1956. A flora of the Temblor Range 
and the neighboring part of the San Joaquin Valley. 
The Wasmann Journal of Biology 14:161—300. 

. 1967. A flora of Kern County, California. The 
Wasmann Journal of Biology 25:1—393. 

WoLF, C. B. 1938. California plant notes II. Occasional 
Papers Rancho Santa Ana Botanic Garden, Series I, 
2:44—-90. 


MaproNo, Vol. 49, No. 1, pp. 25-32, 2002 


POLLINATION OF CYTISUS SCOPARIUS (FABACEAE) AND 
GENISTA MONSPESSULANA (FABACEAE), TWO INVASIVE 
SHRUBS IN CALIFORNIA 


INGRID M. PARKER! AND ALEXANDRA ENGEL 
Department of Ecology and Evolutionary Biology, University of California, 
Santa Cruz, CA 95064 


KAREN A. HAUBENSAK 
Department of Integrative Biology, University of California, Berkeley, CA 94720 


KAREN GOODELL 
Department of Ecology, Evolution, and Natural Resources, Rutgers University, 


New Brunswick, NJ 08903 


ABSTRACT 


Mutualistic interactions between natives and non-natives, and between different introduced species, can 
play an important role in the invasion process. The facilitation of a new introduced species by a previous 
invader could either accelerate an invasion or exacerbate its impact, providing a positive feedback loop 
in heavily invaded ecosystems. Open grasslands in Marin County, CA, are being invaded by two closely 
related, introduced legumes, Cytisus scoparius (Scotch broom) and Genista monspessulana (French 
broom). These non-clonal shrubs have been shown to be non-autogamous and pollen limited, underscoring 
the potential importance of pollinators to their fecundity and spread. The flowers of both are fused shut 
and require forced “tripping” by a pollinator. We measured floral characters and pollen production to 
make predictions about which species would be most attractive, and most accessible, to bee visitors. 
Cytisus flowers were an order of magnitude larger and produced four times as many pollen grains, 
suggesting that they should be more attractive and rewarding than Genista flowers. However, Cytisus 
flowers also required significantly more force to open, suggesting that less powerful pollinator species 
might be excluded from visiting. We tested these predictions by quantifying visitation rates and directly 
observing pollinators at two sites where the invaders co-occur. Consistent with the mechanical assay, 
pollinators were more successful at accessing flowers of the small-flowered Genista than the large-flow- 
ered Cytisus; however, Cytisus was more frequently visited than Genista, suggesting that pollinators 
preferred the larger and more rewarding species. We did not find evidence to support the prediction that 
the small-flowered Genista was accessible to a greater diversity of pollinator species. Although introduced 
from the same native range as the two plant invaders, honey bees (Apis mellifera) were not “‘better”’ 
pollinators than native bumble bees (Bombus vosnesenskii) in terms of effectiveness at tripping flowers 
or the number of flowers visited per plant. However, Apis was the numerically dominant pollinator at 
both sites, underscoring the potential conservation implications of local Apis introduction for the spread 


of noxious weeds in natural ecosystems. 


Key words: exotic, introduced, alien, mutualism, plant-pollinator, invasional meltdown 


INTRODUCTION 


Biological invasions have become one of our 
most alarming conservation issues (Williamson 
1996; Parker and Reichard 1998; Mooney and 
Hobbs 2000). Negative interactions such as com- 
petition and predation between introduced and na- 
tive species (“‘biotic resistance’’) have long been 
thought to play a central role in determining wheth- 
er an invasion will succeed or fail (Darwin 1865; 
Elton 1958; Levine 2000; Maron and Vila 2001). 
However, mutualistic interactions between natives 
and non-natives may also be important in the in- 
vasion process (Simberloff and Von Holle 1999; 
Richardson et al. 2000). Interactions between dif- 


‘Author to whom correspondence should be addressed. 
Email: parker @biology.ucsc.edu 


ferent introduced species can be important too, as 
suggested by the idea of “‘invasional meltdown” 
(Simberloff and Von Holle 1999; Parker et al. 
1999), where the facilitation of a new introduced 
species by a previous invader could either accel- 
erate an invasion or exacerbate its impact, provid- 
ing a positive feedback loop in heavily invaded 
ecosystems. For example, spread of the nitrogen- 
fixing tree Myrica faya in Hawai’i appears to occur 
primarily through dispersal by an introduced bird 
species, Zosterops japonica (Vitousek and Walker 
1989). 

The interaction between plants and their polli- 
nators is an example of a mutualism that can exert 
strong control over the fitness of the interacting 
partners and is thought to have driven the evolution 
of many floral traits (Darwin 1877; van der Pijl 
1961). Pollinator limitation of plant reproduction is 


26 MADRONO 


common (Burd 1994) and could be especially im- 
portant for introduced species. A successful inva- 
sive plant must be able to colonize new territory, 
usually in small numbers, and in the absence of its 
original pollinators. Such a scenario would appear 
to favor species that are self-fertile and capable of 
autonomous self-pollination, or autogamy (Allard 
1965; Baker 1965; Brown and Marshall 1981; Bar- 
rett 2000). 

Open grasslands in Marin County, California, are 
being invaded by two closely related European 
shrubs, Cytisus scoparius (L.) Link (Scotch broom) 
and Genista monspessulana (L.) L. Johnson 
(French broom). Both species were first introduced 
to California as ornamentals in the 1860’s and 
1870’s (Hoshovsky 1986). Cytisus and Genista are 
non-clonal legumes, are very similar in appearance, 
and are often lumped together in discussions of 
their ecological role as invaders (McClintock 1985; 
Hoshovsky 1986). Despite the above prediction that 
successful invaders should be self-fertile and au- 
togamous, in previous work we found that neither 
of these species showed substantial autogamy when 
pollinators were excluded, and that both exhibited 
reduced seed set when fertilized with self pollen 
(Parker and Haubensak 2002). We also found that 
both species were significantly pollen limited in 
populations on the Marin Peninsula (Parker and 
Haubensak 2002). These results demonstrate that 
the abundance and behavior of pollinators have im- 
portant consequences for the reproductive success 
of both these invaders. At the same time, however, 
there are apparent differences in floral biology be- 
tween the two species which we predicted could 
differentially affect their pollination success. Flow- 
ers of Cytisus appear larger than those of Genista. 
Reproductive plants of both species vary in size 
from 0.5 m to >2 m, so although Genista produces 
more of its small flowers for a branch of the same 
size (Parker and Haubensak, unpublished data), 
both plants can have from several dozen to several 
thousand yellow flowers. Therefore the primary dif- 
ference in floral display that distinguishes the spe- 
cies appears to be flower size. Neither plant pro- 
duces nectar, so pollen is the only reward for insect 
visitors. 

Theoretical expectations led us to pose two al- 
ternative hypotheses about how differences be- 
tween the two species could influence their repro- 
ductive success. The first hypothesis involves the 
ease of handling of flowers by pollinators. Larger 
flowers like those of Cytisus may be more difficult 
to handle, preventing the access of certain insect 
species and leading to specialization on a narrower 
set of pollinators. Both Cytisus and Genista have 
typical papilionaceous legume flowers, but with 
fused keel petals that an insect visitor must split in 
order to release the style and anthers. Flower visi- 
tors may have to apply more force to trip larger 
flowers (Westerkamp 1997), which could restrict 


[Vol. 49 


the diversity and number of visitors to the larger 
Cytisus flowers. 

The second, alternative, hypothesis is that the 
larger Cytisus flowers are more attractive and re- 
warding than the smaller Genista flowers. Pollina- 
tors often prefer larger flowers or flowering dis- 
plays (Campbell 1989; Ohara and Higashi 1994; 
Schemske and Agren 1995), or larger rewards such 
as nectar or pollen (e.g., Neiland and Wilcock 
1998; Robertson et al. 1999). Flower size can be 
correlated with nectar production (Brink and de 
Wet 1980; Harder and Cruzan 1990; Cresswell and 
Galen 1991), although little is currently known 
about how flower size correlates with pollen quan- 
tity or quality among related species. A large-flow- 
ered species like Cytisus should attract more pol- 
linators than an otherwise similar, small-flowered 
species, especially if it offers larger pollen rewards. 

We measured floral characters and pollen pro- 
duction in Cytisus and Genista. Armed with infor- 
mation on the basic floral biology of the two spe- 
cies, we then tested whether patterns in pollinator 
visitation, the identity and diversity of visiting in- 
sect species, and pollinator behavior were consis- 
tent with the first or second hypothesis. The pur- 
pose of this study, then, was: 1) to investigate floral 
traits that may influence attraction, specialization, 
and pollinator effectiveness; 2) to determine wheth- 
er visitation rates to these two plant invaders were 
consistent with predictions based on their floral 
characters; 3) to determine the identity and origin 
(native vs. non-native) of insect species pollinating 
Cytisus and Genista in central California, and 4) to 
compare the relative “‘quality”’ of pollinators from 
the introduced range vs. the native range of the in- 
vaders. 


METHODS 
Study Sites 


We conducted our study at two sites in Marin 
County, CA. The Mt. Tamalpais site is along the 
eastern edge of Mount Tamalpais State Park, on an 
exposed, west-facing slope. This site was dominat- 
ed by non-native grasses, with patches of Baccharis 
pilularis, Vinca major, Cotoneaster pannosa, Plan- 
tago lanceolata, Rubus laciniatus, and Conium ma- 
culatum. Cytisus and Genista grew interleaved, 
with Cytisus individuals somewhat more sparse and 
spread out than Genista. Overall cover of Cytisus 
and Genista at the site was approximately 15% and 
30%, respectively. 

The China Camp site is located 12 km away from 
the first site, near China Camp State Park. The site 
is located in a wide ravine, with mixed evergreen 
woodland on one side and a steep, NE-facing slope 
on the other side. Cytisus and Genista grow in 
patches and as scattered individuals on the ravine 
bottom and up the NE-facing slope, with a total 
cover of approximately 15% and 35%, respectively. 
Other common species included Heteromeles ar- 


2002] 
ia ap 
, thes 
IE, 
Fic. 1. Schematic to illustrate size measurements con- 


ducted on the flowers of Cytisus and Genista. Flower vol- 
ume was approximated from a polyhedron using length 
and width of the keel and height of banner petal. 


butifolia, Toxicodendron diversilobum, Cortaderia 
selloana, Baccharis pilularis, Carduus nutans, and 
non-native grasses. 

Temperature probes at each site recorded air tem- 
peratures at 15 min intervals from April 2 through 
May 31. The two sites were similar for daily max- 
imum, minimum, and mean termperatures between 
9 am and 6 pm. For example, the average daily 
temperature (CC) was 21.6 in April and 23.1 in May 
at China Camp, and 21.3 in April and 24.4 in May 
at Mt. Tamalpais. 


Floral Characters: Size, Pollen Grain Number, 
and Accessibility 


We compared the two species for aspects of their 
floral biology important to pollinator attraction, re- 
ward, and utilization. To compare flower sizes, we 
took three measurements per flower (n = 10 flow- 
ers for Genista, n = 14 for Cytisus): length of the 
Keel petal, width of the keel petal, and height of 
the banner petal. The approximate volume of each 
flower was calculated as a polyhedron with the ban- 
ner and keel acting as two planes at right angles, 
with a third connecting plane (Fig. 1). 

To quantify pollen production, we collected a 
single, unopened bud from each of 16 Cytisus (6 
Mt. Tamalpais, 10 China Camp) and 19 Genista (7 
Mt. Tamalpais, 12 China Camp) plants. We sus- 
pended and sonicated fully dehisced anthers in a 
1% saline solution. We then estimated the number 
of pollen grains per flower using an Elzone 280-PC 
electronic particle counter (Micromeritics, Nor- 
cross, GA). 

Larger flowers might be more difficult to trip 
open, limiting access to the flower by smaller pol- 
linators. To quantify the force required to open the 
flowers, we hung paper clips sequentially from the 
keel petals until the flower was tripped open. The 
paperclips were then weighed and the total mass 
converted to force. Unpaired t-tests were used to 
compare the two species for all floral characters. 


Visitation Rates to Cytisus and Genista 


Flowers can be tripped only once, after which 
they remain in an open position. In order to deter- 
mine the proportion of flowers tripped (an index of 
Visitation), we marked branches on five individuals 


PARKER ET AL.: POLLINATION OF CYTISUS AND GENISTA ZT 


of each species at the two sites. From March 19 
through May 21, at one- to two-week intervals, we 
examined every flower on each branch, recording 
the number of tripped and untripped flowers. We 
then marked every open flower with Wite-Out™ to 
avoid recounting the same flowers at the next cen- 
sus. Wite-Out@ had no negative effect on fruit pro- 
duction (Parker and Haubensak 2002). Some un- 
tripped flowers may have been tripped after being 
marked and counted (as untripped), leading to an 
underestimate of the tripping rate. However, we 
chose this method over the alternative (regular cen- 
suses without marking flowers), because floral lon- 
gevity differs between tripped and untripped flow- 
ers, leading to the possibility that untripped flowers 
would be double-counted more often than tripped 
flowers, if not marked. We summed the number of 
tripped flowers and untripped flowers over the en- 
tire flowering season (mean flowers per plant = 55 
for Cytisus, 153 for Genista), then calculated the 
total proportion tripped. We used ANOVA to test 
for differences among plant species and sites, with 
both species and site treated as fixed effects. For 
comparison, we plotted the data alongside results 
from a similar study done in 1997 (Parker and Hau- 
bensak 2002). 


Pollinator Observations 


Throughout the flowering season of 2000 (March 
19—May 21), we observed pollinators of Cytisus 
and Genista at the two sites. The two sites were 
visited alternately, either on the same day or on 
consecutive days, and observations were made dur- 
ing sunny, warm conditions between 9:00 a.m. and 
6:00 p.m. In all, we completed 72 hours of obser- 
vation (36 hours at each site), over 17 days. To 
maximize our sample size under low visitation 
rates, we walked haphazard (non-random) transects 
through the sites, watching and listening for flying 
insects. We focused on areas where we could see 
both plant species from the transect. A three-way 
(pollinator, plant, site) log linear model was used to 
test for homogeneity of preferences of the two pol- 
linators (Sokal and Rohlf 1995). 

For each pollinator, we recorded each attempt to 
trip a flower, whether that flower was successfully 
tripped, whether the next flower visited was on the 
same or a different individual plant, and the dis- 
tance to the subsequent plant. We calculated the 
proportion of flowers tripped and the mean number 
of flowers per plant for each pollinator observed, 
then averaged these across pollinators for each pol- 
linator type and each plant species. Only two Bom- 
bus individuals were observed visiting Cytisus, 
therefore we could not do a full, two-way analysis 
using plant species and insect species as factors. 
Instead, we compared the two plant species for the 
behavior of Apis only, and compared the two pol- 
linator species for Genista only. Because the vari- 
ables (proportion successfully tripped and number 


DR MADRONO 


2000 + - - 

gin (a) 
& 1600 - Me 
S : ) 
oO 
iS 1200 + 
= 
fo) 
> 
5 800 - | | 
= | 
2 | | 
Fo 4004 | | 

0 a 

| (b) | 

© 100000 + | 
3 | | 
é | | 
= S00004 | | 
o | 
2 ) | 
Z 60000 ~ 
‘3S | 
5p | | 
= 40000 | | | 
2 

(0) = — 2 

Cynsus Genista 
Fic. 2. Floral traits for Cytisus and Genista: (a) Mean 


estimated flower volume (n = 14 and 10 for Cytisus and 
Genista, respectively) and (b) mean number of pollen 
grains per flower (n = 16 and 19). Error bars represent 1 
SE. 


of flowers tripped per plant) were not normally dis- 
tributed, we analyzed the data using nonparametric 
Mann-Whitney U tests. 

We collected specimens of each pollinator spe- 
cies and identified them following Thorp et al. 
(1983). 


RESULTS 


Flower Size, Pollen Grain Number, 
and Accessibility 


Cytisus flowers were significantly larger than 
Genista flowers; length of the keel petal (mm + 1 
SD) was 18.8 + 1.4 compared to 6.7 + 0.5 for 
Genista (df = 22, t = 27.1, P < 0.0001). The es- 
timated volume of Cytisus flowers (1450 mm? + 
364 SD) was an order of magnitude larger than that 


0.035 + — 


0.030 - ] 


00254 | 
0.020 - 
0.015 - 


0.010 5 
| 
| 


0.000 -—1— 


0.005 + 


Force required to trip flower (Newtons) 


Genista 


Cyrsus 


Fic. 3. Force required to access (trip open) flowers of 
Cytisus and Genista, determined with a mechanical assay. 
Error bars represent 1 SE. 


[Vol. 49 
IO = 5 oa 
VISUS 
Bo Genista a) 2000 

08 4 é 

06 5 
ws) OA 4 
oO 
a. | 
= 
= O24 
o 
3 
© 
rm 00 
az 4 
fo) 
& b) 1997 
& (2 = a 
a. 
2 
Ou 

06 5 | e 

| = 
O04 - 
02 4 : 
| 
09 +1 . 
China Camp Mt. Tamalpais 


Fic. 4. Proportion of flowers tripped for Cytisus (open 
bars) and Genista (filled bars) in two populations. a) Data 
from 2000, collected from 5 individuals per population; 
b) Data from 1997, collected from 40—60 individuals per 
population. Data from 2000 redrawn from Parker and 
Haubensak (2002). Error bars represent 1 SE. 


for Genista-(i32 = 27; Bigs 2:'dti = 2207 — isles 
P < 0.0001). Cytisus produced more than four 
times as many pollen grains per flower as Genista 
(Fig. 2, df = 33, t = 8.0, P < 0.0001). 

In the mechanical assay, the two species required 
significantly different amounts of force to trip the 
flowers (df = 56, t = 2.80, P = 0.007). We used 
more than twice as much force in opening flowers 
of Cytisus (0.026 N + 0.005 SD) as in opening 
flowers of Genista (0.013 + 0.006) (Fig. 3). 


Visitation Rates to Cytisus and Genista 


At both sites, large-flowered Cytisus had a sig- 
nificantly higher proportion of tripped flowers than 
did small-flowered Genista (Fig. 4a, Table 1). 
These results from the 2000 season confirmed the 
same pattern seen in 1997 (Fig. 4b, Parker and 
Haubensak 2002). Therefore, the greater pollination 
success of Cytisus seems to be generalizable over 
both space and time. There was no significant main 
effect of site (Table 1). Rather, for Cytisus, more 


TABLE 1. ANALYSIS OF WARIANCE OF PROPORTION OF 
FLOWERS TRIPPED (I.E., WISITED) OVER THE ENTIRE SEASON 
FOR CYTISUS AND GENISTA AT TwoO SITES. 


Source df SS F Pp 
Species 1 0.80 61.1 <0.0001 
Site 1 0.01 0.53 0.48 
Species X Site 1 0.07 5.07 0.039 
Residual 16 0.21 


2002] 


TABLE 2. NUMBERS OF POLLINATORS OBSERVED IN 72 
HOURS OF OBSERVATION AT TwoO SITES ON THE MARIN PEN- 
INSULA. Observations were done along haphazard transects 
through the sites, in areas where both plant species were 
within visual and auditory range. 


Mt. Tamalpais China Camp 


Insect species Cytisus Genista Cytisus Genista 


Apis mellifera 19 3} q. 12 
Bombus vosnesenskii 1 i>) 1 0 
Xylocopa californica 0) 1 0 0 
Total 20 19 8 12 


flowers were tripped at China Camp, while for Gen- 
ista, more flowers were tripped at Mt. Tamalpais 
(Fig. 4a), resulting in a significant population xX 
species interaction effect (Table 1). 


Pollinator Observations 


In 72 hours of observation at the two sites, three 
bee species were seen pollinating Cytisus and Gen- 
ista: Apis mellifera, Bombus vosnesenskii, and Xy- 
locopa californica (Table 2). We observed a total 
of 59 pollinators, which made 342 effective visits 
to flowers (in which the flower was tripped). Given 
that hundreds to thousands of flowers were being 
watched at any one moment, this represents a very 
low pollination rate in terms of visits/flower/hour. 

Apis was the numerically dominant pollinator ob- 
served at both sites, although Genista at Mt. Tam- 
alpais was visited most by Bombus (Table 2). Xy- 
locopa was observed only once and is left out of 
subsequent analyses. Results of the three-way log 
linear model indicated that bee species preferences 
for the two broom species differed across the two 
sites (G[Williams] = 8.18, df = 3, P < 0.005). 
Therefore, we examined these preferences for each 
site separately. At Mt. Tamalpais, Apis primarily 
visited Cytisus, while Bombus primarily visited 
Genista (G[Williams] = 26.48, df = 1, P < 0.001). 
At China Camp, there were too few Bombus visits 
to include in the analysis, but a chi-square test in- 
dicated that Apis had no significant preference for 
Cytisus or Genista (chi-square = 1.32, df = 1, P > 
0.10). Therefore, we did not find evidence to sup- 
port the prediction that the small-flowered Genista 
was accessible to a greater number of pollinator 
species. Nor did we find that the larger pollinators 
(Bombus) tended to visit the larger flowers (Cyti- 
SUS). 

Before flying to a new individual, bees (all spe- 
cies combined) visited on average 6.6 (+10.6) 
flowers/plant of Cytisus and 3.9 (+3.8) flowers/ 
plant of Genista, a difference that was not statisti- 
cally significant (Z = 0.25, P = 0.80). When con- 
sidered alone, Apis did not differ in its pattern of 
movement on the two plant species (Fig. 5, Z = 
0.22, P = 0.83). Although Apis tended to visit more 
flowers per plant than Bombus (Fig. 5), this differ- 


PARKER ET AL.: POLLINATION OF CYTISUS AND GENISTA 29 


10 


[|] Cynisus 
8 i Genista 


Number of flowers tripped per plant 


Bombus 


Apis 


Fic. 5. Number of flowers successfully visited (tripped) 
per individual plant before flying to a new plant, for Bom- 
bus observed on Cytisus (n = 2) and Genista (n = 15) 
and Apis observed on Cytisus (n = 22) and Genista (n = 
15). Error bars represent 1 SE. 


ence was not Statistically significant (Genista only, 
Z = 1.20, P = 0.23). 

We used Apis visitation success (ratio of flowers 
tripped to flowers attempted) as a measure of the 
accessibility of the two flower types; pollinators 
were more successful at accessing flowers of the 
small-flowered Genista than the large-flowered Cy- 
tisus (Fig. 6, Apis only, Z = 2.13, P = 0.03). The 
proportion of Genista flowers successfully tripped 
did not differ for the two pollinator species (Fig. 6, 
Z = 1.0, P = 0.32). The proportion of Cytisus flow- 
ers successfully tripped appeared to be higher for 
Bombus than for Apis (Fig. 6). This difference was 
statistically significant (Z = 2.09, P = 0.03), but 
the small sample size for Bombus (N = 2) suggests 
caution in interpreting this result. 


DISCUSSION 


Do Floral Characters Predict Which Invasive 
Plants Should Be Most Attractive to Pollinators 
and Which Should Be Most Pollen Limited? 


We started with two alternative hypotheses for 
how floral traits could affect pollinator attraction 
and visitation in this system. The first hypothesis 


(—— 2 ios —— = _— 


| Cytisus 
Bi Cenista 


Prop. of flowers successfully tripped 
oO 
Oo 


Bombus Apis 


Fic. 6. The mean proportion of flower visits attempted 
in which the flower was successfully tripped open, for 
individual Bombus and Apis pollinators observed at flow- 
ers of Cytisus (open bars) or Genista (filled bars). Each 
pollinator observed could involve visits to flowers on one 
plant or multiple plants. Error bars represent 1 SE. 


30 


predicted that because of the unusual “‘tripping”’ 
mechanism in these two species, the larger Cytisus 
flowers could be more difficult for pollinators to 
handle, which would reduce the number of insect 
species capable of visitation. Cytisus would then be 
relatively specialized on larger insects, and polli- 
nator limitation should be more prevalent in Cytisus 
than in Genista (assuming an equal and variable 
pool of pollinator species for the two plants). In 
fact, we did find evidence that Cytisus flowers are 
more difficult to access than Genista flowers, both 
from our mechanical assay and from observations 
of Apis visitation. However, this difference did not 
result in a difference in the pollinator assemblage 
for the two plants. We did not find that small soli- 
tary bees or flies visited Genista, although they are 
common in the area (G. Lebuhn, personal com- 
munication), and we did not find that larger bees 
specialized on Cytisus. Rather, while Apis com- 
monly visited both plant species, the larger Bombus 
and Xylocopa visited primarily Genista. Recent 
work in Great Britain (Stout 2000) found that very 
large Bombus individuals (>20 mm) were less ad- 
ept at accessing Cytisus flowers than were smaller 
Bombus. This suggests, in concert with our results, 
that the largest bees may have trouble handling Cy- 
tisus flowers and may in fact prefer Genista flowers 

While we did not find that the differences in flo- 
ral accessibility between the two invaders translated 
to an effect on pollinator composition in Marin 
County, this effect could be important in other sys- 
tems. For example, one of us (Parker 1997) ob- 
served that Cytisus was rarely successfully tripped 
by Apis in Washington State. Temperature can af- 
fect floral accessibility: Cytisus flowers are more 
difficult to trip in cold compared to warm temper- 
atures in the lab (B. Burley, R. Martin, and K. M. 
Karoly unpublished data). The Washington research 
suggests that in colder climates, the difficulty of 
tripping Cytisus flowers could contribute to pollen 
limitation. 

Our second hypothesis postulated that pollinator 
visitation would be based on floral attractiveness. 
The prediction was that the larger flowers and 
greater pollen rewards of Cytisus could increase its 
attractiveness to pollinators, increasing its success 
in competing with resident plants for pollinator ser- 
vices relative to Genista. We found, both in 1997 
and 2000, that visitation rates to the two plant spe- 
cies were consistent with this hypothesis. Cytisus 
flowers were more frequently tripped in both years 
and at both study sites. Given that Genista tends to 
produce more flowers than Cytisus on plants of the 
same size (Parker and Haubensak unpublished 
data), pollinators in this study appeared to be more 
attracted by the rewards of single flowers than by 
the size of the overall display. 

Genista might have ways of compensating for its 
lower pollinator visitation. Small flowers some- 
times represent reduced allocation to attraction in 
an evolved syndrome including increased selfing 


MADRONO 


[Vol. 49 


and autogamy and lower inbreeding depression 
(Piper et al. 1986; Rathcke and Real 1993; Parker 
et al. 1995; Brunet and Eckert 1998). Genista does 
not exhibit high levels of autogamy (Parker and 
Haubensak 2002). However, compared to Cyfisus, 
it does show less inbreeding depression at seed-set, 
consistent with the syndrome of increased selfing 
and decreased allocation to attractive structures 
(Parker and Haubensak 2002). Thus when pollina- 
tors tend to visit multiple flowers on a plant, Gen- 
ista may be able to take advantage of those visits 
more effectively than Cyfisus. 


Are Plants Better “‘Matched”’ to Pollinators from 
Their Native Range? 


If the floral traits of plants have evolved in re- 
sponse to pollinators in their native range, one 
might expect to see that a pollinator introduced 
from the same region as an introduced plant would 
be a higher quality pollinator, or better match, for 
that plant. We found no evidence to support this 
prediction in our study. Apis mellifera, native to 
Europe, was not better at tripping flowers than the 
locally native Bombus vosnesenskii. We did not fol- 
low the fate of each tripped flower to determine 
relative pollen transfer and fruit set for the different 
pollinators (see Schemske and Horvitz 1984). How- 
ever, the European Apis and native Bombus did not 
differ significantly in their tendency to promote out- 
crossing by moving between plants instead of with- 
in plants. 

Given that most plant-pollinator interactions are 
thought to have arisen from diffuse coevolution of 
guilds rather than coevolution between pairs of spe- 
cies (Jordano 1987; Pellmyr 1992), these findings 
are not particularly surprising. European species of 
Bombus probably occurred throughout the evolu- 
tionary history of these broom species, and may 
have influenced the evolution of their floral mor- 
phology, and North American and European Bom- 
bus are probably similar in terms of their pollina- 
tion value to broom. Although we lack extensive 
information about the degree to which Cytisus and 
Genista are pollinated by Apis in their native range, 
one recent study in Great Britain recorded that 
flowers of Cytisus there were tripped primarily by 
Bombus, while Apis only visited previously tripped 


flowers (Stout 2000). More comparative studies of | 


plant-pollinator interactions in native vs. introduced 
ranges are needed to better understand the role of 
these mutualisms in the invasion process. We are 
aware of only one case in which a highly specific 
pollinator was left behind during invasion (e.g., fig 
wasp invasion in Florida, Nadel et al. 1992). 


Invasional Meltdown? 


Both Cytisus and Genista can be pollen limited 
(Parker 1997; Parker and Haubensak 2002); there- 


fore, attracting pollinators plays a critical role in | 


assuring reproduction. We found that both native 


——SSSEeeee SSS 


2002] 


and non-native insects pollinate these invaders. 
Apis mellifera, native to Europe, was by far the 
dominant visitor at the China Camp site, and was 
also more common than Bombus at Mt. Tamalpais. 
It has recently been suggested that positive inter- 
actions among invaders may commonly facilitate 
new invasions or exacerbate the spread or impacts 
of established invaders, leading to non-linear “‘in- 
vasional meltdown’? (Simberloff and Von Holle 
1999). In our system, an introduced pollinator ap- 
pears to be facilitating the invasion of two intro- 
duced shrubs. : 

Recent concern over the environmental impacts 
of Apis introduction has focused on the fates of 
native pollinators (Roubik 1982; Buchmann and 
Nabhan 1996; Goodell 2000). We point to another 
potential conservation issue, the effect of Apis in- 
troduction on the spread of noxious weeds in nat- 
ural ecosystems. In keeping with this warning, re- 
cent work by Barthell et al. (2001) found that pol- 
lination by Apis contributed up to half of the seed 
produced by the noxious invader Centaurea solsti- 
tialis (yellow star thistle) in California. Beekeeping 
is common at the suburban-rural interface. The bees 
at our sites may have come from feral colonies, or 
beekeepers may have been responsible for domestic 
colonies in nearby residential or agricultural areas. 
In light of the results presented here, the potential 
negative impacts of beekeeping on weed manage- 
ment should play a role in policy decisions having 
to do with local introduction and control of Apis 
colonies. 


ACKNOWLEDGMENTS 


We would like to thank Carla D’ Antonio and Greg Gil- 
bert for helpful discussions, and the California State Park 
System for permission to work on their protected lands. 
K. M. Karoly provided access to unpublished data. Part 
of this work was supported by NSF grant DEB-9808501 
to IMP. 


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MApRONO, Vol. 49, No. 1, pp. 33-45, 2002 


FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY IN THE 
KLAMATH MOUNTAINS AND SOUTHERN 
SIERRA NEVADA, CALIFORNIA 


ANDREW J. ECKERT! AND JOHN O. SAWYER 
Department of Biological Sciences, Humboldt State University, Arcata, CA 95521 


ABSTRACT 


We sampled 15 foxtail pine stands located in the central ridges of the Klamath Mountains in order to 
estimate conifer density, basal area, and importance values. We compared these estimates to previous 
research in the southern Sierra Nevada. Our analyses revealed a lack of interregional divergence of stand 
characteristics between the Klamath Mountains and southern Sierra Nevada despite subspecific designa- 
tion and recently identified genetic divergence. Bray-Curtis ordination and hierarchical cluster analyses 
identified four stand types—1) stands dominated by foxtail pine, 2) stands with foxtail pine and whitebark 
pine, 3) stands with foxtail pine and red fir, and 4) mixed stands with foxtail pine, red fir, and western 
white pine. Sub-regions within the Klamath Mountains differed in foxtail pine relative density, conifer 
diversity, and substrate heterogeneity. Further analyses of the Klamath Mountains stands identified an 
inverse relationship between foxtail pine importance and conifer diversity. Interactions between the moun- 
tain island effect and substrate heterogeneity were inferred as regulatory mechanisms for foxtail pine 
importance and conifer diversity, but further research is needed to determine causal relationships from 
our correlations. 


Key words: foxtail pine, conifer diversity, mountain island effect, substrate heterogeneity, Klamath Moun- 


tains 


Foxtail pine (Pinus balfouriana Grev. & Balf.) is 
a California endemic conifer found in two isolated 
areas of the state separated by 500 kilometers. In 
extreme northern California it grows in the central 
ranges of the Klamath Mountains, while the south- 
erm population is centered on the Cottonwood Basin 
and surrounding areas in the southern Sierra Ne- 
vada. The Scottish botanist and explorer John Jef- 
fery discovered it in 1852 on a botanical recon- 
naissance of the Klamath Mountains. Professor 
John H. Balfour and Dr. R. K. Greville subsequent- 
ly prepared the original description and illustrations 
for the Oregon Association (Mastrogiuseppe 1972). 
Since its discovery, however, there has been a pau- 
city of ecological research into the biology of this 
subalpine tree. Ball (1976) highlights the need for 
research into the factors determining the ecological 
patterns of foxtail pine. 

Foxtail pine is a five-needle haploxylon pine 
placed within the subsection Balfourianae Engelm. 
along with Great Basin bristlecone pine (Pinus lon- 
gavea D. Bailey) in the western Great Basin and 
the Rocky Mountain bristlecone pine (Pinus aris- 
tata Engelm.) in the eastern Great Basin and Rocky 
Mountains. Mastrogiuseppe (1972) argued that two 
allopatric subspecies of foxtail pine exist upon dif- 
ferences in mean cone length, seed wing length, 
cotyledon number, and needle resin duct spacing. 
Mastrogiuseppe and Mastrogiuseppe (1980) subse- 


' Present address: Department of Botany, Box 355325, 
University of Washington, Seattle, WA 98195. 


quently named the southern population Pinus bal- 
fouriana ssp. austrina. 

An 11 loci allozyme analysis of stands within the 
northern population revealed high levels of genetic 
differentiation among stands (Oline et al. 2000). 
This genetic diversity was significantly greater than 
the differentiation between the northern and south- 
ern populations and was hypothesized to be a func- 
tion of the mountain island effect, genetic drift, 
possible serpentine soil adaptation, and high conifer 
richness (Hamrick et al. 1994; Oline et al. 2000). 

Foxtail pine and Great Basin bristlecone pine 
from the White Mountains exhibit strong anatomi- 
cal and morphological similarities suggesting hy- 
bridization (Mirov 1967; Bailey 1970). Successful 
experimental crosses support this proposed hybrid- 
ization based on morphological continuity (Critch- 
field 1977). Numerous hypotheses concerning the 
origin of the subsection Balfourianae, foxtail pine, 
and its subspecies have been offered (Mastrogiu- 
seppe 1972; Critchfield 1977; Raven and Axelrod 
1978). Most hypotheses, however, date the initial 
divergence of the bristlecone/foxtail pine ancestor 
to the Oligocene and the disjunction of foxtail pine 
within California to the Xerothermic period of the 
Holocene approximately 8000 years ago (Critch- 
field 1977; Raven and Axelrod 1978) or the Sierran 
orogeny approximately 2 million years ago (Mirov 
1967; Bailey 1970). 

Population and community-level research on 
foxtail pine is sparse. This pine is intolerant of 
shade at all stages of growth, an inhabitant of var- 
ious high elevation substrates, and is a major to 
minor component of subalpine forests and wood- 


34 MADRONO 


lands in both the Klamath Mountains and southern 
Sierra Nevada (Ryerson 1983; Sawyer and Thorn- 
burgh 1988). Extreme old age is achieved on xeric 
high altitude locations and trees can reach a maxi- 
mum age of approximately 2000 years in the south- 
ern population (Ball 1976; Scuderi 1987). Trees 
from the northern population, however, only attain 
maximum ages of approximately 800 to 1,000 years 
as a result of widespread heart rot and periodic fire 
(Mastrogiuseppe 1972). 

Foxtail pine typically occurs in low diversity 
stands with whitebark pine (Pinus albicaulis En- 
gelm.), lodgepole pine (Pinus contorta Loudon ssp. 
murrayana Critchf.), western white pine (Pinus 
monticola Douglas), red fir (Abies magnifica Andr. 
Murray), and mountain hemlock (Tsuga mertensi- 
ana Carriere) (Mastrogiuseppe 1972; Ryerson 
1983). Common ground layer associates include 
dry site graminoids, herbs, and shrubs such as 
Agrostis, Arabis, Arcotstaphylos, Carex, Ceano- 
thus, Eriogonum, Festuca, Juncus, and Sedum 
(Ryerson 1983; Sawyer and Keeler-Wolf 1995). 

Foxtail pine in the Klamath Mountains forms ex- 
tensive stands at high elevations on isolated peaks 
and ridges with mafic and ultramafic substrates 
where it is the dominant tree. Exceptions occur in 
the China Peak and Russian Peak areas where it is 
associated with as many as six other conifer species 
in a mixed subalpine forest type (Sawyer and Kee- 
ler-Wolf 1995). Conifers such as Jeffrey pine (Pi- 
nus jeffreyi Grev. & Balf.), incense-cedar (Caloced- 
rus decurrens (Torrey) Florin), and Douglas-fir 
(Pseudotsuga menziesii (Mirbel) Franco) form a 
minor component in these stands. In the southern 
Sierra Nevada, foxtail pine forms extensive low di- 
versity stands at high elevations on granitic and 
metamorphic substrates. Common associates in- 
clude whitebark pine, limber pine (Pinus flexilis 
James), red fir, and western white pine as described 
by Vankat (1970) in Sequoia National Park as a 
foxtail pine forest type. 

Limited research suggests that foxtail pine’s dis- 
tribution is determined by variation in substrate, cli- 
mate, and interspecific interactions with other co- 
nifers (Mastrogiuseppe 1972; Ryerson 1983; Scu- 
deri 1987; Oline et al. 2000). The low competitive 
ability of foxtail pine and the high regional conifer 
species richness of the Klamath Mountains may il- 
lustrate restriction of foxtail pine to safe sites on 
ultramafic substrates and granitic boulder fields 
where shade tolerant conifers cannot shade out in- 
tolerant foxtail pine trees (Ryerson 1983; Sawyer 
and Thornburgh 1988). 

Previous research has identified examples of eco- 
logical differences between the two populations. 
Foxtail pine stands in the southern Sierra Nevada 
achieve their highest densities on northern slopes 
(Ryerson 1983), which contradicts the typical 
southern and eastern slope success observed in the 
Klamath Mountains (Mastrogiuseppe 1972). Sur- 
prisingly the range of foxtail pine, commonly de- 


[Vol. 49 


scribed as a Pleistocene relict, has been document- 
ed as expanding to lower elevations in the southern 
Sierra Nevada (Ryerson 1983). This expansion may 
be a function of climate change, low conifer rich- 
ness, demographic and environmental stochasticity, 
or a combination of these processes (Mastrogiusep- 
pe 1972; Ryerson 1983; Scuderi 1987). 

We documented environmental and composition- 
al gradients related to the level of dominance of 
foxtail pine among stands in the Klamath Moun- 
tains. We also compared these stands to those of 
the southern Sierra Nevada using the studies of 
Vankat (1970) and Ryerson (1983). These compar- 
isons allowed us to analyze the disjunct foxtail pine 
populations at several levels and to make inferences 
about the role of interspecific interactions, geologic 
substrate, and climate in the distribution and dom- 
inance of foxtail pine at the intraregional and local 
levels in the Klamath Mountains. 


METHODS 


Study areas. The Klamath Mountains are com- 
posed of a complex set of predominantly southwest 
to northeast trending mountain ridges separated by 
deep canyons and valleys that encompass approxi- 
mately 30,300 km? of northern California (Fig. 1). 
The main mountain ranges, from south to north, are 
the South Yolla Bolly Mountains, North Yolla Bol- 
ly Mountains, Trinity Alps, Salmon Mountains, 
Scott Mountains, Trinity Mountains, Marble Moun- 
tains, and Siskiyou Mountains. Summits average 
from 1500 to 2100 m in elevation, with a maximum 
of 2750 m at Mount Eddy. The major watersheds 
include the Sacramento, Trinity, Salmon, Scott, 
Klamath, and Smith Rivers. Climate patterns con- 
form to a modified mediterranean climate type with 
long, wet winters and generally dry summers. Ay- 
erage precipitation ranges from 125 cm to 175 cm, 
with thunderstorms and lightning developing in late 
August and early September (Major 1988). Plant 
assemblages range from low elevation chaparral, 
woodlands and forests, to patches of alpine plants, 
with a number of endemic and relict populations of 
vascular plants on ultramafic substrates (Walker 
1954; Kruckeberg 1992). 

The southern Sierra Nevada is a linear north- 
west-southeast trending mountain range bounded 
on the west by the San Joaquin Valley and on the 
east by the White Mountains and the Great Basin 
(Fig. 1). Over 285 peaks reach elevations of 3600 
m, over 140 exceed 3900 m, while 11 peaks top 
4200 m (Kruse 1990). Major watersheds include 
the many forks of the Kern, Kings and San Joa- 
quin Rivers. Regional climates are influenced by 
local topography, but all resemble a modified 
Mediterranean climate with moderately wet win- 
ters and dry summers broken by periods of after- 
noon thunderstorms. Annual precipitation ranges 
from 35 cm to 115 cm on the western crest and 
from 50 cm to 75 cm on the eastern crest (Major 


) 


. 
| 
. 
| 


| 
| 
| 
| 


2002] 


32 


pha GMM Pinus balfouriana 


ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 35) 


Ee eines i x $32 
: Qo 
x isolated occurrence | o 
“_ 2 
Limits of Pras aristetet aS 
ow 
-a | 
= 
9 Igo 200 300 MILES oe 
- - se a 
. vad OO 260 $00 400 KILOMETERS | | We 
jams t = 7 720 ne 1S ; 


Fic. 1. 


Distribution of foxtail pine in the Klamath Mountains and Sierra Nevada by Griffin and Critchfield 


(1972). MM = Marble Mountains, RP = Russian Peak, TA = Trinity Alps, TM = Trinity Mountains, YB = Yolla 


Bolly Mountains. 


1988). Plant assemblages range from low eleva- 


_ tion pinyon pine woodlands to alpine tundra on 


the eastside and from foothill woodlands, mon- 
tane and subalpine forests, to alpine vegetation 


_ types on the westside. 


Sampling techniques. We identified five sub-re- 
gions where foxtail pine was dominant within the 
Klamath Mountains—the Yolla Bolly Mountains, 
the Trinity Alps, the Trinity Mountains, Russian 
Peak, and the Marble Mountains (Table 1). Each 


26 MADRONO 


TABLE 1. SUB-REGIONAL LOCALITIES OF 15 FOXTAIL PINE 
STANDS IN THE KLAMATH MOUNTAINS. SUB-REGIONS AND 
LOCALITIES WERE BROADLY DEFINED AND DO NOT HAVE DISs- 
TINCT BOUNDARIES. 


Sub-region Locality Stands 
Marble Mountains Lake Mountain 2 
Russian Peak Russian Peak 1 
Trinity Alps Granite Peak 2 

Seven-Up Peak 1 
Union Creek D 
Trinity Mountains Mt. Eddy 2 
China Mountain/ 
Crater Lake 2 
Yolla Bolly Mountains North 
South—Mt. Lynn 1 


sub-region contained distinct localities where fox- 
tail pine density was sufficiently high for sampling. 
At each locality we randomly identified stands pri- 
marily through homogeneity of species composi- 
tion and secondarily through homogeneity of geo- 
morphology. Stand areas ranged from 8 ha to 28 
ha. A total of 15 stands were sampled from the 10 
localities in the 5 sub-regions from May 22 through 
September 16, 2000. 

We used the point centered quarter (PCQ) meth- 
od to sample each of the 15 stands in order to es- 
timate tree density, basal area, frequency, and im- 
portance values by species. This method is a plot- 
less sampling technique based upon the placement 
of random points along pre-determined transects 
perpendicular to contour lines within each stand. 
Trees were defined as single conifer individuals 
with diameter measured at breast height for trees 
taller than 1.37 m, at ground level for trees below 
1.37 m in height, and as an average for trees with 
multiple trunks. 

Ryerson (1983) sampled 15 marginal foxtail pine 
stands located in the southern Sierra Nevada using 
the PCQ method in a similar manner to estimate 
conifer importance values. Vankat (1970) conduct- 
ed linear transect sampling of foxtail pine forest 
type within Sequoia National Park to estimate stand 
and species-specific tree density, basal area, and 
frequency. 

To determine the number of points needed to ad- 
equately estimate stand density, we conducted pre- 
sampling of two stands in the North Yolla Bolly 
Mountains following Bonham’s (1989) method. We 
determined that 22 points were adequate to estimate 
stand-level parameters with an error of 10%. We 
increased this number to 25 to ensure reliable es- 
timation, but were only able to establish 15 points 
in the Marble Mountains stands due to their small 
Size. 

To generate estimates of tree density, basal area, 
and frequency, we placed 25 points along four to 
six transects that were evenly spaced throughout 
the stand. Spacing among points was dependent 
upon stand area. We followed Bonham’s (1989) 


[Vol. 49 


protocol at each point, yielding 100 sampled trees 
per stand. The diameter at breast height (DBH) was 
measured for each tree, and the average diameter 
and stem number were obtained for trees with mul- 
tiple trunks. We measured percent slope, aspect, el- 
evation, and identified bedrock type. Cover of boul- 
ders (>50 cm), cobbles (10 cm—S50 cm), gravels 
(<10 cm), and organic material was estimated us- 
ing the Braun-Blanquet scale (Bonham 1989). 


Estimators and statistical analyses. The PCQ 
method allowed unbiased estimation of tree density 
and basal area within each stand (Pollard 1971). 
From these values we calculated stand and species- 
specific tree density, basal area, frequency, and im- 
portance values (Kent and Coker 1992). Importance 
values are composite estimates based upon the 
summation of relative estimates of species density, 
frequency, and basal area (Bonham 1989). These 
values could therefore range from a minimum of 
zero, if the tree species was absent, to a maximum 
of 300 if it was the only species present. 

We placed foxtail pine trees into ten size classes 
based upon diameter measurements. Using a simple 
linear regression model constructed from tree ring 
counts obtained from Mastrogiuseppe (1972), we 
related logarithmically transformed values of tree 
diameter to age by the following equation: 


Log(age) = 1.1571(Log[DBH]) + 0.3762 (1) 


This model was significant (ANOVA: F,4,; = 
277.0459, P = 0.000001) and had a high degree of 
explanatory power (R* = 0.8738). Hundred-year 
age classes could be correlated to size classes with 
confidence and reproductive success inferred from 
relative density of the first age class (Ryerson 
1983). Survival among age classes was calculated 
as the ratio of the density of age class x to the 
density of age class x + 1. 

We used the Shannon-Wiener Index (10g,)) to 
measure substrate heterogeneity and tree diversity 
using median species-specific relative density val- 
ues for conifer diversity. We computed Jaccard sim- 
ilarity coefficients, weighted by species-specific im- 
portance values, among all stands to obtain average 
within and among sub-regional stand similarities 
within the Klamath Mountains. We subjected the 
data to a variety of statistical analyses. Significance 
levels for hypothesis tests were set at a = 0.05 or 
0.10. A modified t-test checked for significant dif- 
ferences among index values (Zar 1999). Two-sam- 
pled t-tests and Mann-Whitney U tests established 
interregional differences and similarities for stand 
characteristics using Ryerson’s (1983) and Vankat’s 
(1970) data sets. Variation around the mean of these 
estimates, if the data were normally distributed, was 
compared between regional populations using Har- 
tley’s equal variance test. We used simple linear 
regression analyses and General Linear Models 
(GLM) ANOVA’s to identify trends within and dif- 
ferences among stand characteristics and environ- 
mental variables by sub-regions within the Klamath 


2002] 


ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 37 


TABLE 2. SUMMARY OF THE ENVIRONMENTAL VARIABLES AND STAND CHARACTERISTICS OF 15 FOXTAIL PINE STANDS FROM 
THE KLAMATH MOUNTAINS BY SUB-REGIONS. Conifer diversity and substrate heterogeneity are reported as Shannon- 
Wiener Index (log,, base) values. Density given in trees/hectare and basal area in m*/hectare. 


Basal Area Elevation Slope Substrate Conifer Conifer 
Stand Density area (ha) (m) Aspect (%) heterogeneity diversity richness 
Yolla Bolly Mountains 
1 238 45 8.6 2365 WSW 28.0 0.09 0.39 3) 
2, Sil 10 25.4 2260 WSW BOF] 0.38 0.44 5 
3 85 14 L523 2335 NNW 42.6 0.18 Oi 3 
Trinity Alps 
4 nO DES 10.1 2240 WSW 29.0 0.29 0.42 5 
5) Di. Oa 11.4 2345 WNW 39:6 0.29 0.44 5 
6 133 36 28.0 DUES WNW 58).3) 0.29 0.26 3 
gl eZ Dey 11.2 2202 WNW 40.1 0.25 0.42 3 
8 190 38 28.0 2004 NNE 153 0.41 0.52 6 
Trinity Mountains 
9 252 44 47.5 2358 WNW 25.8 O33 OZ 6 
10 207 8 24.5 2376 SSE 32) 0.41 0.61 6 
iN 92 26 28.1 2500 WNW S57 0.41 0.42 5 
12 381 I 14.0 2205 ESE Te 0.41 0.69 6 
Russian Peak 
13 315) 24 12.8 2180 NNW S85 0.44 0.84 8 
Marble Mountains 
14 all 25 8.2 2003 SSE BES) 0.41 0.66 6 
15 198 10 Sal 2033 NNE 35.0 0.20 0.50 6 


Mountains. Average within and among sub-region- 
al Jaccard similarity coefficients were compared us- 
ing GLM ANOVA with Fischer’s LSD grouping 
algorithm among sub-regions with more than two 
stands. 

We used hierarchical cluster and Bray-Curtis or- 
dination analyses to reduce dimensionality within 
the data sets and to analyze multivariate correla- 
tions among stand compositional and environmen- 
tal variables (Kent and Coker 1992). Hierarchical 
cluster analyses provided dendrograms based on 
Euclidean distance, scaled by standard deviations, 
using species importance values. Clusters were 
identified with the group average (unweighted pair- 
groups) algorithm. We employed Bray-Curtis ordi- 
nation analyses to evaluate the results of the hier- 
archical cluster analyses and to correlate stand 
groupings with environmental variables using the 
Sorenson similarity coefficient based upon species 
importance values. Biplots were overlain with com- 
positional and environmental variable vectors and 
correlations with the first two axes were computed 
(McCune 1993). 


RESULTS 


Summary. Approximately 1,420 trees were sam- 
pled in the determination of importance values for 
11 conifer species sampled within 15 stands located 
in the Klamath Mountains. Stand densities ranged 
from 50 trees/ha in the North Yolla Bolly Moun- 
tains to 381 trees/ha at China Mountain (Table 2). 
Stand basal area ranged from a minimum of 8 m?/ 
ha at China Mountain to a maximum of 45 m?/ha 
at Mt. Lynn (Table 2). Foxtail pine was the most 


important conifer in 14 out of the 15 stands with a 
regional average importance value of 168, a mini- 
mum of 46 at Crater Lake, and a maximum of 241 
at Seven-Up Peak (Table 3). The first age class 
(<100 years) dominated (=50% relative density) 
the age class distribution of foxtail pine in 9 out 15 
stands. Maximum tree age approached 1000 years 
for a few foxtail pine trees scattered throughout 
stands. Average stand abundances by age class 
within the Klamath Mountains assumed J-shaped 
distributions and differed (ANOVA: F,14;; = 
120.06, P = 0.000001). This pattern was also ob- 
served in the southern Sierra Nevada (Ryerson 
1983). 


Klamath Mountains. The common tree species 
were foxtail pine, red fir, and western white pine 
with importance value dependent upon geographic 
sub-region. Foxtail pine achieved higher densities 
on steep, high elevation slopes with western aspects 
(Tables 2, 3). Whitebark pine was primarily found 
in stands on high elevation peaks with granitic or 
glacial till substrates. Mountain hemlock tended to 
be most important on northern slopes at lower el- 
evations and east slopes at higher elevations. Co- 
nifers typical of lower elevations, such as incense- 
cedar, Jeffrey pine, and Douglas-fir, were found in 
stands with elevations below 2200 m in elevation 
or western and southern aspects. 

Conifer diversity was variable throughout the re- 
gion, but was greatest in northern-most stands (Ta- 
ble 2). Stands in the Yolla Bolly Mountains and 
southern Trinity Alps had lower diversity values, 
while stands in the Trinity and Marble Mountains 
had higher diversity values. Diversity values ranged 


38 MADRONO 


[Vol. 49 


TABLE 3. CHARACTERISTICS OF FOXTAIL PINE WITHIN 15 STANDS FROM THE KLAMATH MOUNTAINS BY SUB-REGIONS. 
Density given in trees/hectare, basal area in m7/hectare, and relative values in percent. 


Relative 
Stand Density density 
Yolla Bolly Mountains 
1 ISS 65 
D 52 63 
3 62 73 
Trinity Alps 
4 98 Syil 
5 2 47 
6 109 82 
d 53 47 
8 116 61 
Trinity Mountains 
9 153 61 
10 79 38 
11 13) 59 
12 69 18 
Russian Peak 
13 69 2D 
Marble Mountains 
14 74 43 
15 109 5) 


from 0.25 at Seven-Up Peak to 0.83 at Russian 
Peak. These two values were statistically different 
(t; = 43.9796, P < 0.00001) according the t-test 
proposed by Hutcheson (1970). Conifer richness 
was as low as 3 species in the Trinity Alps stands 
and as high as 8 species in the Russian Peak stand. 

Foxtail pine inhabited slopes with a variety of 
aspects. A large proportion of stands had northwest 
and southwest aspects with foxtail pine importance 
higher on southwest facing slopes (Tables 2, 3). 
Substrate heterogeneity measured with the Shan- 
non-Wiener Diversity Index ranged from a low of 
0.09 in the North Yolla Bolly Mountains to 0.44 at 
Russian Peak (Table 2). Inceptisols developed from 
mafic and ultramafic geologic substrates were the 
primary soil type within stands. Pockets of granitic 
and metasedimentary rocks were prominent in the 
Yolla Bolly Mountains stands and at Russian Peak. 
Evidence of glaciation was present in high eleva- 
tion stands with glacial till as an important substrate 
at stands in the Trinity Alps, Russian Peak, and 
Trinity Mountains. Elevations ranged from a low of 
2000 m at Lake Mountain to a high of 2500 m at 
China Peak with a regional average of 2247 m (Ta- 
blew): 

Simple linear regression analyses identified nu- 
merous compositional and environmental gradients 
among stands located in the Klamath Mountains 
(Fig. 2). A south to north latitudinal gradient was 
identified as significantly influencing foxtail pine 
relative density, conifer diversity, species richness, 
substrate heterogeneity, and foxtail pine survival to 
the second age class (F,,4 > 4.35, P < 0.05, R* > 
0.30). Conifer diversity negatively influenced fox- 
tail pine importance (Fig. 2A). Substrate heteroge- 
neity, in turn, was identified as positively influenc- 


Basal Relative Relative Importance 
area frequency dominance value 
38 46 95 207 

7 48 89 199 
10 56 90 220 
ts) 48 Syl 156 
13 44 51 142 
31 61 98 241 
20 46 80 7/3) 
33 44 81 187 
38 45 96 202 

5) 55 68 139 
21 46 88 195 

3) 18 10 46 

9 26 38 85 
7 31 89 163 

4 40 64 IS) 


ing conifer diversity (Fig. 2B). This relationship 
was dependent upon Pleistocene glaciation with 
stands on glaciated surfaces exhibiting a stronger 
correlation with high conifer diversity (glaciated R? 
= 0.6181, unglaciated R? = 0.3002). 

GLM ANOVA’s of compositional and environ- 
mental variables revealed sub-regional differentia- 
tion. Elevation, conifer species richness and diver- 
sity, whitebark pine relative density, and foxtail 
pine survival to the second age class significantly 
differed among the five sub-regions (ANOVA: F,,, 
> 3.25, P < 0.05). In general, foxtail pine survival 
was highest in stands with lower conifer diversity. 

Stands were more homogeneous, as measured 
with average sub-regional Jaccard similarity coef- 
ficients, within sub-regions versus among sub-re- 
gions (Fig. 3). In general, within sub-region Jaccard 
similarity coefficients were greater than among sub- 
region coefficients with the Marble Mountains be- 
ing similar to all other sub-regions (ANOVA: F3 157 
= 17.50, P = 0.0000009). Russian Peak was most 
similar to the Trinity Mountains and Marble Moun- 
tains, while least similar to the Trinity Alps and 
Yolla Bolly Mountains. Similarity was not corre- 
lated to distances among sub-regions, however the 
extreme northern and southern stands were among 
the least similar. The recently disturbed stand (e.g., 
Crater Lake, stand 12) was the least similar to oth- 
ers at both within and among sub-regional levels. 

The Bray-Curtis ordination of the Klamath 
stands differentiated stands with high conifer rich- 
ness from those with high foxtail pine importance 
values (Fig. 4). Axis I in the ordination was highly 
and variously correlated with mountain hemlock (R 
= —(0.837), western white pine (R = 0.564) and 
foxtail pine (R = 0.547) importance values. Axis II 


mG 2. 


_ 0.6399). Stands with serpentine soils are represented by filled points. (B) Relationship between substrate heterogeneity 
_ and conifer diversity grouped by glaciation history (dashed line = 


2002] 


(A) 
250 5 


225 - 


200 - “< oO e 


Foxtail pine importance 
N 
(Sy) 


205 


(0) - 7s T T 


ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 


38) 


T U T T 


0.35 0.40 


T T 


0.20 0.25 0.3 


I T 


0.45 0.50 055 060 065 070 0.75 0.80 0.85 0.90 


Conifer Diversity 


‘= 
fo) 
1 


Conifer Diversity 
—) 
(3)] 
| 


0.4 5 


oO Glaciated 


m Unglaciated 


- R*=06181 


T 


0.20 0.25 0.30 0.35 0.50 


Substrate Heterogeneity 


0.15 0.40 0.45 


(A) Relationship between conifer diversity and foxtail pine importance (F, ,, = 23.096, P = 0.000343, R? = 


unglaciated regression, solid line = glaciated 


regression). See text for definitions of conifer diversity and substrate heterogeneity. 


was similarly correlated with western pine (R = 
—0.617), whitebark pine (R = —0.616), and foxtail 
pine (R = 0.603) importance values. These two 
axes had relatively high explanatory power (R? = 
0.739). Stands tended to be broadly grouped by 
sub-region and composition (Fig. 4). Crater Lake 
was unique with extremely low foxtail pine impor- 


_ tance and high conifer species richness. This stand 


was selectively logged and burned in the last half 
century. 


Southern Sierra Nevada. Using Ryerson’s (1983) 
data set, stand density ranged from a low of 72 


trees/ha to a high of 881 trees/ha, while stand basal 
area ranged from a low of 9 m*/ha to 646 m*/ha 
(Table 4). Extremely high density and basal area 
values were associated with whitebark pine domi- 
nance and may have been an artifact of sampling 
trees with numerous trunks. A wide range of im- 
portance values for foxtail pine, low species rich- 
ness, and high elevations characterized these stands 
(Table 4). 

Using Vankat’s (1970) data set, stand density 
ranged from a low of 200 trees/ha to a high of 700 
trees/ha, while stand basal area ranged from 15 m7/ha 


40 | MADRONO [Vol. 49 


97.5 
95.0 
92.5 
90.0 - 
87.5 


85.0 


Average Jaccard Similarity (%) 


82.5 - 


80.0 
77.5 - 
75.0 
Within Among Within Among Within Among Within Among 
Yolla Bolly Trinity Alps Trinity Mtns. Marble Mtns. 


Fic. 3. Comparison of average within and among sub-regional Jaccard similarity coefficients for 15 foxtail pine stands 
in the Klamath Mountains. Differences are significant (ANOVA: F; 5, = 17.50, P = 0.0000009). 


N 
NM Northt 
or 
< 
North2 
North3 
North6 


North? 


North15 
Oe O Axis 1 


V/ North3 


North11 


North13 


North12 Ey 
o 


North10 


Fic. 4. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 15 foxtail pine stands 
from the Klamath Mountains. Symbols are proportional to foxtail pine importance. Marble Mountains (closed dia- 


monds), Russian Peak (open triangles), Trinity Alps (open circles), Trinity Mountains (open squares), and Yolla Bolly 
Mountains (closed triangles). 


2002] 


ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 41 


TABLE 4. CHARACTERISTICS OF 15 FOXTAIL PINE STANDS IN THE SOUTHERN SIERRA NEVADA. Density given in trees/ 
hectare, basal area in m?/hectare, and relative values in percent. Data are from Vankat (1970) and Ryerson (1983). 


Averages +1 SE in parentheses. 


Vankat 
Stand Density Basal area 
1 500 220 
2 0 0 
3 500 161 
4 300 182 
5) 600 67 
6 400 66 
1 200 97 
8 600 n/a 
2 200 151 
10 — —: 
11 — — 
1192 — — 
13 — — 
14 — — 
15 — — 
Average 367 (69) 118 (24) 


to 56 m*/ha (Table 4). Stand elevations ranged from 
2900 m to 3650 m. Importance values for foxtail 
pine were at or approached 300 with western white 
pine, whitebark pine, and lodgepole pine as minor 
stand components. This prompted Vankat to iden- 
tify a foxtail pine forest type. Tree ring counts in- 
dicated that foxtail pine cover and density began 
increasing approximately 110 years ago in accor- 
dance with decreased grazing levels by sheep. 


Interregional comparison. A comparison to 
Ryerson’s data set showed that stand basal area, 
foxtail pine basal area, elevation, and conifer rich- 
ness differed between regions, but importance val- 
ues did not (Table 5). A comparison to Vankat’s 
data set showed that stand density, foxtail pine den- 
sity, and foxtail pine relative basal area differed be- 
tween regions (Table 5). Variances around these es- 
timates also differed between regions, with the 
southern Sierra Nevada having significantly greater 
variance estimates (Table 5). Comparisons between 
Ryerson’s and Vankat’s data sets showed differenc- 
es in stand density, foxtail pine relative density, and 
foxtail pine relative basal area (Table 5). 

The Bray-Curtis ordination using all stands did 
not cluster stands exclusively by region (Fig. 5), but 
there was broad separation of the majority of Klam- 
ath Mountain stands from a bifurcated grouping of 
southern Sierra Nevada stands dependent upon 
dominance of either foxtail or whitebark pine (Fig. 
5). The stands with the highest importance values 
for foxtail pine were exclusively identified in the 
Sierra. Axis I in the ordination using all stands was 
highly correlated with foxtail pine importance (R = 
0.855) and axis II with whitebark pine importance 
(R = —0.833). These axes had relatively high ex- 
planatory power (R? = 0.757). Four major clusters 
were identified: 1) stands dominated by foxtail 


Ryerson 

Foxtail pine 

Density Basal area importance 
78 38 276 
193 9 296 
34 13 122 
178 36 120 
54 6 68 
a2 18 90 
74 N7/ 59 
74 35 VS 
DAS 61 DW 
DDD 75 285 
11 2 16 
61 26 273 
111 4 44 
29 4 25) 
51 15 181 

100 (20) 24 (6) 145 (26) 


pine, 2) stands with foxtail pine and whitebark pine, 
3) stands with foxtail pine and red fir, and 4) mixed 
stands with foxtail pine, red fir, and western white 
pine (Fig. 5). 

Stands with red fir and western white pine were 
associated with the lower elevations, while stands 
with whitebark pine were associated with higher 
elevations or northern stands. Incense-cedar, Doug- 
las-fir, and mountain hemlock were sampled only 
within the Klamath Mountains, while limber pine 
and mountain juniper (Juniperus occidentalis 
Hook.) were exclusively present in the Sierra Ne- 
vada. Red fir was present in the Sierra as the typical 
variety (Abies magnifica var. magnifica) and in the 
Klamath by Shasta red fir (Abies magnifica Andr. 
Murray var. shastensis Lemmon). An inverse rela- 
tionship between foxtail pine importance and co- 
nifer species richness was evident in the results of 
both procedures (Figs. 4, 6). 


DISCUSSION 


Recent genetic research has documented inter- 
regional and intraregional divergences of foxtail 
pine populations (Oline et al. 2000). We expected 
similar conclusions when compositionally compar- 
ing stands at these levels. Therefore we expected 
foxtail pine stands in close proximity within the 
Klamath Mountains to be similar, especially if they 
were located on the same substrate. Likewise, we 
expected the Klamath Mountains and southern Si- 
erra Nevada stands to differ in several ways. 

Both regions had simple stands dominated by 
foxtail pine, but this organization was found more 
often in the southern Sierra Nevada. Mixed stands 
with red fir or whitebark pine cut across regions. 
As found here, past studies suggested the lack of 
ecological differences between stands with different 


42 MADRONO 


[Vol. 49 


TABLE 5. STAND AND FOXTAIL PINE CHARACTERISTICS COMPARED USING TWO-SAMPLE T-TESTS (T), MANN-WHITNEY U 
TESTS (Z), AND HARTLEY'S EQUAL VARIANCE TESTS BETWEEN KLAMATH AND SIERRA REGIONS. Comparison to Ryerson 
(1983) is indicated by R and to Vankat (1970) by V. Equal variance tests were not conducted on non-normal data. 


Two-sample T and Mann-Whitney U tests 


Hartley’s test 


Variable Source Sample sizes Critical value P-value F-value P-value 
Conifer richness R n, = 15 T = 3.543** 0.00141** 1.600 0.389848 
Elevation (m) R a s fe Z = 4.668** 0.000003** — — 
Importance value R a ic ET = —0:814 0.424904 4.026** 0.013294 
Basal area R -, a a T = 1.847*** 0.08583*** 248.941 ** 0.000001 
Density R = a ie T = 0.248 0.806775 264.552** 0.000001 
Relative basal area R ze ie Z = —0.560 OD75511 — — 
Relative basal area* Vv : is Thy = PIS NEKO 0.003388** — — 
Relative density R ey z ie Z = —0.477 0.63325 a — 

Ny = 15 
Relative density* Vv n, = 15 Tie eo nly lee 0.001521** — — 
Relative frequency R a a ie 1S) 02135 0.27096 Daten | 0.002213 
Stand basal area R Fs is T= 22i0F= 0.036552** 264.552** 0.000001 
Stand density R rf, B ie T = 0.848 0.405853 339582 0.029873 
Stand density* Vv - = 13 Ti=s— 3.65338 0.004152** 4.039** 0.037774 
ny = 


* Tests for statistical differences between these values from Ryerson (1983) and Vankat (1970) were significant with 


a = 0.05. ** Significant at a = 0.05. *** Significant at a = 0.10. 


red fir varieties (Barbour and Woodward 1985). 
Mixed foxtail pine stands with western white pine 
grew on sites with similar conditions to those for 
unmixed stands in both regions suggesting similar 
habitat needs by both species within and between 
regions. This result was expected because of west- 
ern white pine’s prevalence in many subalpine for- 
est types within the Californian mountains (Sawyer 
and Keeler-Wolf 1995). As expected there were 
species exclusive to each region (Griffin and 
Critchfield 1972), but they were not important in 
differentiating stands at the regional level. 
Ryerson found that mixed stands with whitebark 
pine were associated with less-developed soils at 
higher elevations typical of xeric, nutrient-limited 
sites, while mixed stands containing red fir were 
associated with deeper and more developed soils 
typical of mesic, nutrient-rich sites. Our analyses 
identified a cluster of foxtail pine-red fir dominated 
stands that cut across both regions. Within the 
Klamath Mountains, red fir was a more common 
associate of species rich stands, but these were nei- 
ther more mesic nor nutrient-rich than stands dom- 
inated by foxtail pine. Stands dominated almost ex- 
clusively by foxtail pine in the Klamath Mountains 
were typically found on sites intermediate to the 


extremes mentioned by Ryerson, on ultramafic 
soils, or homogeneous substrate compositions. 

An alternate explanation to Ryerson’s site limi- 
tation hypothesis may better explain the foxtail pine 
stands mixed with red fir. These stands may be the 
product of species-specific elevation limits. Within 
the Sierra Nevada, conifer species zone over a con- 
siderable elevation range. Much of the upper mon- 
tane red fir forests are replaced with lodgepole pine 
and whitebark pine forests as elevation increases 
(Potter 1998). Individual red fir trees are found in 
these types illustrating broad ecotones between for- 
est types. This is not the case in the compressed 
forest zones found in the Klamath Mountains. 

The comparison of Ryerson’s and Vankat’s data 
sets illustrated important differences suggesting that 
different sampling methods accounted for the dif- 
fering estimates of basal area. Stand selection cri- 
teria may have been even more important. Ryerson 
picked stands at marginal locations, while Vankat 
chose stands in areas with extensive foxtail pine 
dominance. The marginal stands of Ryerson were 
expected to have high variances if foxtail pine 
stands exhibited a core-periphery spatial structure. 
Comparisons reveled that Ryerson’s estimates were 
more variable and could be attributed to sampling 
from different core and peripheral populations. 


2002] ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 43 


Fic. 5. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 30 foxtail pine stands in 
the Klamath Mountains and southern Sierra Nevada. Open circles represent the Klamath Mountains stands. Closed 
triangles represent the southern Sierra Nevada stands. 1) stands dominated by foxtail pine, 2) stands with foxtail pine 
and whitebark pine, 3) stands with foxtail pine and red fir, and 4) mixed stands with foxtail pine, red fir, and western 
white pine. ABMA = red fir density, BA = stand basal area, PIAL = whitebark pine density, PIBA = foxtail pine 
density. 


Fic. 6. A Bray-Curtis ordination using Sorenson similarity coefficients showing patterns for 30 foxtail pine stands in 
the Klamath Mountains and southern Sierra Nevada. Open circles represent the Klamath Mountains stands. Closed 
triangles represent the southern Sierra Nevada stands. Symbol sizes are proportional to the conifer species richness 
found within each stand. 


44 MADRONO 


The finding that called most for an interpretation 
was the inverse relationship between foxtail pine 
importance and conifer diversity prominent in the 
Klamath Mountains. Because conifer diversity was 
also correlated to substrate heterogeneity, simple 
causal explanations are difficult. Several authors 
have an explanation for this pattern that might be 
called the marginal hypothesis (Wright and Mooney 
1965; Ryerson 1983; Sawyer and Thornburgh 
1988; Kruckeberg 1992; Oline et al. 2000). They 
offer that within marginal habitats decreased biotic 
interactions lead to increased species diversity. 
Marginal habitats have been hypothesized as com- 
posed of nutrient limited soils (Sawyer and Thorn- 
burgh 1988; Kruckeberg 1992; Oline et al. 2000), 
extreme climate (Ryerson 1983), and cyclically dis- 
turbed habitat (Murray et al. 2000). Two possible 
marginal site types exist within the distributional 
limits of foxtail pine—high elevation sites and ultra- 
mafic soils. 

Ryerson proposed that the highest elevation 
stands within the Sierra Nevada simulated marginal 
habitat, but foxtail pine acts as a timberline species 
in the center of its range when south of whitebark 
pine’s range (Scuderi 1987). These stands are sim- 
ple with one or two species, rather than mixed as 
proposed by the marginal hypothesis. This may be 
a function of species distribution ranges along the 
Sierran crest. Similar elevations are absent in the 
Klamath Mountains, and elevation was not corre- 
lated to species richness. 

Ultramafic soils have been argued as marginal 
sites that isolate species and promote increased di- 
versity. This soil type is absent within the range of 
foxtail pine within the southern Sierra Nevada. 
Stands located on marginal ultramafic soils in this 
study were not more diverse than stands located at 
better sites (see Fig. 2). Nor were they less diverse 
with respect to conifer diversity. Stands with high 
conifer diversity were found on schists, gabbro, and 
peridotite in the Yolla Bolly Mountains, Trinity 
Alps, and Trinity Mountains, as were the stands 
dominated by foxtail pine. The most diverse stand, 
sampled at Russian Peak, was on glaciated granite. 
These results suggest that ultramafic soils do not 
directly control conifer species diversity as pro- 
posed in the marginal hypothesis. Nor do they con- 
trol foxtail pine importance (see Fig. 2). 

Conifer diversity in the Klamath Mountains was 
variable, and was greatest in the northern-most 
stands. The stands in the Yolla Bolly Mountains 
were well south of the range of lodgepole pine and 
whitebark pine and at the range limits for mountain 
hemlock and western white pine, reducing the spe- 
cies pool in the south. In the northern stands, these 
subalpine species mixed with montane conifers 
such as Douglas-fir, incense-cedar, and white fir. 
Within the Sierra Nevada, the limits of these mon- 
tane species are well below that of foxtail pine. 

Analysis of Jaccard similarity coefficients by 
sub-region reveled that foxtail pine stands in close 


[Vol. 49 


proximity were more similar to one another than to 
stands in distant sub-regions. This pattern indicated 
an island-like distribution of foxtail pine and other 
species as predicted by the mountain island effect 
hypothesis (Brown 1971; Hamrick et al. 1994). 
This hypothesis argues that mountain tops are iso- 
lated collections of species experiencing ecological 
processes commonly observed within insular sys- 
tems. These patterns may be the result of slow dis- 
persal rates inherent to conifers and/or historical 
climate fluctuations. Mohr’s et al. (2000) recreation 
of post-glacial vegetation history at two lakes in the 
Trinity Mountains supports this possibility. The 
vegetation patterns and histories differed as much 
as they were similar even though these lakes are 
approximately 4.5 km apart. 

Inferences with our data suggest that the moun- 
tain island effect enhanced by substrate heteroge- 
neity better explains conifer species diversity pat- 
terns than does the marginal hypothesis. Substrate 
heterogeneity was positively correlated with conifer 
diversity and glaciated substrates. Rocky conditions 
such as moraines would isolate individual trees in 
favorable microsites if the species was in the area 
to take advantage of these sites. The high diversity 
stand at Crater Lake was partially logged and it had 
high diversity. Qualitative observations suggested 
that boulders isolated the new seedlings and sap- 
lings establishing after the disturbance. 


SUMMARY 


Foxtail pine stands within the Klamath Moun- 
tains and southern Sierra Nevada did not differ dra- 
matically in structure. Foxtail pine importance was 
constant across both regions, but species richness, 
stand basal area, and elevation differed between re- 
gions. Within the Klamath Mountains, conifer di- 
versity, foxtail pine relative density, and substrate 
heterogeneity differed among five sub-regions. Co- 
nifer diversity and foxtail pine importance were not 
correlated to substrate type (e.g., ultramafic soils) 
as previously hypothesized. Significant correlations 
were observed among foxtail pine importance, co- 
nifer diversity, and substrate heterogeneity. These 
correlations were more significant on glaciated sub- 
strates versus unglaciated substrates. Inferences 
from our data suggest that foxtail pine importance 
and conifer diversity in the Klamath Mountains 
may be regulated by a mountain island effect en- 
hanced through substrate heterogeneity, but more 
research is needed to tease apart the causal mech- 
anisms within these correlations. 


ACKNOWLEDGMENTS 


We would like to thank the Department of Biological 
Sciences at Humboldt State University, the Hayfork Rang- 
er District of Shasta-Trinity National Forest, and Linda 
Peak for use of sampling equipment. My fiancé Melissa 
L. Postler and coworker Vin D’ Angelo provided invalu- 
able field assistance, and Bob’s Auto and Tire Repair Ser- 
vice located in Red Bluff, California fixed numerous flat 


a 


2002] 


tires obtained on the back roads of Siskiyou, Trinity, and 
Tehema counties. 


LITERATURE CITED 


BaiLey, D. K. 1970. Phytogeography and taxonomy of 
Pinus subsection Balfourianae. Annals of the Mis- 
souri Botanical Gardens 57:210—249. 

BALL, J. T. 1976. Ecological survey of Last Chance Mead- 
ow Candidate Research Natural Area. Unpublished 
report. USDA Forest Service, Pacific Southwest Re- 
search Station, Albany, CA. 

BarsBour, M. G. AND R. A. WOODWARD. 1985. The Shasta 
fir forest of California. Canadian Journal of Forestry 
15:570—576. 

BONHAM, C. D. 1989. Measurements for terrestrial vege- 
tation. John Wiley & Sons, New York. 

Brown, J. H. 1971. Mammals on mountaintops: Non- 
equilibrium insular biogeography. American Natural- 
ist 105:467—478. 

CRITCHFIELD, W. B. 1977. Hybridization of foxtail and 
bristlecone pines. Madrofio 24:193—211. 

GRIFFIN, J. R. AND W. B. CRITCHFIELD. 1972. The distri- 
bution of forest trees in California. Research Paper 
PSW-82. USDA Forest Service, Pacific Southwest 
Research Station, Albany, CA. 

Hamrick, J. L., A. E SCHNABEL, AND P. V. WELLS. 1994. 
Distribution of genetic diversity within and among 
populations of Great Basin conifers. Pp. 147—161 in 
K. T. Harper, L. L. St. Clair, K. H. Thorne, and W. 
M. Hess (eds.), Natural history of the Colorado Pla- 
teau and Great Basin. University of Colorado Press, 
Niwot, CO. 

HUTCHESON, K. 1970. A test for comparing diversities 
based on the Shannon formula. Journal of Theoretical 
Biology 29:151—154. 

KENT, M. AND P. COKER. 1992. Vegetation Description and 
Analysis: A Practical Approach. Belhaven Press, 
London. 

KRUCKEBERG, A. R. 1992. Plant life of western North 
American ultramafics. Pp. 31—74 in B. A. Roberts and 
J. Proctor (eds.), The ecology of areas with serpen- 
tinized rocks: A world view. Kluwer Academic Pub- 
lishers, Dordrecht, The Netherlands. 

KRusE, S. M. 1989. Climatic water budgets in the southern 
Sierra Nevada, California. Thesis, California State 
University, Fresno. 

Major, J. 1988. California climate in relation to vegeta- 
tion. Pp. 11—74 in M. G. Barbour and J. Major (eds.), 
Terrestrial vegetation of California (Expanded ver- 
sion), California Native Plant Society, Sacramento. 

MASTROGIUSEPPE, R. J. 1972. Geographic variation in fox- 
tail pine, Pinus balfouriana Grev. & Balf. Thesis, 
Humboldt State University. Arcata, CA. 

AND J. D. MASTROGIUSEPPE. 1980. A study of Pi- 


ECKERT AND SAWYER: FOXTAIL PINE IMPORTANCE AND CONIFER DIVERSITY 45 


nus balfouriana Grev.& Balf. (Pinaceae). Systematic 
Botany 5:86—104. 

McCune, B. 1993. PC —Ord. Version 3.11 computer pack- 
age. 

Mone, J. A., C. WHITLOCK, AND C. N. SKINNER. 2000. 
Postglacial vegetation and fire history, eastern Klam- 
ath Mountains, California, USA. The Holocene 10: 
587-601. 

Mirovy, N. T. 1967. The Genus Pinus. The Ronald Press 
Company, New York. 

Murray, M. P.,, S. C. BUNTING, AND P. MorGAn. 2000. 
Landscape trends (1753—1993) of whitebark pine (Pi- 
nus albicaulis) forests in the West Big Hole Range of 
Idaho/Montana, U.S.A. Arctic, Antarctic, and Alpine 
Research 32:412—418. 

OLINE, D. K., J. B. MITTON, AND M. C. GRANT. 2000. Pop- 
ulation and subspecific genetic differentiation in the 
foxtail pine (Pinus balfouriana). Evolution 54:1813-— 
1819. 

POLLARD, J. H. 1971. On distance estimators of density in 
randomly distributed forests. Biometrics 27:991— 
1002. 

Potter, D. A. 1998. Forested communities of the upper 
montane in the central and southern Sierra Nevada. 
General Technical Report PSW-GTR-169, USDA 
Forest Service, Pacific Southwest Research Station, 
Albany, CA. 

RAVEN, P. H. AND D. I. AXELROD. 1978. Origin and rela- 
tionships of the California flora. University of Cali- 
fornia Publications in Botany 72:1—134. 

RYERSON, D. 1983. Population structure of Pinus balfour- 
iana Grev. & Balf. along the margins of its distribu- 
tion area in the Sierran and Klamath Regions of Cal- 
ifornia. Thesis, Sacramento State University. Sacra- 
mento, CA. 

SAWYER, J. O. AND D. THORNBURGH. 1988. Montane and 
subalpine vegetation of the Klamath Mountains. Pp. 
699-732 in M. G. Barbour and J. Major (eds.), Ter- 
restrial vegetation of California, Expanded ed., Cali- 
fornia Native Plant Society, Sacramento. 

AND T. KEELER-WOoLF. 1995. A Manual of Cali- 
fornia Vegetation. California Native Plant Society, 
Sacramento. 

SCUDERI, L. 1987. Late-Holocene upper timberline varia- 
tion in the southern Sierra Nevada. Nature 325:242— 
244. 

VANKAT, J. L. 1970. Vegetation change in Sequoia Na- 
tional Park, California. Dissertation, University of 
California, Davis. 

WALKER, R. B. 1954. Factors affecting plant growth on 
serpentine soils. Ecology 35:259—266. 

WRIGHT, R. D. AND H. A. Mooney. 1965. Substrate-ori- 
ented distribution of bristlecone pine in the White 
Mountains of California. The American Midland Nat- 
uralist 73:257—284. 

ZAR, J. H. 1999. Biostatistical Analysis. Prentice Hall, Up- 
per Saddle River, NJ. 


MApRONO, Vol. 49, No. 1, pp. 46-47, 2002 


REVIEW 


The manzanitas of California, also of Mexico and 
the world, by Phillip V. Wells. 2000. Published by 
P. V. Wells, Department of Ecology and Evolution- 
ary Biology, Haworth Hall, University of Kansas, 
Lawrence KS 66045. Available from Cody’s 
Books, Berkeley, CA for $53.00. ISBN: 0-933994- 
DD. 


The recent review of Trees and Shrubs of Cali- 
fornia (Stuart and Sawyer 2001) by Rejmanek 
(2001) underscores the need for a review of The 
manzanitas of California (Wells 2000). Rejmanek 
points out that there are “‘more than 40 excluded 
species of Arctostaphylos” in Stuart and Sawyer 
(2001) and he suggests that people would be better 
off getting Wells (2000) if they have an interest in 
this quintessentially Californian group of woody 
shrubs. To a degree, we agree—but with some im- 
portant caveats that should be kept in mind. 

P. V. Wells has made the study of manzanitas a 
lifetime work and this self-published, un-peer re- 
viewed book demonstrates both the best and the 
worst of this kind of situation. Most of the infor- 
mation in the book, for example, can be found in 
previous publications of his, including most of the 
figures (See, e.g., Ecological Monographs 32:79-— 
103 [1962], Evolution 23:264—267 [1969], The 
Four Seasons 7:17—21 [1987], 8:46—70 [1990], 9: 
64-69 [1992]). Wells published his treatment on 
Arctostaphylos in the 1993 Jepson Manual (Hick- 
man 1993) (Chapter V). He also published a phy- 
logenetic hypothesis for Arctostaphylos in 1992 
(Chapter IV) (Wells 1992) in which he divides the 
genus into two subgenera and six sections. This lat- 
est book presents this treatment in detail and ig- 
nores later publications that cast serious doubt on 
Wells’ phylogenetic hypothesis (Markos et al. 
1998). He then goes on to trivialize important new 
findings in the genus by Keeley and his students 
(Keeley 1994; Keeley et al. 1997a, b). Figures in 
this book come from the original publications and 
many of the species names are woefully out of date, 
no longer valid even in the treatment by Wells. In 
Chapter V, he includes small, fuzzy, black and 
white photos of herbarium specimens to illustrate 
species; none of these are sharp enough or show 
the appropriate characters to be of much use. While 
he provides some interesting discussions, for ex- 
ample, concerning reticulate evolution, Wells has 
not published data that would enable objective 
evaluation of his conclusions, nor does it exist as a 
table or appendix in this book. The overall impres- 
sion is that Wells has decreed an ideal manzanita 
world that he can perceive and which now has been 
formally revealed. Indeed, but if only those plants 
in the field would behave! 


With these minor criticisms aside, we acknowl- 
edge that the majority of the taxa recognized by 
Wells are probably distinct lineages and this vol- 
ume provides a wealth of information about them. 
Indeed, we agree with Wells that manzanitas are the 
most diverse and fascinating genus of woody 
shrubs in the California Floristic Province (one of 
25 global “‘hotspots’’ on the planet [Mitermeier et 
al. 2000]). Most importantly, this book distills 
Wells’ long years of scholarly research into the ge- 
nus and provides an invaluable resource for any 
serious manzanita student. As mentioned above, 
Wells develops an intriguing chapter on reticulate 
evolution in Arctostaphylos (Chapter VII). This in- 
formative discussion concerning hybridization and 
speciation in manzanitas articulates his hypothesis 
concerning diploid hybridization. Wells presents 
scatter diagrams and multivariate-analytic figures 
that suggest evidence of hybrid origin between spe- 
cies such as A. canescens and A. andersonii (lead- 
ing to intermediate species such as A. glutinosa and 
A. auriculata). Unfortunately, the data supporting 
these studies is missing, and in tables speculating 
on species of possible hybrid origin, the inclusion 
of many plausible parents (e.g., A. glutinosa from 
A. canescens X A. andersonii) are undermined by 
a number of frankly preposterous suggestions (e.g., 
A. pilosula from A. wellsii X A. glauca). 

Wells also provides a lot of practical material. 
He gives interesting regional keys to manzanitas in 
Chapter VI, a chapter rich in historical lore. The 
Introduction contains a discussion of generic rela- 
tionships between Arctostaphylos and other mem- 
bers of the Arbutoideae (Arbuteae) distilled from 
Diggs and Breckon (1981) through Wells’ own per- 
spective. Hileman et al. (2001) provide a molecular 
counterpoint that reinforces the general accuracy of 
the generic circumscriptions that Wells describes 
for this monophyletic group. 

The Character Analysis (Chapter II) of Arcto- 
staphylos illustrates the fundamental flaw in Wells’ 
classification system. He lists 70 morphological 
traits that he claims he has analyzed for 61 species. 
He then describes character states for these traits 
and goes on to fashion elaborate descriptions of 
species that are infinitely complex based upon this 
enormous data set (again, which is not tabulated in 
the book). These species descriptions are enumer- 
ated in Chapter IV and ordered into the same phy- 
logenetic scheme that Wells (1992) proposed eight 
years ago. The flaw is that manzanitas don’t obey 
the world according to P V. Wells, as was illus- 
trated by Keeley (1994), Markos et al. (1998), Va- 
sey and Parker (1999), and our extensive field ob- 
servations. Wells continues with the belief that pan- 
icles can be described as racemes (with up to 7 


2002] 


branches!), and that nascent inflorescences with 
variable bract characters (leafy vs. scaly bracts) can 
be classified as either one state or the other exclu- 
sively. As a consequence, we have found that a 
great deal of confusion and mystification is created 
by a treatment that tends to elicit more frustration 
than enlightenment. 

And so, we caution, if you buy this book, focus 
on the real entities and not necessarily on the bi- 
or trinomial classification system offered by Wells. 
Luxuriate in the lore but take lightly the conclu- 
sions. Particularly, defer judgments regarding phy- 
logenetic relationships until an alternative and 
probably molecular phylogeny is worked out that 
meshes with morphology, cytology, and ecological 
information and that will also employ a more con- 
temporary species concept. There is a rich literature 
on manzanitas by some great California botanists 
and evolutionary biologists (e.g., Eastwood [1934], 
Jepson [1922], Dobzhansky [1953], Stebbins and 
Major [1965], etc.). These scientists recognized the 
extraordinary importance of Arctostaphylos to un- 
derstanding driving forces in the evolution of Cal- 
ifornia’s remarkable flora. Wells’ book culminates 
a wild and woolly era in the history of manzanita 
taxonomy that diverged from this venerable tradi- 
tion. Will Wells (2000) be the last word on man- 
zanitas? We don’t think so (Keeley 1998; Keeley 
et al. 1994, 1997a, b; Markos et al. 1998; Vasey 
and Parker 1999; Hileman et al. 2001). 


—MICHAEL C. VASEY AND V. THOMAS PARKER, Depart- 
ment of Biology, San Francisco State University, 1600 
Holloway Avenue, San Francisco, CA 94132. mvasey @ 
sfsu.edu; parker@sfsu.edu 


LITERATURE CITED 


Diccs, G. M. AND G. J. BRECKON. 1981. Generic circum- 
scription in the Arbuteae (Ericaceae). In G. M. Diggs, 
Systematic studies in the Arbuteae. Ph.D. dissertation. 
University of Wisconsin, Madison, WI. 

DOBZHANSKY, T. 1953. Natural hybrids of two species of 


REVIEW 47 


Arctostaphylos in the Yosemite region of California. 
Heredity 7:73—79. 

Eastwoop, A. 1934. A revision of Arctostaphylos with 
key and descriptions. Leaflets of Western Botany 1: 
105-127. 

HICKMAN, J. C. (ed.). 1993. The Jepson manual: Higher 
plants of California. University of California Press, 
Berkeley. 

HILEMAN, L. C., M. C. VASEy, AND V. T. PARKER. 2001. 
Phylogeny and biogeography of the Arbutoideae (Er- 
icaceae): Implications for the Madrean-Tethyan hy- 
pothesis. Systematic Botany 26:131—143. 

JEPSON, W. L. 1922. Revision of Californian Arctostaphyli. 
Madrono 1:78—86. 

KEELEY, J. E. 1994. Arctostaphylos rainbowensis, a new 
burl-forming manzanita from northern San Diego 
County, California. Madrono 41:1—12. 

, L. BOYKIN, AND A. MASssIHI. 1997a. Phenetic 

analysis of Arctostaphylos parryana: (1) Two new 

burl-forming subspecies. Madrono 44:253—267. 

, A. MAssiHt, J.D. RODRIGUEZ, AND S.A. HIRALES. 
1997b. Arctostaphylos incognita, a new species and 
its phenetic relationship to other manzanitas of Baja 
California. Madrofio 44:137—150. 

MARKOs, S., L.C. HILEMAN, M.C. VASEY AND V.T. PARKER. 
1998. Phylogeny of the Arctostaphylos hookeri com- 
plex (Ericaceae) based on nrDNA data. Madrono 45: 
187-199. 

MITERMEIER, R.A., N. MEYERS, C.G. MITERMEIER, AND N. 
MEYERS. 2000. Hotspots: Earth’s biologically richest 
and most endangered terrestrial ecosystems. Univer- 
sity of Chicago Press, Chicago, IL. 

REJMANEK, M. 2001. Trees and shrubs of California by 
John D. Stuart and John O. Sawyer: A book review. 
Madrono 48:128—129. 

STEBBINS, G. L. AND J. Mayor. 1965. Endemism and spe- 
ciation in the California flora. Ecological Monographs 
35:79-102. 

STUART, J. D. AND J. O. SAwYER. 2001. Trees and shrubs 
of California. University of California Press, Berke- 
ley. 

VASEY, M. C. AND V. T. PARKER. 1999. Nascent inflores- 
cences in Arctostaphylos pringlei: response to Keeley 
and Wells. Madrofo 46:51—54. 

WELLS, P. V. 1992. Subgenera and sections of Arctostaph- 
ylos. The Four Seasons 9:64—69. 

. 2000. The manzanitas of California, also of Mex- 

ico and the world. Published by the Author. 


MADRONO, Vol. 49, No. 1, p. 48, 2002 


Illustrated field guide to selected rare plants of 
northern California. Edited by Gary Nakamua and 
Julie Kiersteand Nelson. 2001. University of Cali- 
fornia Agriculture and Natural Resources Publica- 
tion 3395, Oakland, CA. 370 pp. Softcover $36.00. 
ISBN 1-879906-5470. 


For those interested in the botany of northern 
California, this field guide is a real treat. This book 
richly illustrates and describes 149 of the rarer plant 
taxa found in northern 10 counties in the state 
(Butte, Del Norte, Glenn, Humboldt, Lassen, Mo- 
doc, Plumas, Shasta, Siskiyou, and Trinity Coun- 
ties). The authors focus mainly on plants presumed 
extinct or rare in California, Lists 1A and 1B re- 
spectively in the 6th edition CNPS Inventory 
(CNPS 2001). Also included are six List 2 (rare in 
California, but more common elsewhere), one on 
List 3 (review list), and four on List 4 (watch list). 
The 149 taxa include three that are state-listed as 
endangered, 12 that are state-listed as rare, eight 
that are federally-list as endangered, and three that 
are federally listed as threatened. 

It is easy to jump over the first 40 pages of in- 
troduction to enjoy the species descriptions. Each 
is a two page treatment. The spiral binding allows 
for quick access. Plants are arranged alphabetically 
by genus, with the name in the upper left and lower 
right corners to make it easy to find a plant. Each 
treatment involves a large photograph of a mature 
plant, a small photograph of its habitat, and a map 
indicating occurrences by USGS quadrangle loca- 
tion. 

On the facing page, a line art illustration accom- 
panies a description of the plant, habitat, and lo- 
cation. Scientific names follow the new edition of 
the CNPS Inventory, as do synonymy, common 
name(s), family names, distribution, elevation, and 
quadrangle codes. Habitat designations generally 
follow those in the CNPS Inventory, but in some 
cases, the habitat descriptions are broader. In the 
Key Feature section, the first paragraph typically 
describes the plant. The second paragraph discusses 
similar taxa, and, if necessary, instructs the reader 
to “‘consult an expert to verify identification.” If 
this is the case, a list of diagnostic features accom- 
panies an expert symbol. Flowering times and iden- 
tification times, which may differ if fruits are re- 
quired, finish the presentation. Illustrations are 
drawn from 26 sources including lovely, original 
art by Linda Vorobik. 

The book has no keys, exhaustive descriptions, 
or complete synonymy. It assumes a basic knowl- 


edge of plant identification and is not intended to 
substitute for standard botanical references or field 
guides. Instead, the book’s purposes are to “. 

help the reader develop an accurate search image 

” and to “... learn how to accurately distin- 
guish rare plants from similar species in the field 

.. The editors involved 28 contributors, who 
make up the Northern California Botanists, an ad 
hoc committee of federal, state, and consulting bot- 
anists. More than other botanists, they are faced 
each field season with a new set of recruits to con- 
duct plant surveys. This book is designed for them, 
but it will be useful to seasoned botanists as well. 

Turning to the introduction, the reader finds the 
expected definitions, a short description of state and 
federal laws concerning protecting and conserving 
plants, and an explanation of how to use guide. 
Next come two extensive tables. The first lists spe- 
cies by geographic subdivisions of the state found 
in The Jepson Manual (Hickman 1993). The second 
table list species by habitat in each region. 

These tables are enlightening. As expected, rare 
plants along the coast were most common on the 
dunes, but the north coast conifer forest is not far 
behind. I expected to find a spate of serpentine spe- 
cies in the Klamath and North Coast Ranges, but 
not necessarily in mid-elevation forests and wood- 
lands. The same conclusion came from reviewing 
the lists for the Cascades, Sierra Nevada, and the 
Great Basin. The point was made most vividly 
when I saw the habitats of Cryptantha crinita and 
Lotus rubriflorus. The traveler on Interstate 5 drives 
by miles of similar looking dry streams and grass- 
lands. This book will help people get over the idea 
that all rare plants only grow in special places. With 
this book in hand and its great photographs, people 
can shed that misconception, as well as develop 
accurate search images for northern California’s 
rare plants. 

I doubt that you will find the book in most book- 
stores, but it can be ordered from the University of 
California, Agriculture and Natural Resources 
(ANR) Catalog at http://anrcatalog.ucdavis.edu/ 


15 


—JOHN O. SAWYER, Department of Biological Sciences, 
Humboldt State University, Arcata, CA 95521. 


LITERATURE CITED 


CNPS. 2001. Inventory of rare and endangered plants of 
California, 6th ed. Rare Plant Scientific Advisory 
Committee, David P. Tibor, Convening Editor. Cali- 
fornia Native Plant Society. Sacramento 

HICKMAN, J. C. (ed.). 1993. The Jepson manual: Higher 
plants of California. University of California Press, 
Berkeley. 


MApDRONO, Vol. 49, No. 1, pp. 49-53, 2002 


NOTEWORTHY BRYOPHYTE RECORDS FROM THE MOJAVE DESERT 


LLoyD R. STARK 
Department of Biological Sciences, University of Nevada, 
4505 Maryland Parkway, Las Vegas, NV 89154-4004 


ALAN T. WHITTEMORE! 
Missouri Botanical Garden, PO. Box 299, St. Louis, MO 63166 


BRENT D. MISHLER 
University Herbarium, Jepson Herbarium, and Department of Integrative Biology, 
1001 Valley Life Sciences Bld., #2465, University of California, 
Berkeley, CA 94720-2465 


ABSTRACT 


Significant range expansions in the northern Mojave Desert are documented for twenty-three species 
of bryophytes, nine of which are new to the region. Barbula convoluta, Claopodium whippleanum, En- 
tosthodon planoconvexus, Fissidens sublimbatus, Grimmia americana, Reboulia hemispherica, Syntrichia 
bartramii, Weissia condensa, and Asterella californica are new to the flora of Nevada. 


The northern Mojave Desert encompasses four 
counties: Clark and Nye Cos., Nevada, Washington 
Co., Utah, and Mohave Co., Arizona (MacMahon 
and Wagner 1985). Recently, a list of bryophyte 
species from this region was compiled from the lit- 
erature (Stark and Whittemore 2000), drawing chief- 
ly from state and regional checklists and treatments 
(Lawton 1958; Haring 1961; Flowers 1973; Spence 
1988). The bryophyte flora of the region consists 
of 75 species of mosses and 5 species of liverworts. 
More than half of the bryophytes from the northern 
Mojave Desert belong to the more xeric families 
Pottiaceae, Grimmiaceae, and Orthotrichaceae. 

Ongoing collecting efforts in southern Nevada 
indicate that this region is not well explored bryo- 
logically, and this applies to the entire state (Heise 
2000). The recent discovery of a new species of 
Didymodon in the remote southeastern portion of 
Nevada (Zander et al. 1995), and an as yet unde- 
scribed species in the liverwort genus Targionia 
that appears to be endemic to the Mojave Desert 
(Whittemore 1996) indicate exploration is needed. 
In the present paper, we discuss several species of 
bryophytes that are reported new to the northern 
Mojave Desert, or whose distributions are consid- 
erably broadened within the region. Nomenclature 
follows Anderson et al. (1990), Zander (1993), and 
Stotler and Crandall-Stotler (1977). 


BRYOPHYTA 


Barbula convoluta Hedw. 


Nevada, Clark Co., southern Gale Hills, lower 
end of Lovell Wash, 0.3 km upstream of confluence 


' Present address: U.S. National Arboretum, 3501 New 
York Avenue NE, Washington, DC, USA 2002-1958 


of Lovell Wash and West End Wash, along steep, 
north-facing outcrop adjacent to dirt road, elev. 550 
m, Stark NV-1941 (UNLV), 1942 (UNLV, BUF). 
Nevada state record. This is the first report of this 
rather wide-ranging species from the Mojave Des- 
ert, with the only other report from the interior ba- 
sins of North America (southern Idaho, Flowers 
1973; Spence 1988). From western North America, 
B. convoluta is known from British Columbia to 
Baja California (Lawton 1971; Zander 1994a). 


Claopodium whippleanum (Sull. in Whipple & 
Ives) Ren. & Card. 


Nevada, Clark Co., Spring Mountains, Red Rock 
Canyon National Recreation Area, steep side can- 
yon near mouth of Red Rock Canyon, near conflu- 
ence with Red Rock Wash; in deep shade beneath 
boulders, on sandstone rock and dead wood, elev. 
1450 m, Stark NV-316 (UNLV). Nevada state rec- 
ord. Found in an area that never receives direct sun- 
light, in a side canyon on a steep slope under boul- 
ders. In North America, the species ranges from 
northwestern Mexico to British Columbia (Crum 
and Buck 1994), and is reported from southern Cal- 
ifornia as occasional in cismontane lowlands (Har- 
thill et al. 1979). It is distributed in western North 
America and also the Mediterranean region (Scho- 
field and Crum 1972). A disjunct population was 
reported from high elevation in northeastern Ari- 
zona (Apache Co., 9500 ft, Haring 1961). 


Coscinodon calyptratus (Hook. in Drumm.) C. 
Jens. ex Kindb. 


Reported from the Mojave Desert of southwest- 
ern Utah (Hastings 1999). Previously reported from 
Mohave County, Arizona (Haring 1961, as Grim- 
mia calyptrata Hook.), and from Lincoln Co., Ne- 


50 MADRONO 


vada (Lawton 1958, as Grimmia calyptrata Hook.). 
This species is broadly distributed in Nevada and 
Utah, reaching its southwesternmost extent in the 
Mojave Desert (Hastings 1999). 


Crossidium seriatum Crum & Steere 


Nevada, Clark County, northern foothills of 
Lime Ridge, ca. 16 km north of Gold Butte, elev. 
487 m, Stark NV-232a (UNLV, MEXU); southern 
Moapa Valley, sandstone bluffs along periphery of 
valley, ca. 8 km south of Overton, along Hwy 169, 
elev. 488 m, Stark NV-872 (UNLV, MEXU); Black 
Mountains, Lake Mead National Recreation Area, 
gypsum formation 2.1 km down Boathouse Cove 
Road from North Shore Road turnoff, elev. 650 m, 
Stark & Bonine NV-3045 (UNLV); Arizona, Mo- 
have County, Lake Mead National Recreation Area, 
Lake Mead Landing, mouth of Kingman Wash, 
elev. 396 m, W. Niles s.n., 24 Feb 1995 (UNLV). 
Recently recorded from the state of Nevada (Zander 
et al. 1995), the range of C. seriatum is expanded 
to a scattering of southern Nevada locations and 
also into Mohave County, Arizona. This globally 
rare species is presently known from only a handful 
of populations worldwide outside of the state of 
Nevada: Mariposa County, Arizona (Zander 1977), 
Cedros Island, Mexico, San Diego County, Cali- 
fornia (Stark and Whittemore 1992), and Spain 
(Cano et al. 1992). 


Didymodon vinealis (Brid.) Zand. 


Nevada, Clark County, Southern Gale Hills, low- 
er end of Lovell Wash, 0.3 km upstream of conflu- 
ence of Lovell Wash and West End Wash, along 
steep, north-facing outcrop adjacent to dirt road, 
elev. 550 m, Stark NV-1946 (UNLV); Lake Mead 
National Recreation Area, 4.3 km south of Rogers 
Spring, narrow canyon on north side of North 
Shore Rd, elev. 750 m, Stark NV-2059a (UNLV); 
Newberry Mountains, Lake Mead National Recre- 
ation Area, “‘Needles Eye’’, ca. 7.2 km north on 
Christmas Tree Pass Rd from Hwy 77E, elev. 817 
m, Stark NV-76 (UNLV). Recently reported from 
the northern Mojave from the River Mountains as 
an incidental species (Stark 1997), the range of D. 
vinealis is considerably broadened here. 


Entosthodon planoconvexus (Bartr.) Grout 


Nevada, Nye County, Nevada Test Site, Rock 
Valley, north-facing foothills of Spectre Range, 
elev. 1159 m, growing with an undescribed species 
of Targionia, Stark NV-724 (UNLV). Nevada state 
record, and one of only four localities worldwide. 
Other known localities include Washington County, 
Utah (Flowers 1973), Pima County, Arizona (Har- 
ing 1961), and the northern Egyptian desert (Shab- 
bara 1999). The specimen cited herein (Stark NV- 
724) differs from descriptions of E. planoconvexus 
in having a pale (not red) seta, a relatively narrow 
capsule that is strongly contracted under the mouth 


[Vol. 49 


when dry, and a rudimentary peristome. The latter 
features align it with E. tucsonii (Bartr.) Grout. 
However, E. tucsonii has spores twice as large as 
Stark NV-724, making this determination improb- 
able. Clearly, a revision of North American Entos- 
thodon is needed. 


Fissidens sublimbatus Grout 


Nevada, Clark County, Newberry Mountains, 
Lake Mead National Recreation Area, Pipe Spring 
Canyon, near Pipe Spring, west-facing soil in rock 
crevice, elev. 732 m, Stark NV-99 (UNLV, PAC). 
New to the northern Mojave Desert and a Nevada 
state record. Known also from Arizona, California, 
New Mexico, and Baja California (Pursell 1994). 


Funaria hygrometrica Hedw. 


Nevada, Clark County, Newberry Mountains, 
Lake Mead National Recreation Area, ‘‘Needles 
Eye,” ca. 7.2 km north on Christmas Tree Pass Rd 
from Hwy 77E, elev. 817 m, Stark NV-SO (UNLV); 
Virgin Mountains, east base of South Virgin Peak 
Ridge, Lime Spring Canyon, abundant in moist 
drainages, W. Niles s.n., 22 Mar 1996 (UNLYV). 
This cosmopolitan species is reported as new to 
southern Nevada, having previously been reported 
from Mohave County, Arizona (Haring 1961). 


Funaria muhlenbergii Turn. 


Nevada, Clark County, northern River Moun- 
tains, Lake Mead National Recreation Area, 5.6 km 
due east of Saddle Island, elev. 610 m, Stark NV- 
144 (UNLV); Eldorado Mountains, Keyhole Can- 
yon Archeological Site, just north of base of Key- 
hole Canyon, elev. 274 m, Stark NV-190b (UNLY); 
Muddy Mountains, southern end of White Basin, 
adjacent to West Longwell Ridge, ca. 6.4 km by 
road northwest of Bitter Spring, elev. 650 m, Stark 
& Bonine NV-3013 (UNLV). Reported from the 
California Mojave (Harthill et al. 1979), and broad- 
ly distributed in western North America (Smith 
1994). 


Grimmia americana Bartr. 


Nevada, Clark County, Newberry Mountains, 
Lake Mead National Recreation Area, Grapevine 
Canyon, on north-facing rock, elev. 793 m, Stark 
NV-16 (UNLV). Nevada State record, and one of 
only three populations of this species known world- 
wide, one from Arizona (Pima Co., Crum 1994), 
and one from western Texas (type locality, Jeff Da- 
vis County, Texas; Crum and Anderson 1981). This 
species is not listed in Haring (1961) as occurring 
in Arizona, and is not yet known from Mexico 


(Crum 1994). Grimmia americana has bistratose 


upper leaf cells, which distinguish it from G. pla- 
giopodia Hedw., and is peristomate, which distin- 
guishes it from G. anodon Bruch & Schimp. in 
B.S.G. Grimmia plagiopodia is listed in Haring 


2002] 


(1961) as occurring only in Yavapai County, Ari- 
zona, from two collections, while G. anodon 1s 
widespread in the southwestern U.S. (Lavin 1982). 


Grimmia moxleyi Williams in Holz. 


Nevada, Clark County, Muddy Mountains, 
southern end of White Basin, adjacent to West 
Longwell Ridge, ca. 6.4 km by road northwest of 
Bitter Spring, elev. 650 m, Stark & Bonine NV- 
3004 (UNLV); foothills of Black Mountains, Lake 
Mead National Recreation Area, along tributary to 
Manganese Wash (from the northwest), ca. 6.4 km 
north of Boathouse Cove, elev. 650 m, Stark & 
Bonine NV-3071 (UNLV). New to the northern Mo- 
jave Desert exclusive of California. Grimmia mox- 
leyi is reported from the California Mojave Desert 
as infrequent (Harthill et al. 1979), and its presence 
is expected based on a Death Valley locality noted 
in Koch (1954). A common associate of G. orbi- 
cularis Bruch ex Wils. in Nevada, G. moxleyi is 
endemic to the southwestern U.S. and adjacent 
northern Mexico (Greven 1999). It has been re- 
ported without specifics from Arizona and Nevada 
(Crum 1994). Recently, Munoz (2000) regarded G. 
moxleyi aS synonymous with the wide-ranging G. 
orbicularis, indicating ample variability in hair- 
point development among southwestern popula- 
tions. 


Homalothecium nevadense (Lesq.) Ren. & Card. 


Nevada, Nye County, Spring Mountains, Wood 
Canyon, in vicinity of Wood Canyon Spring, Stark 
& Landau NV-1794 (UNLV). First report for the 
Mojave Desert. This species is reported from south- 
ern California (Harthill et al. 1979) as frequent in 
the region, but excluding deserts. The species 
reaches its southernmost extent in northern Arizona 
(Flowers 1973). 


Hypnum vaucheri Lesq. 


Nevada, Clark County, Spring Mountains, Red 
Rock Canyon National Recreation Area, steep side 
canyon near mouth of Red Rock Canyon, near con- 
fluence with Red Rock Wash, on shaded rock, elev. 
1450 m, Stark NV-315b (UNLV). New to the Mo- 
jave Desert. Frequent in southern Utah along the 
Colorado-Green River Basin (Flowers 1973). 


Microbryum starkeanum (Hedw.) Zand. 


Nevada, Clark County, lower end of Borax 
Wash, southern Gale Hills, Stark NV-1944 (UNLV). 
New to the Mojave Desert. Guerra and Cano (2000) 
prefer to retain this species in Pottia, as Pottia 
starckeana (Hedw.) Miill., because of its stegocar- 
pous sporophyte. 


Pseudocrossidium crinitum (Schultz) Zand. 


Nevada, Clark County, Muddy Mountains, Val- 
ley of Fire State Park, 0.5 km south of ‘“‘Mouse’s 


STARK ET AL.: MOJAVE DESERT BRYOPHYTES 51 


Tank,’’ sandstone bluffs on west side of road, ele- 
vation 700 m, Stark & Bonine NV-3076, 3084 
(UNLV). Second report for state of Nevada. The 
locality cited may be the same populations noted 
by Lawton (1958, as Tortula aurea Bartr.). This 
report thus confirms the only known population 
from the Mojave Desert, with the nearest known 
locality just outside the Mojave Desert in southern 
Utah (Spence 1987). This species is fairly common 
in Mexico, according to Zander (1994b), occurring 
in several Mexican states, and is listed from five 
counties in Arizona (Haring 1961). Despite the 
abundance of P. crinitum at this site, no sporo- 
phytes were found, consistent with the pattern of 
an absence of male plants in North America. 


Pterygoneurum subsessile (Brid.) Jur. 


Nevada, Clark County, Eldorado Mountains, 
Keyhole Canyon Archeological Site, just north of 
base of Keyhole Canyon, elev. 274 m, Stark NV- 
192a (UNLV); Newberry Mountains, Lake Mead 
National Recreation Area, Grapevine Canyon, 3.2 
km north on Christmas Tree Pass Rd from Hwy 
77E, beyond petroglyphs to the east, elev. 854 m, 
Stark NV-22c (UNLV). Second report from the 
northern Mojave Desert; found previously in Wash- 
ington County, Utah (Flowers 1973). 


Syntrichia bartramii (Steere in Grout) Zand. 


Nevada, Clark County, Newberry Mountains, 
Lake Mead National Recreation Area, Grapevine 
Canyon, 3.2 km north on Christmas Tree Pass Rd 
from Hwy 77E, beyond petroglyphs to the east, 
elev. 854 m, Stark NV-32B, 38B (UNLV). Nevada 
state record. Previously reported from the southern 
Californian Mojave Desert (Harthill et al. 1979), S. 
bartramii occurs in northwestern Mexico and in the 
bordering states of Arizona, New Mexico, and Tex- 
as, with infrequent reports north of this region 
(Mishler 1994). One of the two specimens cited 
above from Nevada was found on juniper growing 
with S. pagorum, noteworthy in that epiphytic 
mosses are exceedingly rare in the Mojave Desert 
below 2000 m. Variation in this species is complex 
and should be studied in association with plants oc- 
curring in southern California, where it is possible 
that an undescribed species is present. 


Syntrichia pagorum (Milde) Amann 


Nevada, Clark County, Newberry Mountains, 
Lake Mead National Recreation Area, “‘Needles 
Eye,” ca. 7.2 km north on Christmas Tree Pass Rd 
from Hwy 77E, elev. 817 m, Stark NV-82A 
(UNLV); Pipe Spring Canyon, in vicinity of Pipe 
Spring, elev. 732 m, Stark NV-91 (UNLV). New to 
the Mojave Desert. Previously reported from Ne- 
vada (Crum and Anderson 1981), this species is 
distributed from eastern North America across the 
southern portion of the USA to the west coast. 
However, reports are lacking for Utah and southern 


52 MADRONO 


California. Syntrichia pagorum is known only from 
female plants in the USA. 


Syntrichia princeps (De Not.) Mitt. 


Nevada, Clark County, Spring Mountains, Red 
Rock Canyon National Recreation Area, near 
mouth of Red Rock Canyon, near confluence with 
Red Rock Wash, on partially shaded rock, elev. 
1450 m, Stark NV-291 (UNLV); Virgin Mountains, 
east base of South Virgin Peak Ridge, Lime Spring 
Canyon, on north-facing slope, at edge of water- 
course on moist drainages, W. Niles s.n., 22 March 
1996 (UNLV). These two localities constitute the 
second and third reports from the northern Mojave 
Desert, with the first from Washington County, 
Utah (Flowers 1973). Syntrichia princeps 1s distin- 
guished from the related S. ruralis (Hedw.) Web. & 
Mohr by its (often) synoicous condition, and is dis- 
junct from the western coast of North America to 
the Spring Mountains and Virgin Mountains of 
southern Nevada and southern Utah, respectively. 
Syntrichia princeps is one of several species in 
Utah known only from Washington County (Flow- 
ers 1973). 


Syntrichia ruralis (Hedw.) Web. & Mohr 


Nevada, Muddy Mountains, Valley of Fire State 
Park, 0.5 km south of ‘““Mouse’s Tank,’’ sandstone 
bluffs on west side of road, elevation 700 m, Stark 
& Bonine 3077, 3078 (UNLV). Oddly, S. ruralis is 
known from only two sites from the northern Mo- 
jave Desert, from Mohave County, Arizona (Haring 
1961), and from the River Mountains in Nevada 
(Stark et al. 1998). The locality given represents 
one of the few sites known to the authors in the 
Mojave Desert where male, female, and sporo- 
phytic plants co-occur. 


Tortula atrovirens (Sm.) Lindb. 


Nevada, Clark County, Bitter Spring Valley, 
Echo Wash, gypsum formation ca. 1.6 km east of 
Bitter Spring, elev. 530 m, Stark NV-2087 (UNLV); 
northern River Mountains, Lake Mead National 
Recreation Area, 5.6 km due east of Saddle Island, 
elev. 610 m, Stark NV-139a (UNLV); Newberry 
Mountains, Lake Mead National Recreation Area, 
Grapevine Canyon, 3.2 km north on Christmas Tree 
Pass Rd from Hwy 77E, beyond petroglyphs to the 
east, elev. 854 m, Stark NV-39a (UNLV). Only a 
single prior report exists from the northern Mojave 
Desert exists, and this as an incidental (Stark et al. 
1998), thus indicating that the species is probably 
much more common than reports indicate. Tortula 
atrovirens was previously known as Desmatodon 
convolutus (Brid.) Grout. 


Weissia condensa (Voit) Lindb. 


Nevada, Clark County, foothills on northwest 
side of River Mountains, ca. 6.4 km from down- 


[Vol. 49 


town Henderson, elev. 671 m, Stark NV-120 
(UNLV, BUF). First report from southern Nevada 
and probably a state record. Reported from Arizo- 
na, southern Utah, Texas, and Mexico, W. condensa 
is a widespread species that also occurs in South 
America, Africa, and Europe (Flowers 1973, as W. 
tortilis (Schwaegr.) C. Muell.; Zander 1994c). 


HEPATICOPHYTA 
Asterella californica (Hampe) Underw. 


Nevada, Clark County, Lake Mead National Rec- 
reation Area, Indian Hills, north of Devil’s Cove, 
Gold Butte area, T20S, R7OE, at base of limestone 
boulders, elevation 650 m, W. E. Niles s.n., 6 
March 1998 (UNLV); Spring Mountains, Red Rock 
National Conservation Area, near mouth of Red 
Rock Canyon near confluence with Red Rock 
Wash, T20S, R58E, S32, damp shaded soil beneath 
overhang at base of north-facing cliff, elev. 1450 
m., A. T. Whittemore 68587 (MO). Nevada state rec- 
ord. Known from Mohave Co., Arizona (Evans 
1917) and from Riverside Co., CA, near Palm 
Springs (S. B. Parish 3890, CAS). 


Reboulia hemispherica (L.) Raddi 


Nevada, Clark County, Spring Mountains, Red 
Rock National Conservation Area, near Willow 
Springs, T21S, R58E, S5, shaded gorge, in cracks 
in cliff, elev. 1500 m, A. 7. Whittemore 6891 (MO). 
Nevada state record. This is the first report of this 
rather wide-ranging species from the Mojave Des- 
ert. Reboulia hemispherica is fairly common in 
New Mexico and eastern Arizona, but it is rare and 
local west of these states, currently known only 
from two collections from the northern Sierra Ne- 
vada in California and a few scattered sites in the 
Pacific northwest. 


ACKNOWLEDGMENTS 


We thank the National Geographic Society (grant no. 
5429-95) for providing funds for travel and the Missouri 
Botanical Garden for administering this grant; the U.S. 
Bureau of Land Management and the National Park Ser- 
vice (Lake Mead) for funding support; Wes Niles for con- 
tributing specimens critical to this study; Mary Bonine, 
Fred Landau, and Robin Stark for assistance in the field; 
Ronald Pursell for identifications of Fissidens and Richard 
Zander for critical determinations in the Pottiaceae; Jesus 
Munoz for information on the distribution of Grimmia 
americana; the granting of collecting permits from the 
U.S. Bureau of Land Management, Las Vegas office (Gay- 
le Marrs-Smith), Lake Mead National Recreation Area 
(Elizabeth Powell and Jennifer Haley), U.S.D.A. Forest 
Service (Kerwin Dewberry), Nevada State Parks, and the 
Nevada Test Site; Harold Robinson and Robert Ireland for 
reviewing the manuscript; and Bruce Allen, Bruce Lund, 
and Philip Medica for logistic support. 


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ico. Pt. 1. Memoirs of the New York Botanical Gar- 

den 69, New York. 

. 1994b. Pseudocrossidium. Pp. 296-299 in A. J. 

Sharp, H. Crum, and P. M. Eckel (eds.), The moss 

flora of Mexico. Pt. 1. Memoirs of the New York 

Botanical Garden 69, New York. 

. 1994c. Weissia. Pp. 213-225 in A. J. Sharp, H. 

Crum, and P. M. Eckel (eds.), The moss flora of Mex- 

ico. Pt. 1. Memoirs of the New York Botanical Gar- 

den 69, New York. 

, L. R. STARK, AND G. MARRS-SMITH. 1995. Didy- 

modon nevadensis, a new species for North America, 

with comments on phenology. Bryologist 98:590— 

595. 


MADRONO, Vol. 49, No. 1, pp. 54-58, 2002 


NOTEWORTHY COLLECTIONS 


CALIFORNIA 


BACCHARIS MALIBUENSIS R. M. Beauch. & Henr. (AS- 
TERACEAE).—Orange Co., [Santa Ana Mountains] 
North Ranch Policy Plan Area [proposed for inclusion in 
the NCCP], Fremont Canyon, N. of Santiago Creek and 
immediately S. of major stream fork [of the Fremont Can- 
yon drainage], 33.7907°N 117.6801°W, UTM Zone 11 
N3740484 E435012, ca. 305 m, 23 Aug 2000, Riefner & 
Wolf 20-732 (RSA). 

Previous knowledge. Baccharis malibuensis was de- 
scribed by Beauchamp and Henrickson in 1996 (Aliso 14: 
197-203) as a narrow endemic restricted to the Malibu 
Creek drainage in the Santa Monica Mountains, extreme 
western Los Angeles County. At that time, the species was 
known from 5 localities on private land within an area 
just over 18 km’, growing on volcanic and sedimentary 
substrates in chaparral, openings in scrub, and in the un- 
derstory of Quercus agrifolia Née (Beauchamp & Hen- 
rickson 1996, loc. Cit.). 

Significance. The Riefner & Wolf collection of B. mal- 
ibuensis from the Santa Ana Mountains represents the first 
record for that range, the first record for Orange County, 
and a disjunction of 93 km south from the southernmost 
station in the Santa Monica Mountains. At the Fremont 
Canyon site, the plant was rare, growing at the base of a 
north-facing slope in the understory of a Q. agrifolia ri- 
parian woodland along an intermittent stream course. As- 
sociated species reported for the site include Symphori- 
carpos mollis Nutt., Heteromeles arbutifolia (Lindley) 
Roem., Rhamnus sp., Toxicodendron diversilobum (Torrey 
& A. Gray) E. Greene, Artemisia douglasiana Besser, Pip- 
tatherum milaceum (L.) Cosson, and Ambrosia psilostach- 
ya DC. This region of Fremont Canyon is primarily un- 
derlain by marine sandstones of the Williams Formation, 
which consists of very resistant, cliff-forming, white to 
brownish-gray feldspathic sandstone, pebbly sandstone, 
and conglomeratic sandstone (Morton 1999, Open-File 
Report 99-172, U.S. Geological Survey). 

Aside from the Palos Verdes headlands, the Santa Ana 
Mountains are the closest near-coastal range south of the 
Santa Monica Mountains. Two other taxa, Dudleya cy- 
mosa (Lem.) Britton & Rose ssp. ovatifolia (Britton) Mor- 
an and Nolina cismontana Dice, exhibit a similar pattern 
of disjunction between the Santa Monica and Santa Ana 
Mountains. It is not completely surprising, therefore, to 
find that B. malibuensis is present in both ranges. In the 
Santa Monica Mountains, B. malibuensis is known only 
from private lands and the documented occurrences are 
represented by populations of very small size, quite vul- 
nerable to extirpation by development (Beauchamp & 
Henrickson 1996, Joc. cit.). For this reason the species was 
added to list 1B of the California Native Plant Society’s 
inventory of rare and endangered vascular plants, with 
rarity-endangerment-distribution (RED) of 3-3-3, their 
highest threat rating (www.cnps.org/rareplants/inventory/ 
6thEdition.htm). At the present time it appears that B. mal- 
ibuensis is also quite rare in the Santa Ana Mountains; 
only a single, pistillate, multi-branched subshrub approx- 
imately 8 dm high and 15 dm wide was located in 2000. 
Owing to the extremely rugged topography and relatively 
undisturbed character of the vegetation of this region it is 
likely that additional plants could be discovered in other 


nearby canyons that were not explored in 2000. Since B. 
malibuensis is also known from west- and south-facing 
slopes in clearings and dense chaparral in the Santa Mon- 
ica Mountains (Beauchamp & Henrickson 1996, loc. cit.) 
further surveys for B. malibuensis in similar habitats of 
the northern Santa Ana Mountains are warranted. 


—STEVE Boyb, Herbarium, Rancho Santa Ana Botanic 
Garden, 1500 N. College Avenue, Claremont, CA 91711. 


COLORADO AND NEw MEXIco 


ERIGERON OCHROLEUCUS Nutt. var. SCRIBNERI (Canby ex 
Rydb.) Cronquist (ASTERACEAE).—COLORADO: 
Weld County, rocky ridge 5.6 km N of Rockport, elev. 
1830 m, TIIN R66W sect. 6 E% NE%, 16 May 2000, 
Dorn 8222 (COLO, RM). 

Previous knowledge. A range extension of 68 km from 
Albany County, Wyoming. 

Significance. First report for Colorado. 


SALIX ARIZONICA Dorn (SALICACEAE).—COLORA- 
DO: Conejos County, wet meadow along streamlet off 
Red Lake Trail Road 0.8 km from Hy. 17, elev. 3140 m, 
37°04.7'N, 106°24.1'W, 6 July 2001, Dorn 8852 (COLO, 
MO, RM). 

Previous knowledge. A range extension of 105 km from 
Rio Arriba and Taos counties, New Mexico. 

Significance. First report for Colorado. 


SALIX DISCOLOR Muhl. (SALICACEAE).—COLORA- 
DO: Larimer County, bank of South Branch Boxelder 
Creek at County Road 37, elev. 2195 m, 40°57.7'N, 
105°14.8’W, 29 May 2001, Dorn 8752, same plant 10 July 
2001, Dorn 8894 (COLO, MO, RM). 

Previous knowledge. A range extension of 50 km from 
Laramie County, Wyoming. 

Significance. First report for Colorado. 


SALIX WOLFI Bebb var. WOLFII (SALICACEAE).—NEW 
MEXICO: Rio Arriba County, meadow along Osier Creek 
ca. 1.2 km SW of confluence with Rio de los Pinos, elev. 
2925 m, 36°59.6'N, 106°20.6’W, 5 July 2001, Dorn 8847 
(MO, NMC, RM, UNM). 

Previous knowledge. A range extension of ca. 40 m 
from adjacent Conejos County, Colorado or 2 km from 
nearest collection site in same county. 

Significance. First report for New Mexico. 


—RoBERT D. Dorn, Box 1471, Cheyenne, WY 82003. 


IDAHO 


CRYPTOGRAMMA STELLERI (S.G. Gmelin) Prantl (PTERI- 
DACEAE)—Boundary Co., Upper Priest Falls on Priest 
River, 1 km south of British Columbia border, 48°59’N, 
116°55'W, rare on seepy, crumbly calcareous rock around 
waterfall in forest of Tsuga heterophylla and Thuja pli- 
cata, with Asplenium viride, ca. 950—1050 m, 20 Jul 2001, 
T. Spribille 11177 (ID). 


2002] 


Previous knowledge. A widespread North American— 
Asiatic fern known from widely scattered localities in the 
western United States (E.R. Alverson, 1993, Cryptogram- 
ma. In: Flora of North America 2: 137—139) and infre- 
quent in adjacent British Columbia (G.W. Douglas et al., 
1991, Vascular Plants of British Columbia). 

Significance. First report for Idaho, a range extension 
of approximately 330 km west from the nearest known 
locality on the east side of Glacier National Park, Glacier 
Co., Montana. 


VIOLA SELKIRKII Pursh ex Goldie (VIOLACEAE)—Bon- 
ner Co., upper end of Priest Lake, along Ruby Creek, ca. 
48°50’N, 116°55’W; frequent in riparian Thuja plicata for- 
est, with Tiarella trifoliata, Viola glabella and Oplopanax 
horridus, ca. 730 m elev., 20 Jul 2001, T. Spribille & R. 
Merkel 11123 (ID). 

Previous knowledge. A widespread circumboreal spe- 
cies, rare in western North America, where it is known 
from widely scattered localities in Alaska, British Colum- 
bia, Alberta, Colorado (E. Hultén, 1968, Flora of Alaska 
and Neighboring Territories), New Mexico (W.C. Martin 
& C.R. Hutchins, 1980, Flora of New Mexico) and Mon- 
tana (T. Spribille et al., 2002, Noteworthy Collections, 
Madronio 49:55—58). 

Significance. First report for Idaho, a range extension 
of approximately 150 km west from the nearest known 
locality in the Whitefish Range, Lincoln Co., Montana. 


—Tosy SPRIBILLE, Kootenai National Forest, Fortine 
Ranger District, PO. Box 116, Fortine, MT 59918 (current 
address: Herbarium, Department of Systematic Botany, 
Albrecht von Haller Institute of Plant Sciences, University 
of Gottingen, Untere Karspiile 2, D-37073 Gottingen, Ger- 
many; e-mail toby.spribille@ gmx.de). 


MONTANA 


ALNUS RUBRA Bong. (BETULACEAE)—Lincoln Co., 
western Cabinet Mountains, Callahan Creek drainage, 
along Callahan Creek and North Callahan Creek at Mon- 
tana/Idaho state line and up to 3 km east into Montana, 
48°26'30"N 115°57'—116°00'W, in alluvial bottoms and on 
moist slopes with Thuja plicata, Tsuga heterophylla and 
Betula papyrifera, 850-915 m, 16 Jun 1998, T. Spribille 
& M. Arvidson 7788 (USFS Fortine District Herbarium, 
COLO, MONTU, MRC). 

Previous knowledge. This primarily coastal species has 
a limited distribution in the interior and is usually asso- 
ciated with inland rainforest communities (Johnson & 
Steele, 1978, Northwest Sci. 52(3): 205—211; C.C. Lorain, 
1988, Floristic history and distribution of coastal disjunct 
plants of the northern Rocky Mountains, M.Sc. thesis, 
Univ. Idaho). It is widespread along the Pacific Coast and 
is known from disjunct inland populations in British Co- 
lumbia, Washington and Idaho. 

Significance. First report for Montana, representing the 
eastern limits of species distribution, a contiguous range 
extension across the state line from nearby Boundary Co., 
Idaho. As in the Idaho locations, there appears to be in- 
trogression with Alnus incana ssp. tenuifolia in at least 
some individuals. The first reports of red alder from this 
area came from forest stand examination contractors who 
reported A/nus in the area as large as 59 cm dbh. 


AZOLLA MEXICANA Presl. (AZOLLACEAE)—Ravalli Co., 
McCalla Creek, app. 2 km west of Stevensville, 


NOTEWORTHY COLLECTIONS 55 


46°30'20"N_ 114°07'40"W, locally common and forming 
small mats, with Callitriche heterophylla and Elodea can- 
adensis, 1030 m, 18 Sep 1999, W.E. Albert 3261 (MRC). 
Verified by PE Stickney (MRC). 

Previous knowledge. Widespread across the western 
states and British Columbia to South America, scattered 
east to the Mississippi River (T. A. Lumpkin, 1993, Azol- 
laceae. In: Flora of North America 2: 338-342). 

Significance. First report for Montana, probably intro- 
duced with recent disturbance, persisting and spreading in 
similar nearby drainages and a nearby slough. The nearest 
known collection stations are 320 km southwest and 
southeast, respectively, in Ada and Bannock Cos., Idaho 
(specimens at ID). 


BOTRYCHIUM PEDUNCULOSUM W.H. Wagner (OPHIO- 
GLOSSACEAE)—Lincoln Co., Big Creek, 48°44'30"N 
115°28’30W, rare in moonwort genus communities on 
floodplains under Thuja plicata, 975 m, 24 Jul 1996, J. 
Vanderhorst 5609 (MONTU); Big Creek, 48°46'’N 
115°27'W, 9 Aug 1997, J. Vanderhorst 5617 (MONTU); 
Cedar Creek, 10 km west of Libby, 48°24’35’N 
115°40'59’W, with Thuja plicata, 1070 m, 1 Aug 1997, 
R. Ferriel s.n. (USFS Kootenai NF Herbarium); Quartz 
Creek, 17 km northwest of Libby, 48°30'50’N 
115°42'07’W, with Tsuga heterophylla, Thuja plicata and 
Athyrium felix-femina, 910 m, 15 Aug 1997, R. Ferriel 
s.n. (USFS Kootenai NF Herbarium); Davis Creek, 20 km 
southwest of Trego, 48°31'37"N 114°57'40’W, in roadbed 
and powerline corridor, 1150 m, 22 Jul 1998, R. Ferriel 
RF98017 (USFS Kootenai NF Herbarium): Sanders Co., 
0.8 km northeast of Noxon Rapids Dam, 47°57'53"N 
115°43'21”W, mesic meadow, 735 m, 27 Jun 1999, R. Fer- 
riel RF99014 (USFS Kootenai NF Herbarium). Vander- 
horst 5609, 5617 determined by W. H. Wagner (MICH). 

Previous knowledge. A rare species of northwestern 
North America from southwestern Saskatchewan west to 
British Columbia and Oregon, east of the Coast/Cascade 
Ranges (W.H. Wagner & ES. Wagner, 1993, Ophioglos- 
saceae. In: Flora of North America 2: 85—106). 

Significance. First reports for Montana, a range exten- 
sion of approximately 130 km east from Pend Oreille Co., 
Washington. 


CAREX CHALCIOLEPIS Holm (CYPERACEAE)—Ravalli 
Co., Anaconda-Pintler Wilderness area, wet meadow that 
parallels both sides of the creek draining Hidden Lake, 
45°55'10’N 113°33'00'W, 2500 m, Mooers & Mooers 998 
(MONT). Determined by T. Spribille, verified by D.E 
Murray (ALA). 

Previous knowledge. A species of subalpine to alpine 
meadows in the southern and central Rocky Mountains, 
described by Holm (1903, Amer. J. Sci. 16: 17—44). Mur- 
ray (1969, Brittonia 21: 55—76) expressed doubt as to the 
occurrence of this species in Montana, because the ma- 
terial available to him was too immature to be certain of 
its identity. 

Significance. First report for Montana, representing the 
northern limits of species distribution, a range extension 
of over 300 km northwest from the nearest reported lo- 
cation in Park County, Wyoming. 


CAREX DEFLEXA Hornem. var. BooTTI L.H. Bailey (CY- 
PERACEAE)—Beaverhead Co., Beaverhead National 
Forest, Stine Mtn., West Pioneer Range, growing in Larix 
lyallii grove, 2640 m, 21 Jul 1968, S.F. Arno 29 (MON- 
TU); Beaverhead Co., Lima Peaks, one mile east of Gar- 
field Mtn., 8 miles south of Lima, common forming small 
patches of turf in stony, quartzite-derived soil of an alpine 


56 MADRONO 


fellfield, 3030 m, 27 Jul 1989, P. Lesica & S. Cooper 
4966 (MONTU); Missoula Co., Flathead National Forest, 
Lindy Peak, Mission Range, beneath stunted Larix lyallii, 
2515 m, | Sep 1968, S.F. Arno 285 (MONTU); Missoula 
Co., Bitterroot Mtns., fellfield on Onehorse Ridge, Lolo 
Peak massif, 2575 m, 21 Jul 1971, K. Lackschewitz 2953 
(MONTU); Ravalli Co., East Boulder Peak, growing in 
talus enclosure within alpine larch [stand], SE slope, 2710 
m, 7 Aug 1968, K. Lackschewitz & T. Fageraas 596 
(MONTU); Ravalli Co., dry, rocky crags, W-slope of the 
Castle Crags, 2590 m, 18 Aug 1970, K. Lackschewitz & 
Stuart 2365 (MONTU); Ravalli Co., NE slope of Canyon 
Peak, Canyon Lakes Basin, beneath alpine larch, 2590 m, 
22 Aug 1971, K. Lackschewitz 3337 (MONTU); Ravalli 
Co., St. Joseph Peak, wind-timber zone, 2740 m, 24 Jul 
1971, K. Lackschewitz & Gouaux 2979 (MONTU). All 
specimens determined by T. Spribille and verified by A.A. 
Reznicek (MICH). 

Previous knowledge. Although previously reported for 
Montana by Rydberg (1900, Flora of the Rocky Moun- 
tains) and EJ. Hermann (1970, Manual of the Carices of 
the Rocky Mountains and Colorado Basin, USDA Agr. 
Handb. 374) as Carex brevipes W. Boott, this species was 
subsumed under Carex rossii Boott in Hook. by C.L. 
Hitchcock et al. (1969, Vascular plants of the Pacific 
Northwest, Vol. 1), and has since escaped mention in the 
floras of Montana. The species is distinct from C. rossii 
both morphologically and ecologically, being a distinctly 
subalpine to alpine taxon with an apparent affinity for La- 
rix lyallii stands near the alpine timberline, in contrast to 
the more montane, xerothermic C. rossii. Furthermore, C. 
deflexa is more widespread across boreal North America 
than the primarily western C. rossi. 

Significance. These reports reaffirm the presence of this 
species in Montana. 


CAREX LACUSTRIS Willd. (CYPERACEAE)—Lake Co., 
small glacial pothole marsh ca. 6 km south of Swan Lake, 
47°52'15"N 113°49'50"W, with Carex utriculata and C. 
lasiocarpa, 945 m, 15 Jul 1989, P. Lesica 4893 (MICH, 
MONTU); Swan River Valley, Lost Creek Fen, ca. 5 km 
south of village of Swan Lake, 47°52'55"N 113°49'42"W, 
in mossy saturated peat of fen, with Betula glandulosa 
and Carex lasiocarpa, 965 m, 19 Jun 1992, J.S. Shelly & 
S. Chadde 1652 (MICH). Both specimens determined by 
A.A. Reznicek (MICH). 

Previous knowledge. A species of the Great Lakes and 
Great Plains, rarely as far west as Idaho (M.L. Fernald, 1942, 
Rhodora 44: 281—331; Great Plains Flora Committee, 1977, 
Atlas of the Flora of the Great Plains), north-central Alberta 
(J.G. Packer, 1983, Flora of Alberta), and reportedly also 
British Columbia, although this report was based on a mis- 
identification (A. Ceska personal communication). 

Significance. First reports of this species for Montana. 
It joins a suite of species of the eastern deciduous wood- 
land region (e.g., Carex comosa, Carex pallescens, Dryop- 
teris cristata, Primula mistassinica) represented by dis- 
junct localities in northwestern Montana, northern Idaho 
and southeastern British Columbia. 


CAREX PALLESCENS L. (CYPERACEAE)—Ravalli Co., 
Bitterroot Valley along Bass Creek, 46°34'35"N 
114°09'11”W, uncommon in moist meadow bordering Pi- 
nus ponderosa and Pseudotsuga menziesii with Juncus 
balticus, 1020 m, 6 Jul 1997, W. E. Albert 3167 (MICH); 
Bitterroot Valley, app. 18.4 km south and 2.4 km west of 
Florence, 46°28'40"N 114°09’'11”W, uncommon in season- 
ally-saturated meadow along a small stream with Populus 
trichocarpa, Pinus ponderosa, Carex illota and C. lanu- 


[Vol. 49 


ginosa, 1085 m, 11 Jun 2000, W. E. Albert & B. Heidel 
s.n. (MONTU); Bitterroot Valley, south of Hamilton at 
Cory Place, 46°12’05’N 114°10’05’W, along waterway, 
1090 m, 27 Jun 1979, J. Cory 1917 (MONTU). Albert 
3167 determined by B. Heidel, verified by A.A. Reznicek 
(MICH); Cory 1917 determined as Carex torreyi by K. 
Lackschewitz, annotated to C. pallescens by A.A. Rezni- 
cek (MICH). 

Previous knowledge. An eastern species, introduced on 
Hornby Island, British Columbia from Europe or eastern 
North America (G.W. Douglas et al. 1994. Vascular Plants 
of British Columbia, Part 4), and recently discovered in 
Stevens Co., Washington, until recently not otherwise 
known from west of Great Lakes region. 

Significance. First reports for Montana, a range exten- 
sion of approximately 335 km southeast of Stevens Co., 
Washington (Bjork 3463, WS, ID). It is otherwise disjunct 
approximately 1700 km west from Duluth, Minnesota. 


CAREX PRAIREA Dewey (CYPERACEAE)—Flathead 
Co., northern Salish Mountains, confluence of Lime and 
Magnesia Creek drainages, approximately 6.4 km south 
of Trego, 48°38'30"N 114°52'30’W, locally common in 
matted clumps over 8—10 acres of bog birch fen complex 
with Potentilla fruticosa and Carex capillaris, 1060 m, 25 
Jul 1995, F.J. Triepke 24 (USFS Fortine District Herbar- 
ium); Flathead Co., northern Salish Mountains, Magnesia 
Creek drainage, Magnesia Fen, 48°37'30"N 114°52'30’W, 
very common and abundant in fen, with Betula glandu- 
losa, Carex leptalea and Tomentypnum nitens, 1145 m, 21 
Jun 1995, 7. Spribille 3355 (MICH), T. Spribille 3358 
(USFS Fortine District Herbarium); same location, 1 Aug 
1995, T. Spribille 3902 (COLO); Flathead Co., northern 
Salish Mountains, Blessed Creek ca. 0.75 km above con- 
fluence with Sunday Creek, 48°49'40’N 114°32'20’W, in 
rich calcareous fen, local in small patches, 1325 m, T. 
Spribille 7749 (COLO). T. Spribille 3355 verified by A.A. 
Reznicek (MICH). 

Previous knowledge. A widespread species of the boreal 
forest, Carex prairea has been reported from the western 
cordillera in Idaho (R. Davis, 1952, Flora of Idaho) and 
Wyoming (E. Hultén & M. Fries, 1986, Atlas of North 
European Vascular Plants North of the Tropic of Cancer, 
3 vol.), although we have been unable to locate vouchers 
for these reports. In western Canada it is known from 
several sites in the Cariboo-Chilcotin region of central 
British Columbia (A. Roberts, 1983, A Field Guide to the 
Sedges of the Cariboo Forest Region, British Columbia. 
B.C. Min. For., Land Management Rep. No. 14; T-M.C. 
Taylor, 1980, The sedge family [Cyperaceae] of British 
Columbia, Royal B.C. Museum Handb. #43) and from 
central Alberta (J.G. Packer, 1983, Flora of Alberta). 

Significance. First reports for Montana, a range exten- 
sion of ca. 420 km south from the nearest known stations 
in west-central Alberta. 


CAREX VAGINATA Tausch (CYPERACEAE)—Lincoln 
Co., northern Salish Mountains, White Creek, just south 
of Forest Service Road 3529, 48°34’00"N, 114°56'30’W, 
1200 m, in midmontane Picea glauca swamp with Rubus 
pubescens, and Carex disperma, 5 Jun 1996, T. Spribille 
& F.J. Triepke 5054 (USFS Fortine District Herbarium, | 
MICH); Lincoln Co., northern Salish Mountains, fen on 
White Creek along FS Rd. 36, 48°33'45’"N, 114°57'00"W, | 
1125 m, 17 Jul 1998, T. Spribille & R.S. Wirt 7938 | 
(MONTU). Spribille & Triepke 5054 verified by A.A. 
Reznicek (MICH). 

Previous knowledge. A widespread pan-continental spe- ~ 
cies of the boreal forest from Alaska to Labrador (A.E. 


2002] 


Porsild & W.J. Cody, 1980, Vascular Plants of Continental 
Northwest Territories, Canada, Natural Museum of Natu- 
ral Sciences, Ottawa), south to New York, Michigan and 
Minnesota. 

Significance. First reports for Montana and the western 
contiguous United States, a range extension of about 120 
km south from the nearest location mapped by T.M.C. 
Taylor (The sedge family [Cyperaceae] of British Colum- 
bia, Royal B.C. Museum Handb. #43, 1980) in the south- 
ern Rocky Mountain Trench of British Columbia. 


CENTAURIUM ERYTHRAEA Rafin. (GENTIANACEAE)— 
Sanders Co., Cabinet Gorge Reservoir, road between Nox- 
on and Heron on south side of reservoir, ca. 5 km south 
of mouth of Elk Creek, locally common where powerline 
maintenance road clearing connects to the main road. 
48°02'N 115°52'30"W, 670-730 m, 25 Aug 1997, T. Spri- 
bille 7429 (COLO). Verified by W.A. Weber (COLO). 

Previous knowledge. A Eurasian meadow species with 
medicinal uses, reported as established in northwestern 
North America from southern British Columbia (G.W. 
Douglas et al., 1990, Vascular plants of British Columbia, 
Part 2) south to California and inland to Idaho (C.L. 
Hitchcock et al. 1959, Vascular plants of the Pacific 
Northwest, Vol. 4). 

Significance. First report for Montana, representing the 
most inland station in western North America. 


ERIOGONUM VISHERI A. Nels. (POLYGONACEAE)— 
Carter Co., Powderville Road badlands, on the divide be- 
tween Dry Creek and Whitetail Creek, 45°46'18"N 
104°55'42”, occasional on outcrops and outwash flats of 
Hell Creek Formation shale with Allium textile, Musineon 
divaricatum and Elymus lanceolatus, 948 m, 6 Jun 1997, 
B. Heidel 1540 (MONT); same location, 12 Jul 1997, J. 
Vanderhorst 5732 (MONT, MONTU). 

Previous knowledge. A regional endemic of the Great 
Plains, previously known only from North and South Da- 
kota (Great Plains Flora Committee, 1986, Flora of the 
Great Plains). 

Significance. First report for Montana, a range exten- 
sion of approximately 100 km southwest from Slope 
County, North Dakota and 150 km northwest from Har- 
ding Co., South Dakota. 


LESQUERELLA DOUGLASII S. Wats. (BRASSICACEAE)— 
Lincoln Co., Lake Koocanusa, Rexford Bench, 48°54’N 
115°10'30"W, just west of Rexford along path; occurring 
infrequently and in small populations on sand in Pinus 
ponderosa/Purshia tridentata community, 775 m, 14 May 
1999, F.J. Triepke & A. Stachurska 233 (COLO), same 
location, fruiting material, 26 Jun 2000, F.J. Triepke 246 
(USFS Fortine District Herbarium). Triepke 246 verified 
by R. Hartman (RM). 

Previous knowledge. A Columbia Basin species of 
Washington, Oregon and British Columbia, known from 
a string of disjunct populations in the Rocky Mountain 
Trench of southeastern British Columbia. Rollins (1993, 
The Cruciferae of Continental North America) suggested 
it was to be expected in northwestern Montana. 

Significance. First report for Montana, a range exten- 
sion of ca. 40 km from the nearest known location along 
the Elk River near Grasmere, British Columbia (Spribille 
#760, UBC). 


MIMULUS RINGENS L. (SCROPHULARIACEAE)—Cho- 
teau Co., south shore of Missouri River, approximately 8 
km east of Virgelle, 48°02'08”"N 110°09'06” W, seasonally 
flooded sandbar with Populus deltoides seedlings, Eleo- 
charis palustris and Helenium autumnale, 767 m, 7 Jul 


NOTEWORTHY COLLECTIONS 57 


2000, B. Heidel 1952 (MONTU). Verified by R. Meinke 
(OSC). 

Previous knowledge. This primarily eastern species is 
known only from widely scattered western stations in Col- 
orado (W.A. Weber & R.C. Wittman, 1992, Catalog of the 
Colorado Flora: A Biodiversity Baseline), Idaho (R. Da- 
vis, 1952, Flora of Idaho), California (J.C. Hickman, ed. 
1993. The Jepson Manual of the Higher Plants of Cali- 
fornia) and Washington (specimen at WTU). 

Significance. First report for Montana, a range exten- 
sion of at least 745 km west from Rolette Co., North Da- 
kota. 


RIBES LAXIFLORUM Pursh (GROSSULARIACEAE)— 
Lincoln Co., West Cabinet Range, south end of Little Spar 
Lake, 29 km SSW of Troy, 48°12'38”N 116°01'06’W, tall 
shrub field, 1675 m, 8 Sep 1997, E. Pederson 500 (MRC); 
same location, 26 Aug 1998, M. Arvidson & L. Ferguson 
10I5 (MRC). Both specimens verified by PE Stickney 
(MRC). 

Previous knowledge. A Pacific coastal species, known 
inland from scattered stations in the Rocky Mountains of 
British Columbia (G.W. Douglas et al., 1990, Vascular 
Plants of British Columbia, Part 2), southwestern Alberta 
(C.L. Hitchcock & A. Cronquist, 1973, Flora of the Pacific 
Northwest), Colorado and New Mexico (A. Cronquist et 
al. 1997, Intermountain flora, Vol. 3, Part A.). 

Significance. First report for Montana, a range exten- 
sion of 13 km east from the nearest known location, at 
Halverson Creek, Bonner Co., Idaho, 1 km from the Mon- 
tana state line (Pederson 26, MRC). 


SENECIO CONGESTUS (R. Br.) DC. (ASTERACEAE)— 
Roosevelt Co., rangeland, no coll. date, received 11 Jun 
1992, location information unavailable, Roosevelt County 
Extension Service s.n. (MONT). Determined by J. H. Ru- 
mely (MONT). 

Previous knowledge. A pan-continental boreal wetland 
species known from Newfoundland to Alaska and south 
from South Dakota and Iowa (Great Plains Flora Com- 
mittee 1986, Flora of the Great Plains) to Michigan (E.G. 
Voss, 1972—1996, Michigan Flora). 

Significance. First report for Montana, a range exten- 
sion of at least 30 km west from Divide Co., North Da- 
kota. 


VENTENATA DUBIA (Leers) Coss. & Dur. (POACEAE)— 
Ravalli Co., Skalkaho Creek, 46°09'57’N 113°55'54’W, 
common on dry roadsides with Poa compressa, Stipa co- 
mata. 1219 m, Jul 1995, W. E. Albert 3131 (MONT). Ver- 
ified by J. R. Rumely (MONT). 

Previous knowledge. A southern European species of 
dry grasslands introduced in western and northeastern 
North America, known from Idaho and Washington (C. L. 
Hitchcock et al. 1969. Vascular Plants of the Pacific 
Northwest, Vol. 1), southwestern British Columbia (G. W. 
Douglas et al., 1994, Vascular Plants of British Columbia, 
Part 4) and Utah (L. Allen & M. Curto, 1996, Madrono 
43:337—338). 

Significance. First report for Montana, a range exten- 
sion of approximately 120 km east from Idaho Co., Idaho. 


VIOLA SELKIRKII Pursh ex Goldie (VIOLACEAE)—Lin- 
coln Co., Whitefish Range, Grave Creek, on south bank 
of creek ca. 2 km downstream of Williams Creek conflu- 
ence, 48°50’45"N_ 114°49’45”, in alluvial mixed forest of 
Betula papyrifera and conifers, with Aralia nudicaulis and 
Symphoricarpos albus; infrequent, only 30—40 plants 
found; 1030 m, 8 Jun 1999, T. Spribille & A. Stachurska 
9081 (BHO). Verified by H. Ballard Jr. (BHO). 


58 MADRONO 


Previous knowledge. This circumpolar boreal species is 
found in North America primarily in the eastern deciduous 
forests, but is also known only from widely scattered lo- 
calities in the Rocky Mountains south to Colorado (cf. E. 
Hultén, 1968, Flora of Alaska and Neighboring Territo- 
ries) and New Mexico (W.C. Martin & C.R. Hutchins, 
1980, Flora of New Mexico). 

Significance. First report for Montana, a range exten- 
sion from southeastern British Columbia. 


—Tosy SPRIBILLE, Kootenai National Forest, Fortine 
Ranger District, RO. Box 116, Fortine, MT 59918 (current 
address: Herbarium, Department of Systematic Botany, 
Albrecht von Haller Institute of Plant Sciences, University 
of Gottingen, Untere Karsptile 2, D-37073 Gottingen, Ger- 
many; e-mail toby.spribille@gmx.de); BONNIE HEIDEL, 
Montana Natural Heritage Program, 1515 E 6th Ave., He- 
lena, MT 59620 (current address: Wyoming Natural Di- 
versity Database, University of Wyoming, P.O. Box 3381, 
Laramie, WY 82071-3381, e-mail bheidel@uwyo.edu); 
WALLACE E. ALBERT, 3653% Silverthorn Drive, Stevens- 
ville, MT 59870; E JAcK TRIEPKE, Kootenai National For- 
est, Fortine Ranger District, PO. Box 116, Fortine, MT 
59918; Jim VANDERHORST, Natural Heritage Program, 
West Virginia Division of Natural Resources, P.O. Box 67, 
Ward Rd. Elkins, WV 26241-0067; and G. MICHAEL AR- 
VIDSON, Kootenai National Forest, Three Rivers Ranger 
District, 1437 Hwy 2 N, Troy, MT 59935. 


OREGON 


HIERACIUM CAESPITOSUM Dumort. (ASTERACEAE).— 
Wallowa Co., along Bear Creek Road, ca. 2 km S of Wal- 
lowa, with Dactylis glomerata, Phleum pratense, and 


[Vol. 49 


Pseudotsuga menziesii. Also in pastures, along logging 
trails but absent in adjacent undisturbed forest, and along 
roads and riparian areas bordering Bear Creek, elev. 1090 
m, T1S R42E sect. 3, Long. 117.55, Lat. 45.48, 23 July 
2000, Brooks (OCS #197099); T1S R42E sect. 15, 17 July 
2001, Dwire 1728 (OSC) (Verified by K. L. Chambers, 
OSC). Distribution extends south along the Bear Creek 
Trail into the Eagle Cap Wilderness, and north in the ri- 
parian areas bordering the Wallowa River. 

Previous knowledge. This species is also known as 
Hieracium pratense Tausch, an outdated synonym ap- 
pearing in Hitchcock and Cronquist (1973) and other 
western floras. Native to Eurasia, meadow hawkweed was 
probably introduced into the United States in the 1820's, 
and was first reported in the Pacific Northwest in Pend 
Orielle Co., Washington in 1969. It has become wide- 
spread throughout Washington, northern Idaho, and north- 
western Montana (Wilson et al. Rangelands 19:18—23, 
1997; Toney et al., Northwest Science 72:198—209, 1998). 
It is spreading rapidly, primarily in montane meadows, 
pastures, and disturbed areas along roads and hillsides. 
Meadow hawkweed is a tenacious invader, and is listed 
as a noxious weed in Washington (Class B), Idaho, and 
Montana (Category 2). 

Significance. First report of the species for Oregon. Al- 
though present in the Bear Creek drainage, Wallowa Co. 
for perhaps 10 years, meadow hawkweed was only re- 
cently distinguished from native hawkweeds. An addition- 
al unvouchered population of Hieracium caespitosum has 
been reported from Hood River Co., Oregon. 


—KATHLEEN A. DwirE, Department of Forest Science, 
Oregon State University, Corvallis, OR 97331-5752, and 
CATHERINE G. PARKS, USDA Forest Service, Pacific 
Northwest Research Station, 
Grande, Oregon 97850. 


1401 Gekeler Lane, La i 


MADRONO, Vol. 49, No. 1, p. 59, 2002 


ERRATUM 


In Volume 48, No. 2, there was a typographical error in the title of the paper by Dieter 
Wilken (pages 116—122). The correct title should be 


A new Ipomopsis (Polemoniaceae) from the southwest USA and adjacent Mexico. 


Volume 49, Number 1, pages 1—60, published 14 August 2002 


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— 


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PG VOLUME 49, NUMBER 2 APRIL-JUNE 2002 


IND 


CATALOGUE OF NoN-NATIVE VASCULAR PLANTS OCCURRING SPONTANEOUSLY IN 

CALIFORNIA BEYOND THOSE ADDRESSED IN THE JEPSON MANUAL—Parrt I 

Fred Hrusa, Barbara Ertter, Andrew Sanders, Gordon Leppig 

FURL ELIE LY TIL SAE RG ROSS COOP 0 Nee ok NN oe A ne 61 
FIELD ASSESSMENT OF THE CALIFORNIA GAP ANALYSIS PROGRAM GIS DaTABASE IN 

CENTRAL CALIFORNIA 

John F. Karlik, Eugene D. Albertson, Y. Jae Chung, Alistair H. McKay 

TRA ERAL EE OTS CNY LEW EDV oe ene er ea eek”. Se ne mn eee 99 
SOME Factors INFLUENCING SEEDLING DENSITY OF CALIFORNIA BLACK OAK 

(QUERCUS KELLOGGII) IN THE CENTRAL SIERRA NEVADA, CALIFORNIA 

Barrett A. Garrison, Robin L. Wachs, James S. Jones 

GRE NIG CW Ti. TTI ES oie ca costes nates OE PR gh vino n 0 deg POD ED cs vesnestn sessions 115 
Canopy MACROLICHENS FROM Four FoREST STANDS IN THE SOUTHERN SIERRA 

MIXED CONIFER FORESTS OF SEQUOIA/KINGS CANYON NATIONAL PARK 

David C. Shaw and St@RCHASNCKET ek Bo Soe. pe NL) ov nccc ese sescnccnsensonss 122 


CALYSTEGIA SILVATICA (CONVULVULACEAE) IN WESTERN NORTH AMERICA 


R. K. Bromine... 4 a IE... eS he. 130 

PNRUZIOIN oo e oo oo acc NT BF os ESN. cscs duacsnnccsececcaes 132 

$ WASHINGTON 2 onc A ESS NS on SSE Sa ROSSA soa vecceesesesaccosess 132 
WIGOREA a DF A cn WSS SSA osc cckeccccnneceesee 132 


SEEING THINGS WHOLE: THE ESSENTIAL JOHN WESLEY POWELL, EDITED BY 
WILLIAM DEBuys 
Wall terran Te ed oon ic oo once cs RUNS Sev ose sesc con ecesweccevene 134 
INVENTORY OF RARE AND ENDANGERED PLANTS OF CALIFORNIA, BY THE CALIFORNIA 
NATIVE PLANT Society, Davip P. TIBOR, CONVENING EDITOR 
ENTE OAD FIG 2 2) DD 12, Re me ets Oy |S A rn eee 135 


PAO 7 US) SCHEDULE EON SPEARGRS 652i ccs coccscccean eceicacevadacccesavitaulecowvinvessdeuecwnndece 136 


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This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper). 


MApRONO, Vol. 49, No. 2, pp. 61-98, 2002 


CATALOGUE OF NON-NATIVE VASCULAR PLANTS OCCURRING 
SPONTANEOUSLY IN CALIFORNIA BEYOND THOSE ADDRESSED IN 
THE JEPSON MANUAL—PART I 


FRED HRUSA 
California Department of Food and Agriculture, Plant Pest Diagnostics Center, 
3294 Meadowview Rd., Sacramento, CA 95832-1448 
FHrusa @cdfa.ca.gov 


BARBARA ERTTER 
University and Jepson Herbaria, University of California, 
Berkeley, CA 94720-2465 


ANDREW SANDERS 
Botany and Plant Sciences Department, University of California, 
Riverside, CA 92521-0124 


GORDON LEPPIG! 
Biological Sciences Department, Humboldt State University, Arcata, CA 95521 


ELLEN DEAN 
UC Davis Herbarium, Section of Plant Biology, University of California, 
Davis, CA 95616 


ABSTRACT 


A catalogue of 315 non-native vascular plant taxa documented as occurring spontaneously in California 
beyond those addressed in The Jepson Manual: Higher Plants of California is presented. The catalogue 
was compiled from new collections by the authors and others, previously existing herbarium specimens, 
formal publications, other printed reports, and direct communications with field botanists. Only reports 
backed by herbarium vouchers are accepted as adequately documented. Of the 315 species, 58 are fully 
or sparingly naturalized in relatively undisturbed wildland habitats, 53 are naturalized in disturbed areas, 
34 are tenuously established or locally persisting, 13 are non-escaped weeds of greenhouse or similarly 
cultivated environments, 43 are presumed to be non-persisting casuals (waifs), for 110 there is no current 
information or observations available, and 4 have likely been extirpated. In addition, 13 reported taxa are 
here specifically excluded as based on erroneous information. Taxa highlighted as already being fully 
naturalized or potential pests are Amaranthus rudis, Brassica fruticulosa, Boehmeria cylindrica, Calys- 
tegia silvatica subsp. disjuncta, Cabomba caroliniana, Cotoneaster lacteus, Crataegus monogyna, Dit- 
trichia graveolens, Fumaria capreolata, Geranium purpureum, Geranium rotundifolium, Hedera canar- 
iensis, Limnobium laevigatum, Maytenus boaria, Pyracantha crenatoserrata, Salvinia molesta, Trifolium 


tomentosum, and Verbascum olympicum. 


Key Words: Weeds, non-native, invasive plants, pest plants, voucher specimens. 


The significance of invasive non-native plants 
has recently gained prominence, as evidence 
mounts for both the environmental and economic 
devastation such invasions can cause. A recent is- 
sue of BioScience (51[2] Feb. 2001), for example, 
is devoted to the topic of ““Global Movements of 
Invasive Plants and Fungi.”’ On the national level, 
various legislative initiatives have been proposed to 
address the problem, such as the Harmful Nonna- 
tive Weed Control Act (S. 198). At the local level, 
Weed Management Areas, established through the 


' Present address: California Dept. of Fish and Game, 
Coastal Timberland Planning Program, 619 Second Street, 
Eureka, CA 95501. 


coordinated efforts of public and private agencies 
and organizations, now blanket most of California. 

Obviously, for all of these efforts to work prop- 
erly, accurate and comprehensive information 
needs to exist on which non-native plants occur 
within the area of concern and what potential level 
of threat they represent. While the average citizen 
might assume that this information is readily avail- 
able, especially in this age of electronic databases, 
the reality is unfortunately otherwise. This is pri- 
marily because, although a broad spectrum of pro- 
fessional biologists and amateur enthusiasts eagerly 
hunt down and keep track of rare native species, 
non-natives have been historically under-reported if 
not outright ignored. As a result, our existing 
knowledge of the identity, occurrence, frequency, 


62 MADRONO 


and distribution of non-native plants is often 
sketchy or preliminary. 

California is by no means an exception to this 
rule, and the incomplete coverage of non-native 
species was one of the acknowledged short-com- 
ings of The Jepson Manual: Higher Plants of Cal- 
ifornia (Hickman 1993), which otherwise repre- 
sented the state-of-the-art coverage of California 
plants at the time of its publication. In a subsequent 
statistical analysis of the numbers and distribution 
of the non-native taxa reported in the Manual (Rej- 
manek and Randall 1994), mention was made of 
some clearly naturalized plants that were missing, 
but no compilation was attempted. As a precursor 
tally by Hrusa and Ertter, over 70 non-native plant 
taxa beyond those included in the Manual were not- 
ed as naturalized in California (Ertter 2000). More- 
over, the composition, frequency and distribution of 
plants in a given area is not static; this is particu- 
larly a feature of the non-native component, but 
even for native plants frequent updates to floristic 
accounts are necessary to maintain currentness 
(Yatskievych and Raveill 2001). The following cat- 
alogue is presented as the first installment of a con- 
tinuing comprehensive effort to follow through on 
this preliminary note by compiling existing reports 
and documenting new occurrences of plant taxa 
that occur spontaneously in California but which 
are not treated in The Jepson Manual. Such cata- 
loguing is intended to serve several functions: To 
help field botanists and consultants identify un- 
known species they come across during survey ac- 
tivities; to assist in the recognition of introduction 
pathways; and finally to provide further raw mate- 
rial for study of the invasion process itself. 


MATERIALS AND METHODS 


Data sources. Records of non-native vascular 
plants reported as growing without cultivation in 
California were reviewed and compiled. Data 
sources consisted of herbarium specimens, formal 
publications, other printed reports, and verbal com- 
munications with field botanists. Primary were our 
own and others’ plant collections; secondary were 
the numerous published local California floras and 
verbal reports provided to one or more of the au- 
thors. We have also included detailed records as 
available for most of those taxa mentioned by Rej- 
manek and Randall (1994) as absent from The Jep- 
son Manual. Because of the inevitable potential for 
misidentification, and following standard taxonom- 
ic practice, only reports that were backed up by 
hard documentation in the form of voucher speci- 
mens deposited in a publicly accessible herbarium 
are included in the current catalogue; this includes 
both those observed by one of the authors, stated 
in publication that such a voucher does exist, or 
verified by herbarium staff that one does exist. For 
example, the following taxa listed in Rejmanek and 
Randall (1994) are not included in Part I of the 


[Vol. 49 


current catalog because testifying vouchers could 
not be located: Cordyline australis (Forst.f.) Endl. 
(Liliaceae s.l.), Cutandia memphitica (Spreng.) 
Richt (Poaceae) and Dodonaea viscosa Jacq. (Sap- 
indaceae). Of these taxa, a Cordyline has been ob- 
served by Hrusa at Salt Point, Sonoma Co., but a 
voucher confirming the specific application has not 
yet been acquired. The other two also may be es- 
tablished in California, but neither have as yet been 
so verified by the deposition of annotated speci- 
mens. 

Many of the printed or verbal reports were from 
federal and state agency botanists, native plant so- 
ciety members, weed control groups, and other un- 
published sources. Of particular importance were 
the plant samples submitted for identification by 
land managers, owners, farmers, ranchers, et al. to 
one or more herbaria. From these sources exact col- 
lection locality and ecological situation were some- 
times difficult to ascertain, and the material provid- 
ed was sometimes too poorly preserved to allow 
for more than an equivocal determination. The re- 
sult was that, until field confirmation or better col- 
lections can be obtained, some reports currently re- 
main unverified and are thus excluded from this 
compilation. Nevertheless, the value of the coop- 
eration of landowners, land managers, or other field 
people cannot be overstated. 

At the same time, the resultant catalogue is more 
than a simple report compilation. All reports were 
subject to verification by one or more authors, who 
also had the responsibility of determining natural- 
ization status. Correspondence with and the assis- 
tance of staff at herbaria worldwide were needed in 
the effort to track down documenting specimens for 
some taxa reported for California. Although one 
cannot absolutely prove the absence of a specimen, 
by a preponderance of evidence some putative re- 
cords were excluded, including several in published 
sources. For those taxa where a specimen was 
available, extensive identification efforts were often 
required to confirm identity, involving both world- 
wide taxonomic literature and herbarium speci- 
mens. In some particularly difficult cases, voucher 
specimens were sent to the appropriate specialist 
for identification or verification. Other problems, 
both taxonomic and nomenclatural, were resolved 
by electronic correspondence with the appropriate 
specialist, including several in Eurasia. Taxa re- 
ported for California, but for which documenting 
vouchers appear to be absent, and others deter- 
mined on examination to be misidentified or erro- 
neously reported, are listed and discussed in the 
RESULTS section. 


Scope of inclusion. Our decision of what to in- 
clude in the catalogue was based on three criteria: 
1) The plant occurs spontaneously in California. 
‘‘Spontaneous”’ as used here encompasses not only 
fully naturalized populations but any evidence of 
successful reproduction or initial appearance inde- 


2002] 


pendent of targeted cultivation, including casuals, 
garden escapes, and weeds of greenhouse, nursery, 
garden, lawn, and agricultural field. This exception- 
ally broad definition was chosen to capture the pos- 
sible first appearance of potentially invasive spe- 
cies. However, long-persisting individuals that were 
intentionally planted, such as trees near abandoned 
homesites are not included unless they show evi- 
dence of successful reproduction, either sexual or 
vegetative; 2) The species was not addressed in The 
Jepson Manual, not even as an equivocal mention 
subordinate to another taxon. For example, Mentha 
spicata L. var. longifolia L. (Lamiaceae) is not in- 
cluded in the catalogue because it is mentioned in 
the Manual under M. spicata var. spicata, even 
though the entry is somewhat ambiguous and the 
variety is now fully documented as being natural- 
ized in California. Likewise, we do not report new 
regional occurrences for species already included in 
the Manual (e.g., plants reported only for northern 
California that are documented from the southern 
part of the state as well); 3) At least one reported 
locality is documented by a voucher specimen de- 
posited in a publicly accessible herbarium. Among 
the authors, at least fifteen vouchers were specifi- 
cally prepared for reports that had not previously 
been so documented. It should be recognized that 
a statement in print that a voucher was made does 
not guarantee that one was indeed deposited and it 
is possible that some taxa included in this catalogue 
will eventually, after further research, be found to 
be unverified. In addition, specimens proving oc- 
currence, but held in private collections were not 
included until a specimen or duplicates were made 
available for public scrutiny. 


Catalogue format. Individual taxon records in 
the catalogue are formatted as follows: 

TAXON NAME: Scientific name and author in 
accordance with the most recently available refer- 
ences (e.g., International Plant Names Index [http:// 
www.ipni.org/]; Flora of North America North of 
Mexico; Flora Europaea on-line [http://www.rbge. 
org.uk/forms/fe.html]; Catalogue of New World 
Grasses _ [http://mobot.mobot.org/W3T/Search/nwgc. 
html]). In cases of conflicting recent treatments, final 
decisions were based on our collective judgement. 

DISTRIBUTION (D): Reported occurrences in 
geographic subdivisions of California as described 
in The Jepson Manual, arranged alphabetically. 

CURRENT STATUS (CS): A brief description 
of current status as can best be determined. There 
is disagreement among invasion ecologists as to the 
scope of the term “‘naturalized’’ (Richardson et al. 
2000). We follow the terminology of Richardson et 
al. but split their “‘naturalized’’ into subcategories: 
1, “‘naturalized in wildlands’’, and 2, “‘naturalized 
(outside of wildlands)’’. It is recognized that dif- 
ferential occupation of these habitats is not mutu- 
ally exclusive, that assignment to one or the other 
category may be subjective, and that it is in addi- 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 63 


tion a rare situation that, in California, has not been 
*“‘disturbed”” at some time in the past. We neverthe- 
less find the additional information provided in a 
classification distinguishing “‘naturalized in wild- 
lands’ and “naturalized (outside of wildlands)”’ 
potentially useful. This is because we feel that 
weedy species with a propensity to invade relative- 
ly natural or undisturbed areas are of different con- 
cern and may exhibit different ecological or repro- 
ductive strategies than are those plants which, al- 
though perhaps well-established, are restricted to 
agricultural conditions, roadsides or other heavily 
disturbed sites. The category ““NCI” (no current 
information) needs some explanation. In general 
these are plants confirmed to have occurred at one 
time in California, but for which no immediate in- 
formation is available as to their continued pres- 
ence, rate of spread, or naturalization. Many re- 
cords placed here could be assigned to another cat- 
egory based on the label data, but we preferred to 
maintain a conservative position on records that 
were often more than thirty years old, that is, these 
plants may be gone, or have spread widely during 
the ensuing years and we would be amiss in imply- 
ing more knowledge about their current status than 
we actually have. 

Naturalization categories are described and stan- 
dardized abbreviations, as used in the catalogue, are 
introduced and described in Table 1. 

DOCUMENTATION (DOC): Reports are ar- 
ranged alphabetically by county (in bold) as either 
a literature citation for previously published reports, 
or as specimen citation for newly documented oc- 
currences. Duplicate specimens might be housed in 
additional herbaria beyond those cited. Herbarium 
abbreviations are as used in /ndex Herbariorum 
(http://www.nybg.org/bsci/ih/ih.html). Generally 
literature cited as documentation includes specimen 
citations; however, collection or accession numbers, 
collectors names, dates and other details of location 
and habitat are only occasionally available in these 
sources. We expect that published floristic compi- 
lations maintain professional standards, and for 
published records where the specimen location is 
not specifically cited, the institution with which the 
author is or was affiliated houses the majority of 
the documented material. Known misapplications 
or misidentifications based on cited literature sourc- 
es alone should be brought to our attention. 

NOTES: Additional discussion or explanation as 
required or available, such as nomenclatural prob- 
lems, history of introduction or evidence of inva- 
siveness. In addition, for taxa believed to be be- 
coming common or rapidly expanding their ranges, 
a short comparison among similar species and/or 
identification clues are provided. 


RESULTS 


A total of 315 non-native vascular plant species 
meeting the criteria for inclusion are listed in Ap- 


64 MADRONO [Vol. 49 
TABLE 1. CURRENT STATUS CATEGORIES. 
Catalogue 
abbrev. Description 
NW Naturalized in wildlands: A population that appears to be successfully propagating (sexual- 


ly or vegetatively) and maintaining itself in a wildland situation. 
N Naturalized (outside of wildlands): A non-wildland population that is apparently of suffi- 
cient size or distribution to make the chances of its disappearance due to stochastic fluc- 


tuation negligible. 


Persistence tenuous: A wildland or non-wildland population that is of few enough individ- 


uals to make the chance of disappearance via stochastic fluctuation a possibility. 
C Casual: Synonymous with “‘waif”’ as used in The Jepson Manual to describe a spontane- 
ous occurrence that shows no evidence of successful propagation and spread. 


Greenhouse, nursery, garden weed: Non-escaped weed of greenhouse, nursery, garden, or 


No current information: Plants confirmed to have occurred at one time in California, but 


for which there is no immediate information available as to the continued presence, rate 
of spread, or naturalization. Many records placed here are assignable to another catego- 
ry based on the label data, but we prefer not to categorize these without more current 


GH/C 

other highly cultivated environment. 
NCI 

information. 
al 


Extirpated: Or reported as such. Habitat alteration or eradication programs in place have 


either eliminated these taxa from California or have nearly done so. 


pendix 1 and itemized in detail (Appendix 2). Both 
lists are organized as in The Jepson Manual, with 
Appendix 1 intended to facilitate the location of 
names in Appendix 2. The classification used in the 
catalogue to convey current naturalization status is 
summarized in Table 2. If a taxon had populations 
that were classifiable in different categories, the 
taxon as a whole was included in the category in- 
dicating more complete naturalization. For exam- 
ple, Salvinia molesta D.S. Mitch. (Salviniaceae) has 
several populations well-naturalized in both wild- 
land (NW) and non-wildland sites (N), plus is 
known several relatively small, localized popula- 
tions from private ponds (classed as TEN); it was 
thus classified as naturalized (NW) for the sum- 
mary Statistics. 

A subset of included species have the potential 
to achieve or have already achieved significant nat- 
uralized distributions or are pest plants, either as 
rapidly spreading invasive species (e.g., Dittrichia 
graveolens (L.) Greuter, Asteraceae) or as first Cal- 
ifornia appearances of well-known pest plants that 
were immediately targeted for eradication (e.g., 
Salvinia molesta). These were taxa rapidly expand- 
ing their former ranges, serious invaders in other 


TABLE 2. CURRENT STATUS SUMMARY. Explanation and 
definition of individual categories are provided in Table 1. 


Catalogue 
Definition abbrev. ‘Total 
Naturalized in wildlands NW 58 


Naturalized (outside of wildlands) N 5)3} 


Tenuous/locally persisting TEN 34 
Greenhouse, nursery, garden weed GH/C 13 
Casual (Cc 43 
No current information NCI 110 
Extirpated EXT 4 


parts of the world, plants already widespread and 
relatively commonly encountered, or plants appar- 
ently widespread but not heretofore correctly iden- 
tified [e.g., Hedera helix in part = Hedera canar- 
iensis Willd., (Araliaceae)]. The plants in this sub- 
set are listed in Table 3. They are provided a longer 
discussion and identification clues within their in- 
dividual Catalogue entries. 


Excluded taxa. It is important to note also the 
number of putative reports that were excluded from 
the current catalogue not from lack of documenta- 
tion, but because the reports were determined by 
the authors to be based on erroneous information. 
The importance of accurate identification of non- 
native naturalizing taxa cannot be overstated, there 
are numerous examples to show how control was 
only achieved once the proper identity of a pest was 
determined. A notable example is Salvinia molesta 
(Thomas and Room 1986); a similar, but currently 
unresolved example from California involves the 
genus Salsola (Chenopodiaceae) (M. Pitcairn per- 
sonal communication). Due to the taxonomic and 
nomenclatural complexities of name application to 
non-native taxa, the following list of thirteen (13) 
excluded reports is certainly not exhaustive: 

Achnatherum clandestinum (Hack.) Barkw. (Po- 
aceae): Misapplied in California to the similar A. 
brachychaetum (Godr.) Barkw. Plants from Mon- 
terey Co. sometimes identified as A. clandestinum 
have been recently redetermined by Barkworth 
(UTC) as A. brachychaetum. The identity of a spec- 
imen reported as this (Barkworth 1993), from San 
Luis Obispo Co., remains unverified, but a possible 
duplicate (or second collection from the same site 
by the same collector) at UC was annotated by 
Barkworth as A. caudatum (Trin.) Jacobs, a taxon 
highly similar to and perhaps conspecific with A. 
brachychaetum (see Vickery et al. 1986) suggesting 


2002] 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 65 


TABLE 3. POTENTIALLY SIGNIFICANT PLANTS. Fully naturalized species or first California appearances of well-known 


pest plants. 


Taxon 


Amaranthus rudis 
Brassica fruticulosa 
Boehmeria cylindrica 


Calystegia silvatica subsp. disjuncta 


Cabomba caroliniana 


Cotoneaster lacteus 
Crataegus monogyna 
Dittrichia graveolens 
Fumaria capreolata 


Geranium purpureum 
Geranium rotundifolium 
Hedera canariensis 


Limnobium laevigatum 


Maytenus boaria 
Pyracantha crenatoserrata 


Salvinia molesta 


Trifolium tomentosum 
Verbascum olympicum 


Summary of current plant behavior and status 


Well-established in widely separated localities. 

Rapidly spreading in mostly disturbed areas. 

Becoming common in the upper Sacramento River delta. 

Widespread in coastal areas, previously misidentified. 

Rapidly expanding, well-established aquatic invader. Dominant in some 
Sacramento River delta locations. 

Becoming locally common, expected more widely. 

Appearing in widely separated regions, often in stable habitats. 

Rapidly expanding. 

Unrecognized, probably more common than previously thought, likely 
expanding. 

Rapidly becoming more abundant. 

Widespread; overlooked previously, or rapidly expanding. 

Persistent invader of forest and woodland understories; previously con- 
fused with or included in H. helix. 

Rapid colonizer of open, still water, at low elevations. Can dominate to 
the exclusion of all other aquatic species. 

Aggressive invader of burns, forming long-lasting seedbanks. 

Widespread; invader of mesic ditchbanks, roadside depressions, open 
shorelines. 

Recent aquatic introduction into California where spreading in the Col- 
orado River and canal system. 

Widespread; previously overlooked or misidentified. 

Naturalized in wildland habitat, expanding for more than 20 years. 


that it may also be that taxon. Relationships among 
A. clandestinum, A. caudatum and A. brachychae- 
tum need critical review. 

Dichondra repens Forst. & Forst.f. (Convolvu- 
laceae): Misapplied in California. Acc. to Hortus 
III (1976) material cultivated as D. repens is actu- 
ally D. micrantha. See comments in the catalogue 
under the latter taxon. 

Glyceria declinata Bréb. (Poaceae): Acc. to Lep- 
pig, all specimens of putative Glyceria declinata 
from California are indistinguishable from G. oc- 
cidentalis. A discussion and treatment of variation 
in California Glyceria occidentalis is currently in 
preparation by Leppig. See also comments by Soza, 
Provance and Sanders (2000). 

Humulus americanus Nutt. sensu lato (Canna- 
baceae): The voucher for the report in Smith (1973, 
pg. 92), has not been located, but a new collection 
from the same locality (Echo Summit Lodge, El 
Dorado Co.) by Steve Matson has been verified by 
E. Small as H. lupulus var. neomexicanus A. Nelson 
& Cockerell. In Small’s treatment of Humulus in 
Flora of North America North of Mexico (3: 384— 
387, 1997), H. americanus has been split among 
several varieties of H. lupulus native to North 
America: var. lupuloides E. Small, var. neomexi- 
canus, and var. pubescens E. Small. Although 
Small’s treatment reports var. neomexicanus from 
California, this apparent introduction is the only 
currently confirmed occurrence of the taxon in Cal- 
ifornia. We are rejecting the presence of H. amer- 
icanus S.\. because this name also applies to forms 


of H. lupulus (vars. pubescens and lupuloides) not 
in California. 

Hydrocharis morsus-ranae L. (Hydrocharita- 
ceae): Reports have all been misidentifications of, 
or misapplications to, Limnobium laevigatum. Both 
of these taxa are often called “‘frogbit’’, probably 
the source of the identity confusion. H. morsus-ran- 
dae is a possible threat to northern and high eleva- 
tion lakes in California, but has not yet been con- 
firmed in California outside of cultivation. 

Nothoscordum bivalve (L.) Britton (Liliaceae 
sensu lato): Apparently misapplied to Nothoscor- 
dum gracile (Ait.) Stearn s.l. (¢ncluding N. borbon- 
icum Kuntze = N. inodorum in the Jepson Manual). 
The cited location in Beauchamp (1986), “‘garden 
weed in Fallbrook”’ has a corresponding specimen 
at CDA of N. inodorum sensu auct. (=N. gracile 
or N. borbonicum). 

Passiflora mollissima (Kunth) L.H. Bailey (Pas- 
sifloraceae): Acc. to determinations by D. Goldman 
(BH) this name has been misapplied to P. tarmi- 
niana Coppens & Barney and P. mixta L.f., both 
of which are established in California. See 
d’Eckenbrugge et al. (2001). 

Polypogon elongatus Kunth (Poaceae): Misap- 
plied to P. imberbis (Phil.) Bjorkm. See catalogue 
for more information. 

Pueraria lobata (Willd.) Ohwi (Fabaceae): Ver- 
bal reports of kudzu vine in California have all 
been traced to native plants of Vitis californica. 
Pueraria has been cultivated in California as a cu- 
riosity, but in general, does not thrive under our dry 


66 MADRONO 


[Vol. 49 


TABLE 4. TIMING OF REPORTING FOR NEW CALIFORNIA RECORDS OF TAXA INCLUDED IN THE CURRENT CATALOGUE IN 


RELATION TO PUBLICATION OF THE JEPSON MANUAL. 


No. of taxa 


Relationship to The Jepson Manual 


129 Discovered prior to the Manual but either not identified or a report not published. 
110 Discovered (published report or not) subsequent to the Manual. 
63 Discovered and a report published prior to the Manual. 
11 Re-determination of pre-Manual species. 
2 Accidentally excluded from the Manual, but the taxonomic treatment was completed 


and submitted prior to publication. 


Mediterranean climate. Locations discovered in 
northern Oregon in 2000 do apparently flower and 
fruit and are currently under pest evaluation. 

Salvia pratensis L. sensu stricto (Lamiaceae): 
Thomas C. Fuller collections from the region of 
Yreka (Siskiyou Co.) cited in Munz, PA. (1968, pg. 
103) as S. pratensis have been redetermined at KW 
as Salvia virgata Jacq. See entry in the catalogue 
under that species for details. 

Sieglingia decumbens (L.) Bernh. (Poaceae): 
Misapplied in Matthews (1997) to Tribolium obli- 
terum (Hemzl.) Renvoize, first reported for North 
America in this catalogue. Cited also in Hitchcock 
and Chase (1950) as “escaped from cultivation, 
Berkeley”’, but no confirming evidence or speci- 
mens have been located. 

Spartina maritima (M.A. Curtis) Fern. (Po- 
aceae): Misapplied to Spartina anglica C.E. Hubb. 
For detailed explanation see Spicher and Josselyn 
(1985). 

Spartina spartinae (Trin.) Merr. (Poaceae): Mis- 
applied to S. densiflora Brongn. Reported from 
Humboldt Bay (Gerish 1979; Spicher and Josselyn 
1985) and in Marin Co. at Greenwood Cove near 
Tiburon [Aug. 11, 1982, M.P. Wells s.n. (CAS, 
CDA)]. 

As a further complication, individual occurrence 
reports can be erroneous even if the plant itself is 
documented to occur spontaneously in California. 
As a particularly convoluted example, Amaranthus 
rudis (Amaranthaceae), which is included in the 
current catalogue on the basis of multiple docu- 
mented reports, had previously been cited in print 
(Munz 1974) only on the basis of a specimen from 
Pala in San Diego County, as A. tamariscinus Nutt. 
(misapplied to A. rudis). As it turns out, a duplicate 
of the voucher specimen (Townsend s.n., housed at 
CDA) has been re-determined as A. palmeri S. 
Wats. In other words, the report of A. rudis from 
near Pala in southern California is erroneous, even 
though other, mostly more recent reports substan- 
tiate the existence of this midwestern native in 
northern California. This example likewise under- 
scores the critical need to have available and ex- 
amine vouchers to confirm their identification. 


DISCUSSION 


Relation to the Jepson Manual. The Jepson Man- 
ual: Higher Plants of California (Hickman 1993) 


has for the past nine years provided California bot- 
anists with a new “‘standard”’ reference, supplant- 
ing the more than thirty-year old A California Flora 
(and its 1968 Supplement) by P.A. Munz and D.D. 
Keck. As earlier noted, non-native plants have his- 
torically been under-collected. Because floristic 
works such as the Manual used herbarium collec- 
tions as the fundamental scientific standard, the re- 
sult has been a general under-representation of non- 
native species. It is hoped the current paper, along 
with the burgeoning interest in non-native taxa as 
potential pest plants, will reverse this tendency. 

In addition to those species that were omitted 
from the Manual because of the lack of accessible 
herbarium documentation, others were not included 
for a variety of reasons. In particular, the Jepson 
Manual Project deliberately excluded ‘casual’ 
(‘waif’) species, unconfirmed naturalization re- 
ports, or material apparently persistent from culti- 
vation. While this was a reasonable decision for the 
Manual, we believe that even these marginal oc- 
currences are worth reporting, given that all taxa 
that are now naturalized and well-established com- 
ponents of the State flora began as rare casuals or 
as small populations. Finally, there are plants in the 
current catalogue that have appeared as spontane- 
ous occurrences in California subsequent to the clo- 
sure of treatments for the Manual. 

Worth mentioning in particular are those natu- 
ralized taxa that were reported in A California Flo- 
ra or adequately documented in the intervening pe- 
riod but which were not treated in the Manual. The 
reasons these taxa were excluded are diverse. Some 
had been included in A California Flora but current 
information as to their naturalization status was not 
available. For others, individual authors may have 
decided that the inclusion of marginally established 
or rarely encountered weeds unnecessarily compli- 
cated identification keys and/or in general did not 
benefit the majority of users. Again, while this ra- 
tionale may have been justified for the purposes of 
the Manual, this excluded group of plants is in- 
cluded in the current catalogue in order to assist 
those who have diverse interests concerning inva- 
Sive taxa. 

The temporal relationship of species in the cur- 
rent catalogue to The Jepson Manual is summa- 
rized in Table 4. 


2002] 


Significance of new reports. The often pestifer- 
ous nature of the most prominent weeds in Cali- 
fornia is well-recognized by the layman and pro- 
fessional alike (e.g., Bossard et al. 2000) What is 
not so universally understood is that these wide- 
spread and/or noxious pests generally began their 
occupation of California as seemingly innocuous 
roadside waifs, occasional garden volunteers, minor 
seed source contaminants, or localized populations 
remaining for some time in other non-prominent 
situations. The California Dept. of Food and Agri- 
culture (CDFA) rates noxious weeds, and applies 
control efforts accordingly, not by their current af- 
fect on agricultural economics, but by their poten- 
tial effect, the position being that control is both 
most biologically and cost effective when the pop- 
ulations are small, before the plant has become a 
widespread, established pest. Data have been as- 
sembled and published in support of this position 
(Bayer 1999). Although CDFA is concerned pri- 
marily with weeds affecting agricultural endeavors, 
wildland pests are no different nor are these con- 
ditions mutually exclusive; it is important to ag- 
gressively control these organisms before they be- 
come too widespread for control to be effective or 
eradication possible. Addressing only the most 
widespread and well-established weeds in Califor- 
nia leaves a gap in our knowledge that may affect 
the future of California’s agricultural activities, 
pristine wildlands and outdoor recreation areas. 
That this is an important gap may be recognized 
when one considers that every presently wide- 
spread weed in California began long ago as a “‘ca- 
sual”? [Centaurea solstitialis L. (Asteraceae), Lep- 
idium latifolium L. (Brassicaceae)], or ““occasional 
garden escape” [Cyftisus scoparius (L.) Link, Gen- 
ista monspessulana (L.) L. Johnson (both Faba- 
Geae)p 

While it cannot be specifically claimed that any 
plant appearing in this catalogue will ultimately be- 
come more than even a local pest, the naturalization 
of non-native plants may ultimately have long-last- 
ing impacts. The truth of this statement is dramat- 
ically evident when perusing S. B. Parish’s 1920 
synopsis of immigrant plants of southern Califor- 
nia, in the number of species that were merely pre- 
sent as scattered waifs in 1920 but which are now 
both fully naturalized and widespread. For exam- 
ple, Potamogeton crispus L. (Potamogetonaceae) 
was known in California from a single collection 
at Corona in 1918, while Pennisetum villosum R. 
Br. (Poaceae) was a “local fugitive from cultiva- 
tion” that Parish was aware of only from Santa 
Barbara and Ventura counties. Even currently ca- 
sual taxa have a potential long-term impact be- 
cause, unlike many kinds of environmental pollut- 
ants that are eventually neutralized within or dis- 
appear from the system upon termination of the 
source, biological pollutants such as naturalizing 
non-native plants are self-perpetuating, often in- 
creasing in prominence and distribution with time 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 67 


regardless of whether the source has been termi- 
nated (O’Kennon et al. 1999). Moreover, as stated 
above, the cost of control rises linearly as popula- 
tion size increases (Bayer 1999; Rejmanek 2000); 
because populations often enlarge logarithmically, 
ultimate control costs for populations left uncon- 
trolled may exceed available resources. As a result, 
awareness of the identity and ecology of a potential 
or incipient weed population is not only important, 
it is also economically imperative—if control or 
eradication are eventual goals. 

We hope that this compilation will spur two ac- 
tivities: first, further exploration for non-native 
spontaneous plants in California; and second, clar- 
ification of the current status of the taxa on this list, 
especially those classified as “‘No Current Infor- 
mation (NCI)’”’. Clearly, the disturbed areas on 
which many of these latter taxa established in the 
earlier decades of this century have subsequently 
been stabilized by parking lots or other intensive 
development; however, other populations may have 
expanded their range, but are on private land no 
longer readily accessible. 


On-going need for vouchered reports. The pre- 
sent compilation is not exhaustive for several rea- 
sons. First and foremost is the rapid rate at which 
new weedy taxa are being found in a state as large 
and geographically diverse as California. Secondly, 
a compilation such as this, which includes many 
records known only from obscure locations or sin- 
gle sites in California, comprises a major effort by 
botanical collectors of modern and previous eras; 
without their past and ongoing efforts this collation 
would not exist. With this in mind, as California’s 
increase in human population and consequent plant 
introductions continues, it is important that the col- 
lection and documentation of non-native taxa, by 
the submission of specimens to public collections, 
continue as well. 

As a final note we wish to again emphasize the 
absolute need for specimen documentation of new 
reports of spontaneous non-native plants (see Dean 
and Hrusa 2000 for instructions on collecting and 
documenting plant occurrences). The fact that de- 
termining and verifying these species does not keep 
pace with the introduction rate is largely a reflec- 
tion of the difficulty in applying names to plants 
whose geographic origins are not known and which 
may be cultivated forms or which have become 
phenetically modified in their new habitats. For this 
reason it has been discouraging to realize how 
many seemingly authoritative reports, via both 
agencies and private organizations, lack confirming 
specimen material. Indeed, some lacked even the 
documentation of who applied the name and when, 
or where the plant was reported to grow. Without 
such information a reliable understanding of which 
taxon is actually represented is not possible. The 
ramifications of this information gap are great; eco- 
logical behavior, environmental tolerances, devel- 


68 MADRONO 


opmental and phenological patterns are often spe- 
cies-specific, and control measures, whether phys- 
ical, chemical, or biological, may not be effective 
if the totality of life history is not correctly under- 
stood. The essential first step toward accessing 
whatever critical information is available is having 
an accurate identification (for a case study, see 
Thomas and Room 1986). 

The addition of so many non-native pest plants 
with the potential for becoming naturalized further 
underscores just how much the weed invasion of 
California should be of continuing economic and 
ecological concern. The current catalogue, while 
the result of intensive research in both the field and 
herbarium, is by no means complete; rather it 
serves to point out the extent to which non-native 
plants must be constantly sought out and reported. 


ACKNOWLEDGMENTS 


The current effort is dedicated to Thomas C. Fuller, 
Elizabeth McClintock, G. Douglas Barbe, and the late 
June McCaskill, our predecessors in the arena of docu- 
menting new occurrences of non-native plants in Califor- 
nia. Sincere appreciation and acknowledgment of our debt 
is also extended to the long list of individuals whose col- 
lective contributions, ranging from first collections of new 
occurrences to identifications of problematic taxa to other 
forms of assistance, made the catalogue possible: Carlos 
Aedo, Lowell Ahart, Mary Barkworth, James Barry, Steve 
Boyd, Robin Breckenridge, Richard Brummitt, Adolf Ces- 
ka, Steve Clemants, D. Gail DeLalla, Thomas Delendick, 
Joe DiTomaso, Ed Finley, Jesse Geissow, Jeff Glazner, D. 
Goldman, Jeff Hart, Evelyn Healy, Nancy Hillyard, Kevin 
Hintsa, Margaret Hurlbert, Lawrence Janeway, John Kar- 
tesz, David Keil, Dean Kelch, Rod Kerr, Dave Kratville, 
Elaine Mahaffey, Steve Matson, Mary Ann (Corky) Mat- 
thews, Dick Moe, Tony Morosco, Sergei Mosyakin, Barry 
Meyers-Rice, Robert Ornduff, Vern Oswald, Elizabeth 
Painter, David Philcox, Mike Pitcairn, Robert Preston, 
John Randall, Marcel Rejmanek, Tom Rosatti, Jake Ruygt, 
Mary DeValle Sanvictores, Rudi Schmid, Steve Schoenig, 
Teresa Sholars, Ernest Small, James P. Smith, John Stroth- 
er, Dean Wm. Taylor, Chris Thayer, Gordon Tucker, Mi- 
chael Vincent, Margriet Wetherwax, Dieter Wilken, Jo- 
seph Wipff, and Vern Yadon. Special thanks to these 
sharp-eyed individuals and any others, inadvertently over- 
looked, who have made vouchers or brought reports of 
new spontaneous non-natives to our attention. We are also 
indebted to Marcel Rejmanek and two anonymous review- 
ers for comments that vastly improved earlier versions of 
this manuscript. 


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BAYER, D. E. 1999. Biological, economic and public pol- 
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APPENDIX 1 
LisT OF TAXA WITH NATURALIZATION CLASS 


List of non-native vascular plant taxa occurring spon- 
taneously in California but not included in The Jepson 
Manual. Organization as in the Manual. Current Status 
subcategories in parentheses: NW = naturalized in wild- 
lands, N = naturalized (outside of wildlands), TEN = per- 
sistence tenuous, C = casual (waif), GH/C = greenhouse/ 
cultivation, EXT = extirpated. See Table 1 and Appen- 
dix 2. 


FERNS AND ALLIES 


Salviniaceae 
Salvinia molesta D.S. Mitch., (NW). 


Selaginellaceae 


Selaginella kraussiana (Kunze) A. Braun, (N). 


Ephedraceae 
Ephedra altissima Desf., (NCI). 
Ephedra distachya L., (NW). 
CONIFERS 
Pinaceae 
Pinus halepensis P. Mill., (N). 
Pinus pinea L., (NW). 
ANGIOSPERMS: DICOTS 
Acanthaceae 
Acanthus mollis L., (C). 


[Vol. 49 


Aceraceae 


Acer campestre L., (N). 
Acer saccharinum L., (NW). 


Aizoaceae 


Galenia pubescens (Ecklon & Zeyher) Druce var. pubes- 
cens, (N). 


Amaranthaceae 


Amaranthus blitum L. subsp. emarginatus (Uline & Bray) 
Carretero, (C). 

Amaranthus rudis Sauer, (N). 

Brayulinea densa (Willd.) Small, (NCI). 

Froelichia gracilis (Hook.) Mog., (NCI). 


Anacardiaceae 


Rhus lancea L.f., (N). 
Schinus polygamus (Cav.) Cabr., (N). 


Apocynaceae 


Nerium oleander L., (NW). 


Aquifoliaceae 


Ilex X attenuata Ashe, (NCI). 


Araliaceae 


Hedera canariensis Willd., (NW). 


Asclepiadaceae 


Asclepias fruticosa L., (C). 
Calotropis procera (Ait.) Ait. f., (C). 
Cynanchum louiseae Kartesz & Gandhi, (N). 


Asteraceae 


Achillea filipendulina Lam., (NCI). 

Artemisia vulgaris L. sensu stricto, (NCI). 

Aster novae-angliae L., (NCI). 

Carthamus oxyacantha M. Bieb., (C). 

Centaurea babylonica L., (C). 

Chrysanthemum balsamita L., (C). 

Cirsium scabrum (Poir.) Bonnett & Barratte, (NCI). 
Coreopsis grandiflora Hogg. ex Sweet, (C). 

Cotula mexicana (DC.) Cabrera, (N). 

Crepis tectorum L., (N). 

Dendranthema X grandiflorum Kitam., (NCI). 
Dittrichia graveolens (L.) Greuter, (NW). 

Emilia fosbergti D.H. Nicolson, (GH/C). 

Emilia sonchifolia (L.) DC. ex Wight, (GH/C). 
Grindelia papposa Nesom & Suh, (NCI). 

Helianthus tuberosus L., (C). 

Helichrysum petiolare Hilliard & B. L. Burtt., (NW). 
Heliomeris multiflora Nutt. multiflora, (TEN). 
Helipterum roseum (Hook.) Benth., (NCI). 
Lasiospermum bipinnatum (Thunb.) Druce, (NW). 
Leontodon muelleri (Schultz-Bip.) Fiori, (NCI). 
Oncosiphon piluliferum (L.f.) Kallersjo, (N). 
Osteospermum fruticosum (L.) Norl., (NW). 

Pericallis cruenta (DC. non Roth) Webb. & Berth., (NCI). 
Porophyllum ruderale (Jacq.) Cass., (C). 

Ratibida columnifera (Nutt.) Wooton & Standl., (NCI). 
Senecio squalidus L., (NCI. 

Tragopogon hybridus L., (NCI). 

Tripleurospermum maritimum (L.) W. D. J. Koch, (TEN). 


2002] 


Balsaminaceae 
Impatiens balfouri J.D. Hook., (N). 
Impatiens noli-tangere L., (N). 

Berberidaceae 


Berberis darwinii Hook., (NCI). 


Betulaceae 


Alnus cordata (Loisel.) Duby, (C). 


Bignoniaceae 

Campsis radicans (L.) Bureau, (TEN). 

Catalpa bignonioides Walter, (NW). 
Boraginaceae 

Echium lusitanicum L., (TEN). 

Pentaglottis sempervirens (L.) Tausch ex Bailey, (NCI). 
Brassicaceae 


Brassica fruticulosa Cyrillo, (N). 

Cardamine flexuosa With., (N). 

Coincya monensis (L.) Greuter & Burdet, (NW). 
Iberis umbellata L., (C). 

Rorippa sylvestris (L.) Besser, (TEN). 


Cabombaceae 


Cabomba caroliniana A. Gray, (NW). 


Campanulaceae 
Campanula medium L., (C). 
Lobelia erinus L., (C). 
Caprifoliaceae 
Leycesteria formosa Wallich., (NCI). 
Viburnum tinus L., (NCI). 
Caryophyllaceae 
Silene pseudatocion Desf., (NCI). 


Celastraceae 


Maytenus boaria Molina, (NW). 


Chenopodiaceae 


Atriplex muelleri Benth., (NCI). 
Chenopodium watsonii A. Nels., (NCI). 
Salsola kali L. subsp. pontica (Pallas) Mosyakin, (NCI). 


Cistaceae 


Halimium lasianthum (Lam.) Spach, (NCI). 


Convolvulaceae 


Calystegia silvatica (Kit.) Griseb. subsp. disjuncta Brum- 
mitt, (N). 

Convolvulus tricolor L., (NCI). 

Dichondra micrantha Urb., (NCI). 

Ipomoea aquatica Forssk., (C). 

Ipomoea lacunosa L., (C). 

Ipomoea quamoclit L., (C). 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 71 


Crassulaceae 


Crassula multicava Lem., (NW). 
Sedum album L., (N). 
Sedum dendroideum Sesse & Moc. ex DC., (NCI). 


Cucurbitaceae 


Cucumis anguria L., (NCI). 
Cucurbita ficifolia Bouche, (NCI). 
Cucurbita pepo L. var. medullosa Alef., (C). 


Cuscutaceae 


Cuscuta reflexa Roxb., (EXT). 


Droseraceae 


Drosera aliciae Hamet, (NCI). 
Drosera capensis L., (NCI). 
Drosera tracyi MacFarlane, (NW). 


Ebenaceae 


Diospyros virginiana L. var. virginiana, (NW). 


Elaeocarpaceae 


Aristotelia chilensis (Molina) Stuntz, (TEN). 
Muntingia calabura L., (GH/C). 


Escalloniaceae 


Escallonia macrantha Hook. & Arn., (NCI). 


Euphorbiaceae 


Euphorbia characias L., (NCI). 
Euphorbia cyathophora Murr., (NCI). 
Euphorbia dendroides L., (NW). 
Euphorbia heterophylla L., (NCI). 
Euphorbia hirta L., (N). 

Euphorbia hypericifolia L., (GH/C). 
Euphorbia marginata Pursh, (NCI). 
Euphorbia myrsinites L., (NCI. 
Euphorbia rigida M. Bieb., (NCI). 
Euphorbia terracina L., (NW). 
Sapium sebiferum (L.) Roxb., (NW). 


Fabaceae 


Astragalus cicer L., (TEN). 

Cassia nemophila A. Cunn., (TEN). 

Ceratonia siliqua L., (NW). 

Coronilla valentina L., (NW). 

Dolichos lignosus Pers., (NCI). 

Genista monosperma (L.) Lam, non Link, nec Del., (NW). 
Gleditsia triacanthos L., (NW). 

Lathyrus sativus L., (C). 

Ononis alopecuroides L., (NW). 

Robinia hispida L., (N). 

Senna artemisioides (Gaudich. ex DC.) Randell, (N). 
Senna obtusifolia (L.) H.S. Irwin & Barneby, (N). 
Sesbania punicea (Cav.) Benth, (N). 

Trifolium alexandrinum L., (NCI). 

Trifolium cernuum Brot., (N). 

Trifolium gemellum Poir. ex Willd., (N). 

Trifolium resupinatum L., (NCI). 

Trifolium retusum L., (NW). 

Trifolium stellatum L., (NCI). 

Trifolium striatum L., (NCI). 

Trifolium tomentosum Willk. ex Nyman, (NW). 


72 


Trifolium vesiculosum Savi, (N). 
Trigonella corniculata L., (NCI). 
Trigonella foenum-graecum L., (NCI). 
Vicia bithynica (L.) L., (NCI). 


Fagaceae 


Quercus ilex L., (N). 


Geraniaceae 


Geranium columbinum L., (NCI). 
Geranium lucidum L., (N). 
Geranium purpureum Vill., (NW). 
Geranium pyrenaicum Burm. f., (C). 
Geranium rotundifolium L., (NW). 


Geranium texanum (Trel.) A. Heller, (NCI). 


Hamamelidaceae 


Liquidambar styraciflua L., (TEN). 


Hydrophyllaceae 
Wigandia caracasana HBK., (NCI). 


Hypericaceae 


Hypericum androsaemum L., (N). 
Hypericum calycinum L., (TEN). 


Hypericum hookerianum Wight & Arn., (TEN). 


Lamiaceae 


Calamintha sylvatica Bromf. subsp. ascendens (JordaN). 


P.W. Ball, (TEN). 


Cedronella canariensis (L.) Willd. ex Webb & Berth., 


(NCI). 
Galeopsis tetrahit L., (NCI). 


Lamiastrum galeobdolon (L.) Ehrend. & Polatsch., (TEN). 


Lavandula stoechas L., (C). 

Mentha X villosa Huds., (NCI). 
Monarda citriodora Cerv., (N). 
Rosmarinus officinalis L., (C). 
Salvia longistyla Benth., (N). 

Salvia microphylla Benth., (NCI). 
Salvia reflexa Hornem., (GH/C). 
Salvia virgata Jacq., (NCI). 
Scutellaria caerulea M. & S., (C). 
Stachys floridana Shuttlew., (GH/C). 


Lauraceae 
Cinnamomum camphora (L.) J. Presl, (C). 
Laurus nobilis L., (TEN). 


Lentibulariaceae 


Utricularia subulata L., (NCI). 


Limnanthaceae 


Limnanthes macounii Trel., (N). 


Linaceae 


Linum trigynum L., (NCI). 


Malvaceae 


Anisodontea capensis (L.) Bates, (C). 
Anoda pentaschista A. Gray, (NCI). 
Gossypium hirsutum L., (NCI). 


MADRONO 


Hoheria populnea A. Cunn., (NCI). 
Lavatera olbia L., (NCI). 

Lavatera trimestris L., (NCI). 
Malva verticillata L., (NCI). 

Sida spinosa L., (NCI). 


Moraceae 


Fatoua villosa (Thunb.) Nakai, (GH/C). 
Ficus palmata Forssk., (NCI). 


Nymphaeaceae 


Nymphaea alba L., (NCI). 


Oleaceae 


Fraxinus uhdei (Wenz.) Lingel., (NW). 
Ligustrum lucidum W.T. Aiton, (NW). 
Ligustrum ovalifolium Hassk., (NW). 
Olea africana Mill., (TEN). 


Onagraceae 


Fuchsia magellanica Lam., (NW). 
Fuchsia X hybrida Voss., (NCI). 


Orobanchaceae 


Orobanche hederae Duby, (TEN). 


Papaveraceae 
Fumaria capreolata L., (N). 
Papaver X hybridum L., (NCI). 

Passifloraceae 


Passiflora caerulea L., (N). 
Passiflora manicata (Juss.) Pers., (NCI). 
Passiflora mixta L. f., (N). 


Passiflora tarminiana Coppens & Barney, (N). 


Pedaliaceae 


Sesamum indicum L., (C). 


Plumbaginaceae 


[Vol. 49 


Limonium ramosissimum (Poir.) Maire subsp. provinciale 


(Pignatti) Pignatti, (NW). 


Polygalaceae 


Polygala myrtifolia L., (NCI). 


Polygonaceae 


Polygonum multiflorum Thunb., (GH/C). 
Polygonum orientale L., (C). 


Ranunculaceae 


Caltha palustris L., (NW). 
Clematis terniflora DC., (TEN). 
Clematis vitalba L., (NCI). 

Nigella damascena L., (N). 
Ranunculus cortusifolius L., (TEN). 


Rhamnaceae 


Ziziphus jujuba L., (C). 


2002] HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 73) 


Rosaceae 


Cotoneaster lacteus W.W. Smith (NW). 
Crataegus monogyna Jacquin, (NW). 
Cydonia oblonga Mill., (NCI). 

Cydonia sinensis Thouin, (NCI). 

Eriobotrya japonica Lindl., (NCI). 
Filipendula vulgaris Moench, (NCI). 
Photinia davidsoniae Rehd. & Wilson, (NCI). 
Potentilla anglica Laicharding, (NW). 
Potentilla reptans L., (TEN). 

Prunus laurocerasus L., (NW). 

Prunus persica (L.) Batsch, (C). 

Prunus serrulata Lindl., (C). 

Pyracantha coccinea M. Roem., (TEN). 
Pyracantha crenatoserrata (Hance) Rehder, (N). 


Pyracantha crenulata (D. DON). M. Roem., (TEN). 


Pyrus communis L., (N). 
Rosa multiflora Thunb. ex Murray, (NCI). 
Rubus ulmifolius Schott var. ulmifolius, (N). 


Rubiaceae 


Coprosma repens A. Rich., (N). 


Salicaceae 


Populus nigra L. cv. “‘Ttalica”’, (TEN). 


Sarraceniaceae 


Sarracenia aff. rubra Walter, (N). 


Scrophulariaceae 


Anarrhinum bellidifolium (L.) Willd., (C). 
Limnophila X ludoviciana Thieret, (TEN). 

Mazus japonicus Kuntze, (GH/C). 

Penstemon strictus Benth., (TEN). 

Penstemon subglaber Rydb., (TEN). 

Scrophularia peregrina L., (N). 

Verbascum olympicum Boiss. non Bunyard, (NW). 


Solanaceae 


Atropa belladonna L., (NCI). 

Capsicum annuum L., (NCI). 

Cestrum parqui LU Her., (NCI). 

Lycium ferocissimum Meirs, (NCI). 
Nicotiana X sanderae Hort. ex Wats., (C). 
Nicotiana tabacum L., (C). 

Petunia violacea Lindl., (NCI). 

Solanum gayanum (Remy) Phil. f., (NCI). 
Solanum scabrum Mill., (C). 

Solanum villosum Mill., (C). 


Urticaceae 
Boehmeria cylindrica (L.) Sw., (NW). 
Laportea aestuans (L.) Chew, (GH/C). 
Verbenaceae 
Verbena rigida Spreng., (NCI). 
Vitex agnus-castus L., (NCI). 
Vitaceae 


Cissus antarctica Venten., (NCI). 
Vitis aestivalis Michx., (NCI). 
Vitis rupestris Scheele, (TEN). 


Zy gophyllaceae 
Peganum harmala L., (EXT). 


ANGIOSPERMS—MONOCOTS 
Alismataceae 


Sagittaria brevirostra Mackenzie & Bush, (NCI). 
Sagittaria rigida Pursh, (NW). 


Araceae 


Arum palestinum Boiss., (TEN). 
Dracunculus vulgaris Schott, (N). 
Pinellia ternata (Thunberg) Makino, (NCI). 


Cyperaceae 


Bulbostylis barbata Kunth, (GH/C). 

Cyperus flavescens L., (NW). 

Cyperus flavicomus Michx., (N). 

Cyperus gracilis R. Br., (NCI). 

Cyperus iria L., (TEN). 

Cyperus owanii Boeck, (NCI). 

Cyperus papyrus L., (NCI). 

Fimbristylis autumnalis (L.) Roem. & Schult., (NW). 
Scirpus cyperinus (L.) Kunth, (NW). 

Scirpus prolifer Rottb., (NCI). 


Eriocaulaceae 


Eriocaulon cinereum R. Br., (EXT). 


Hydrocharitaceae 


Limnobium laevigatum (Humb. & Bonpl. ex Willd.) Hei- 
ne, (NW). 


Iridaceae 


Tris foetidissima L., (N). 

Iris germanica L., (TEN). 

Tris orientalis L., (NCI). 

Ixia polystachya L., (NCI). 

Ixia speciosa Andrews, (NCI). 

Moraea collina Thunb., (N). 

Moraea polystachya Ker Gawl., (NCI). 


Juncaceae 


Juncus nodatus Cov., (N). 


Liliaceae (sensu lato) 


Agapanthus praecox Willd., (NCI). 
Allium cepa L., (NCI). 

Allium sativum L., (C). 

Amaryllis belladonna L., (TEN). 
Chlorophytum capense (L.) Druce, (C). 
HAyacinthus orientalis L., (C). 
Kniphofia uvaria (L.) Hooker, (NW). 
Leucojum aestivum L., (NCI). 
Narcissus pseudonarcissus L., (NW). 
Narcissus tazetta L., (NW). 
Ornithogalum umbellatum L., (GH/C). 
Pancratium maritimum L., (N). 

Tulipa clusiana DC. in Redoute, (TEN). 


Poaceae 


Acrachne racemosa (Roem. & Schult.) Ohwi, (TEN). 
Aira caryophyllea L. var. cupaniana (Guss.) Fiori, (NCI). 


74 MADRONO 


Amphibromus neesii Steud., (NW). 

Aristida dichotoma Michx., (NW). 

Chloris truncata R. Br., (N). 

Echinochloa crusgalli subsp. spiralis (Vasing.) Tzvelev, 
(N). 

Echinochloa esculenta (A. Br.) H. Scholz, (NCI). 

Ehrharta longiflora Sm., (NW). 

Eragrostis curvula (Schrad.) Nees var. conferta Nees, (N). 

Eremochloa ciliaris (L.) Merr., (EXT). 

Gaudinia fragilis (L.) P. Beauv., (NW). 

Glyceria fluitans (L.) R. Br., (NW). 

Hordeum vulgare L. sensu lato, (C). 

Leptochloa dubia (Kunth) Nees, (N). 

Nassella tenuissima (Trin.) Barkworth, (C). 

Panicum maximum Jacq., (GH/C). 

Panicum repens L., (NCI). 

Panicum rigidulum Bosc ex Nees var. rigidulum, (NCI). 

Panicum texanum Buckl., (TEN). 

Pennisetum glaucum (L.) R. Br., (C). 

Pennisetum latifolium Spreng., (NCI). 

Phalaris coerulescens Desf., (C). 

Phyllostachys aurea A. & C. Riviere, (NCI). 

Phyllostachys bambusoides Siebold & Zuccarini, (NCI). 

Piptochaetium stipoides Hackel ex Arech. sensu lato, 
(NW). 

Polypogon imberbis (Phil.) Bjorkm., (NCI). 

Pseudosasa japonica (Sieb. & Zucc. ex Steud.) Makino 
ex Naka, (NCI). 

Schedonnardus paniculatus (Nutt.) Trel., (NCI). 

Spartina anglica C.E. Hubb., (NW). 

Stipa capensis Thunb., (NW). 

Themeda quadrivalvis (L.) Kuntze, (TEN). 

Tribolium obliterum (Hemzl.) Renvoize, (NW). 

Triticum aestivum L., (C). 


Pontederiaceae 


Heteranthera rotundifolia (Kunth) Griseb., (N). 


APPENDIX 2 
ANNOTATED CATALOGUE 


Family circumscriptions and organization follow The 
Jepson Manual. Generic and specific applications reflect 
published treatments by specialists, modified only if clear- 
er information was gained by utilizing an alternative no- 
menclature. Abbreviations are as follows: DISTRIBU- 
TION (DIST) with geographic subdivisions as used in 
The Jepson Manual; CURRENT STATUS (CS); Current 
Status subcategories: NATURALIZED IN WILDLANDS 
(NW); NATURALIZED (OUTSIDE OF WILDLANDS) 
(N); PERSISTENCE TENUOUS (TEN); CASUAL (waif) 
(C); GREENHOUSE/CULTIVATION (GH/C); EXTIR- 
PATED (EXT); DOCUMENTATION (DOC). For details 
and descriptions of these categories see Tables 1 and 2. 


FERNS AND ALLIES 
Salviniaceae 


Salvinia molesta D.S. Mitch.: DIST: CCo, DSon, SCo: 
CS: NW, N (DSon), TEN (SCo): DOC: Imperial Co.: N 
side Hwy 98 in E Highline Canal nr. Winterhaven. T16S, 
R20E, Sec. 01, SB. Aug. 25, 1999, Johnson s.n. (CDA); 
All American Canal at Drop 1 nr. Coachella Canal. Sam- 
ple from equipment which at this point collects vegetation 
floating down the canal. Winterhaven area. T16S, R20E, 
Sec. 31, SB. Aug. 25, 1999, Johnson s.n. (CDA); River- 
side Co.: Drain canal of Palo Verde Irrig. District, Blythe 


[Vol. 49 


region. TO6S, R23E, SB. Aug. 20, 1999, R. O'Connell 
s.n. (CDA); San Diego Co.: Private pond in vicinity of 
Fallbrook off Harris Truck trail. Oct. 28, 1999. S. Riviera 
s.n. (CDA); San Luis Obispo Co.: Private pond off Price 
Cyn. Rd. T30S, R12E, MD. Nov. 27, 2001, S. Stoltz s.n. 
(CDA, OBI); NOTES: Reported from San Diego River 
(San Diego Co.) Aug., 1999 but no specimen has been 
seen. Fallbrook location may have been purposely planted 
for increase and resale. Eradication attempts are currently 
underway by CDFA, USDA-APHIS in all sites. A Federal 
Noxious Weed, sale or growth is prohibited. 


Selaginellaceae 


Selaginella kraussiana (Kunze) A. Braun: DIST: 
NCo: CS: N: DOC: Humboldt Co.: Arcata. Damp, shad- 
ed areas near Humboldt State University Conservatory. 
April 5, 2000, G. Leppig 1286 (HSC); redwood forest be- 
hind Humboldt State University, along stream at Fern 
Lake. April 12, 2000, G. Leppig 1290 (CDA, HSC); Son- 
oma Co.: Best, C., et al. (1996, pg. 25). 


Ephedraceae 


Ephedra altissima Desf.: DIST: SCo: CS: NCI: DOC: 
San Diego Co.: Spreading from root shoots and climbing 
to 7 m to top of adjacent macadamia tree, site of old 
USDA Exp. Station, N of Science Park Dr., E of North 
Torrey Pines Rd, N of La Jolla. T15S, RO4W, Sec. O1, 
SB. Oct. 1, 1974, 7.C. Fuller 20049 (CDA). 

Ephedra distachya L.: DIST: SCo: CS: NW: DOC: 
Santa Barbara Co.: Smith, C.E (1998, pg. 372): 
NOTES: Reported as ‘established’ in Smith (1998), but 
by our criteria Wilken (SBBG) reports that it is naturalized 
in the oak woodlands about the Trout Club in the Santa 
Ynez Mtns. (pers. comm.). Det. by D. Wilken, verified 
also by S. Carlquist (SBBG). 


CONIFERS 
Pinaceae 


Pinus halepensis P. Mill.: DIST: CCo, SnFrB: CS: N: 
DOC: Contra Costa Co.: Mount Diablo, W slope Mount 
Zion, upper extent of Kaiser quarry reddish chert-like 
rocks on steep 25% W facing slope; mature trees (planted 
in rows), with abundant reproduction, juveniles to 10 ft 
tall. Elev. 1550 ft, TOIN, ROIW, Sec. 22, MD. Oct. 9, 
1996, D.W. Taylor 15896 (UC); San Mateo Co.: Mc- 
Clintock, E., et al. (1990, pg. 62): NOTES: Seedlings are 
occasionally encountered about cultivated trees but are 
seldom allowed to mature. 

Pinus pinea L.: DIST: n Chl, SnFrB: CS: naturalized: 
DOC: Contra Costa Co.: Walnut Creek, Lakewood sub- 
division, on hillside. June 21, 1950, W.S. Malloch s.n. 
(UC); Santa Barbara Co.: Junak. S. et al. (1995, pg. 64). 


ANGIOSPERMS—DICOTS 
Acanthaceae 


Acanthus mollis L.: DIST: SnFrB: CS: C: DOC: San 
Mateo Co.: McClintock, E., et al. (1990, pg. 63): 
NOTES: Probably originating via garden waste; individ- 
uals sites often long-persistent, but permanence tenuous. 


Aceraceae 


Acer campestre L.: DIST: SnFrB: CS: N: DOC: Ala- 
meda Co.: lower Strawberry Canyon firetrail behind UC 
Berkeley campus, a few small trees at edge of woodland, 


2002] 


June 11, 1998, B. Ertter & A. Rusev 16152 (UC): 
NOTES: Verbal reports indicate it may be sparingly nat- 
uralized elsewhere in the East Bay Hills. 

Acer saccharinum L.: DIST: ScV: CS: NW: DOC: 
Glenn Co.: Sacramento River at Butte City boat launch 
site, small tree on silty river bank. Oct. 22, 1998, Ertter 
16448 (UC); Bank btwn. levee and W side of Butte Creek 
approx. % mi NE of confluence with Howard Slough, and 
immed. below McPherrin Dam (private and scheduled for 
removal). Two multi-trunked trees, + 12 meters tall, both 
apparently sterile, possibly persisting from old plantings. 
59-2 TIN] P2153" We Sept 6s’ 1996hG.F* “Hrasa’ 13571 
(CDA): Sacramento Co.: Sacramento River in Sacramen- 
to near the end of 10th St., elev. 85 ft. Apr. 28, 1997, D. 
Kelch DGK97.012 (UC); Betw. N side American River 
and William Pond, American River Parkway at Arden 
Way entrance. 38°33’N; 121°22’W. June 28, 1998, B. Mey- 
ers-Rice MR980601 (CDA, DAV); N bank American Riv- 
er, American River Parkway, few hundred meters down- 
river of the Estates Dr. entrance. June 28, 1998, B. Mey- 
ers-Rice MR980602 (DAV); American River Recreation 
Trad; E’ oi Union Pacific trestle. TO7N, ROSE, Sec. 31, 
MD. April 13, 1984, G.D. Barbe 4142 (CDA, DAV). 


Aizoaceae 


Galenia pubescens (Ecklon & Zeyher) Druce var. pu- 
bescens: DIST: SCo: CS: N: DOC: Los Angeles Co.: 
Ross, T. (1993, pg. 226-228); Ross, T. and S. Boyd (1996, 
pg. 432—433); Riverside Co.: U.S. Forest Fire Lab, Cyn. 
Crest Dr. El. 1200 ft. Fairly common in experimental 
planting of Artemisia californica, Eriogonum fasciculatum 
etc., Doubtless progeny of plants grown by E.C. Nord in 
1970. Aug. 9, 1996, J. Beyers s.n. (UCR): NOTES: A 
specimen from San Diego Co. at UCR has not yet been 
accessioned. 


Amaranthaceae 


Amaranthus blitum L. subsp. emarginatus (Uline & 
Bray) Carretero: DIST: SCo: CS: C: DOC: Los Angeles 
Co.: waste ground at intersection of Beach Blvd. (Hwy 
39) and Rosecrans Ave, at the Orange County line, La 
Mirada. Oct. 29, 1988, T. Yutani s.n. (CDA); Solitary 
small weed in a potted plant purchased at The Farm Store, 
Cal Poly Pomona. Jan. 10, 2002, A.C. Sanders 2489] 
(UCR): Riverside Co.: Moreno Valley, weed in untended 
planter beside Home Depot at Pigeon Pass Rd. and Hwy 
160. Nov. 25, 2001, A.C. Sanders 24887 (UCR): NOTES: 
Determination by S. Mosyakin (KW, Mar. 2001), [=A. 
blitum subsp. polygonoides (Mogq.-Tandon) Carretero]. 
See Costea et al. (2001) for taxonomic discussion. Occa- 
sionally intercepted during nursery stock inspections on 
material from the US southeast, esp. Florida. Adapted to 
tropical and subtropical climates, this species would not 
be expected to be more than a minor or casual weed of 
greenhouse, garden or nursery. Sometimes treated (and 
reported) as A. lividus L. 

Amaranthus rudis Sauer: DIST: SCo, ScV: CS: N: 
DOC: Butte Co.: E side Sac. R. NW Parrott Landing, 1 
mi. SE Ord Ferry, 12 mi. SW Chico. Riparian woodland. 
Sept. 21, 1999, L. Ahart 8267, 8266 (CDA, CHSC); 6 mi. 
W Chico, gravel bar, Sac. R. Sept. 26, 1981, L. Ahart 3205 
(CAS, CDA, CHSC), det. by J.T. Howell 12/81; Sacra- 
mento Co.: NW corner Int. US 50 and Howe Ave., Sac- 
ramento. Single plant. Sept. 22, 1985, A. Shapiro s.n. 
(CDA, DAV); Santa Barbara Co.: SPRR, Carpenteria. 
Sept. 5, 1957, H.M. Pollard (CAS, CDA), det. J. Sauer 5- 
1959 (as A. tamariscinus Nutt.); SPRR yards, Santa Bar- 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA > 


bara. Sept. 26, 1957, H.M. Pollard s.n. (CAS, CDA), det. 
J. Sauer 5/1959; loc. cit. Oct. 8, 1957, H.M. Pollard s.n. 
(DAV), det. J. Sauer 2/1974; SPRR yards, Santa Barbara. 
Oct. 8, 1957, H.M. Pollard s.n. (DAV): NOTES: Similar 
among dioecious California Amaranthi to A. arenicola. 
Pistillate plants readily distinguished among the California 
taxa by their two, rather than five pistillate sepals. Sta- 
minate plants of A. rudis have acute to acuminate sepal 
tips in contrast to the obtuse to retuse sepal tips in A. 
arenicola. Amaranthus tamariscinus Nutt. misapplied. 
Staminate and pistillate plants comprising Townsend s.n., 
Aug. 2, 1968, (CDA, RSA) collected nr. Pala in San Diego 
Co. and the source of the citation for A. tamariscinus in 
Munz (1974) USDA, NRCS (2001) and Beauchamp 
(1986), are, acc. to Hrusa, misidentified A. palmeri S. 
Wats. 

Brayulinea densa (Willd.) Small: DIST: SCo: CS: 
NCI: DOC: Santa Barbara Co: Howell, J.T. (1966, pg. 
256); NOTES: Probably a roadside casual, not since re- 
ported in California. 

Froelichia gracilis (Hook.) Mog.: DIST: SCo: CS: 
NCI: DOC: Los Angeles Co.: Santa Fe RR, San Dimas. 
June 23, 1955, G.W. Garrettson s.n. (CDA). Det. by PA. 
Munz. 


Anacardiaceae 


Rhus lancea L.f.: DIST: SCo, DSon: CS: N: DOC: 
Riverside Co.: Coachella Valley, Palm Springs, W side 
of Palm Canyon Dr. just above (S of) junction with Hwy 
111, weedy disturbed vacant lot, clearly spontaneous. 
33°47.75'N, 116°32.52'W; TO4S, RO4E, Sec. 27, SB. EI. 
140 m. Mar. 17 1996, A. C. Sanders and G. Helmkamp 
17982 (UCR); Palm Springs, Murray Canyon, off Palm 
Canyon, a solitary arborescent shrub 4—5 m tall at the 
edge of the stream between palm groves, elev. 245 m, far 
from any cultivated plants, 33°45.5’N, 116°33’W, TO4N, 
RO3E, Sec. 10, SB. Feb 8, 1997, A. C. Sanders et al 19686 
(UCR); Riverside, S side of the U.C.R. campus, a solitary 
shrub, apparently spontaneous, growing wedged between 
a large pecan tree and the wall of a concrete reservoir. 
Elev. 400 m. Feb 26, 1997, A. C. Sanders 19688 (UCR); 
Ventura Co.: Hills north of Moorpark, 0.5 mile west of 
Happy Camp Canyon, 1.9 miles north of Arroyo Simi 
Channel, TO2ZN R19W sec. 33, elev. 245-275 m, invasive 
in coastal sage scrub, particularly in small drainages. Jul. 
27, 1995, C. Jones and R. Ramirez 8 (RSA, UCR): 
NOTES: Native to S. Africa and widely cultivated in Cal- 
ifornia, these are the first records of naturalized plants in 
California. Doubtless naturalized elsewhere in S. Calif. as 
the locations where it has been found represent much of 
the range of environmental conditions in lowland southern 
California, from the Sonoran Desert to the mild maritime 
influenced climate of Ventura County. 

Schinus polygamus (Cay.) Cabr.: DIST: SCo: CS: N: 
DOC: Los Angeles, Riverside, San Bernardino Cos.: 
Sanders, A.C. (1996, pg. 530): NOTES: Widespread in 
urban So. Calif., but is widely ignored by collectors. Still 
scarce in + natural areas, but often encountered in dis- 
turbed sites. 


Apocynaceae 


Nerium oleander L.: DIST: SCo, ScV: CS: NW: DOC: 
Los Angeles Co.: San Gabriel Mtns., in and near Glen- 
dora Wilderness Park, Harrow Canyon at the third debris 
basin. Elev. 1450 ft, Jul. 14, 1989, D. Swinney s.n. (UCR); 
Riverside Co: Temple, PJ. (1999, pg. 55); San Bernar- 
dino Co.: Waterman Canyon Road at old Arrowhead Hot 


716 MADRONO 


Springs Resort, 7 km north Hwys 30 and 18 intersection. 
Sandy/rocky riparian with Artemisia californica and Vitis 
girdiana, May 9, 1972, E. Trubschenck 28 (UCR); Wa- 
terman Canyon, 1 km W of the old Arrowhead Hot 
Springs Hotel, 117°16'W, 34°11.5’'N, TOIN, RO4W, Sec. 
11, SB. Elev. 1900 ft, abundant shrub in moist areas along 
the rocky canyon bottom along creek, numerous seedlings 
and plants of all sizes. Apr. 27, 1993, A. C. Sanders et al. 
13824 (UCR); along Colorado River north of Parker 
Bridge, TOIN, R26E, Secs. 17 and 18, SB. Elev. 350 ft, 
mostly alkali and disturbed sites. May 2, 1978, Faulkner 
572 (UCR).: Shasta Co.: Keeley, J., (1992, pg. 157): 
NOTES: Naturalized at the Waterman Canyon site for at 
least 30 years. Plants are abundant there, forming large 
thickets along the creek and obviously reproducing sex- 
ually. There is considerable variation in flower color, in- 
cluding shades of pink never seen in cultivation. Also es- 
tablished in nearby Hot Springs and Strawberry Canyons. 
The Swinney collection certainly represents a naturalized 
population as it shows (dried, no color notes) the mottled 
pink flower color that is common in the reproducing pop- 
ulation at Waterman Canyon. The Faulkner collection 
might be questioned because the dried flowers appear to 
be the pure white which is common in cultivated plants. 
There is nothing on the label which indicates that this 
collection was from cultivated or persisting plants, but 
there also is nothing eliminating that possibility. This re- 
port needs confirmation. 


Aquifoliaceae 


Ilex X attenuata Ashe: DIST: ScV: CS: NCI: DOC: 
Sacramento Co.: Volunteer tree on creek bank. Hoffman 
Lane, Fair Oaks. Jan. 30, 1977, Bly s.n. (CDA). 


Araliaceae 


Hedera canariensis Willd. (incl. H. algeriensis Hibb.): 
DIST: CCo, SCo, SnBR, SnFrB, SnGB: CS: NW: DOC: 
Alameda Co.: Albany Hill, abundant in oak forest. Jan. 
14, 1995, B. Ertter 13918 (UC); Orange Co.: Trabuco 
Canyon. May 13, 1966, E.W. Lathrop 6297 (UCR); San 
Bernardino Co.: upper Waterman Canyon, San Bernar- 
dino Mtns. Dec. 23, 1998, A.C. Sanders 22369 (UCR); 
Dick Stoddard Canyon, San Gabriel Mtns. Jan. 15, 1994, 
Swinney 2695 (UCR); NOTES: Differs from H. helix in 
having larger leaves with fewer (most often only 3), more 
rounded lobes and fainter veins; more robust stems that 
are more shallowly rooted at the nodes; and rusty hairs 
on the growing tips that are basally fused into tightly ap- 
pressed, stellate-peltate trichomes (vs. white and + 
spreading in H. helix). Some naturalized forms not clearly 
separable from H. helix, needs further study. Occurring 
throughout the San Francisco Bay Area, where capable of 
being a serious pest plant; probably more common than 
H. helix in that region. 


Asclepiadaceae 


Asclepias fruticosa L.: DIST: deltaic GV: CS: C: 
DOC: Contra Costa Co.: Martinez Regional Shoreline, 
Granger’s Wharf Park, north end of Berrelessa St. Aug. 
21, 1977, Walter and Irja Knight 3158 (CAS): NOTES: 
Narrow leaves like A. fascicularis, but pods ovate with 
long bristles. Not relocated in 2001, but site had been 
recently denuded by heavy grading. 

Calotropis procera (Ait.) Ait. f.: DIST: DSon: CS: C: 
DOC: Imperial Co.: agricultural area in the Imperial Val- 
ley. August, 1987, F. Laemmlen s.n. (UCR): NOTES: A 


[Vol. 49 


solitary shrub, grower was concerned about its potential 
as a weed and is reported to have destroyed it after send- 
ing material for determination. 

Cynanchum louiseae Kartesz & Gandhi: DIST: SCo: 
CS: EXT: DOC: Riverside Co.: Sanders, A.C. (1996, pg. 
526, 527); NOTES: The site of this population, which had 
been established for many years, was re-landscaped re- 
cently and the plants could not be relocated in 2001. 


Asteraceae 


Achillea filipendulina Lam.: DIST: NCo, SCo: CS: 
NCI: DOC: Humboldt Co.: dense growth in unkempt 
yard, McKinleyville. July 7, 1975, F. Bapeaux s.n. 
(CDA); Ventura Co: light infestation, 0.01 acre net over 
1.0 acre gross, vacant lot, Poncho Rd., Camarillo. TOIN, 
R20W, Sec. 05, SB. June 20, 1979, H. Carpenter s.n. 
(CDA): NOTES: Gen. persistent and spreading vegeta- 
tively from cultivation, probably rarely reproducing by 
seed. 

Artemisia vulgaris L. sensu stricto: DIST: SCo, SNE: 
CS: NCI: DOC: Ventura Co.: Waste area of old habita- 
tion site between Ventura Ave. and Southern Pacific RR 
hear Wadstrom. Oct. 10, 1969, H.M. Pollard s.n. (CAS, 
CDA, SBBG); Mono Co.: Rock Creek Basin Rd. 0.1 mile 
N of Mono/Inyo Co. line, just S of Pine Grove Camp- 
ground, Inyo National Forest. TO5S, R30E, Sec. 31, MD. 
Aug. 12, 1981, G.D. Barbe 3532 (CDA): NOTES: De- 
terminations by S. Mosyakin (KW), 3/2001. Acc. to Mo- 
syakin, several other non-native taxa of the A. vulgaris 
alliance are represented at CDA but none can at present 
be associated with a specific name. 

Aster novae-angliae L.: DIST: KR: CS: NCI: DOC: 
Siskiyou Co.: Single plant in roadside ditch, escape from 
garden across the rd, Etna, Scotts Valley. Oct. 6, 1966. 
T.C. Fuller 15244 (CDA). 

Carthamus oxyacantha M. Bieb.: DIST: CCo: CS: C: 
DOC: Monterey Co.: Waif, screening disposal area, va- 
cant field south of spice processing plant, Schilling Place, 
Salinas. T15S, RO3E, Sec. 03, MD. Aug. 2, 1978. G.D. 
Barbe 2421 (CDA): NOTES: Carthamus oxyacantha is 
on the Federal Noxious Weed Act quarantine list (see Fed- 
eral Register, May 25, 2000, p. 33741-33743). Native to 
South Africa. Related species are aggressive invaders of 
pastures in New Zealand, Australia, California. 

Centaurea babylonica L.: DIST: n SNH: CS: C: DOC: 
Plumas Co.: Spontaneous at edge of lawn, to 6 ft tall, 
large rosette. County Hospital, Quincy. Aug. 7, 1972. F.H. 
Surber s.n. (CDA). 

Chrysanthemum balsamita L.: DIST: CaR: CS: C: 
DOC: Siskiyou Co.: volunteer, street side, Hennesy St., 
McCloud. Sept. 17, 1976, F.D. Horn s.n. (CDA). 

Cirsium scabrum (Poir.) Bonnett & Barratte: DIST: 
CCo: CS: NCI: DOC: Santa Cruz Co.: Howell, J.T, 
(1959, p. 27): NOTES: Collection made by A. Eastwood 
in 1900 and filed under the synonym Cnicus giganteus 
(Desf.) Willd. (UC). Acc. to J. Kartesz, this has also been 
reported as Cirsium giganteum (Desf.) Spreng. 

Coreopsis grandiflora Hogg. ex Sweet: DIST: CaR, 
SnFrB: CS: C: DOC: Alameda Co.: vacant lot, Appian 
Way, Union City. June 3, 1976, E. Whitaker s.n. (CDA); 
Plumas Co.: Disturbed roadside along Squirrel Ck, USFS 
road leading to Argentine Rock. Ca. 1 mi NE of Hwy 70, 
and ca. 7 mi E of Quincy. Elev. 1360 m. 39°55'N; 
120°47'30"W. Aug. 22, 1996, G.F. Hrusa 13532 (CDA); 
San Bernardino Co.: a single waif, distinct from dwell- 
ings, San Bernardino Valley. June 10, 1909, S.B. Parish 
TA SI SIE): 


2002] 


Cotula mexicana (DC.) Cabrera: DIST: CCo, ScV, 
SnFrB: CS: N: DOC: Alameda Co.: golf course green, 
Livermore. Apr. 20, 2000, E. de Villa s.n. (CDA); golf 
course green, Hayward. Apr. 26, 2000, G. Ingram s.n. 
(CDA); golf course green, Castro Valley. Apr. 26, 2000, 
G. Ingram s.n. (CDA); Marin Co.: golf course green, No- 
vato. Apr. 29, 2000, G. Ingram s.n. (CDA); Monterey 
Co.: golf course green, Fort Ord. Nov. 12, 2001, S. Fen- 
nimore s.n. (CDA, DAV); Sacramento Co.: golf course 
green, Elverta. Dec. 23, 1999. R. Chavez s.n. (CDA); golf 
course green, Sacramento. Jan. 21, 2000, V. Nyvall s.n. 
(CDA); golf course green, Sacramento. Jan. 9, 2000, F. 
Carl s.n. (CDA); golf course green, Galt. Feb. 9, 2000, 
D. Thompson s.n. (CDA); golf course green, Elk Grove. 
March 16, 2000, F. Carl, V. Nyvall s.n. (CDA); San Ma- 
teo Co.: golf course green, Pacifica. Mar. 21, 2002, D. 
Pendleton s.n. (CDA); Loc. cit., Apr. 18, 2002, (UCR); 
Siskiyou Co.: golf course green, Mt. Shasta region. Dec. 
1, 2000, D. Smith s.n. (CDA): NOTES: A large infestation 
observed also in Napa Co. on a golf course near Pope 
Valley. Rapidly spreading perennial capable of competing 
and establishing by seed in mature low cut turf of greens 
and the adjacent collar. Not yet seen in taller mowed turf, 
or in wild situations, but expected in the latter. Easily 
overlooked due to diminutive stature. Native southern 
Mexico to Boliva, apparently at elevations above 3000 m. 

Crepis tectorum L.: DIST: SnBr, s SNH: CS: N: DOC: 
Inyo Co.: Mammoth Lakes, 37°38.8’N, 118°58.5'W, elev. 
8100 ft. Uncommon in one local area on roadside among 
pines. Sept. 26, 1996, G. Helmkamp 1218 (UCR); San 
Bernardino Co.: Sanders, A.C. (1997b, pg. 307). 

Dendranthema X grandiflorum Kitam.: DIST: SCo: 
CS: NCI: DOC: Ventura Co.: heavy infestation in yard, 
Santa Paula. October 27, 1964, C.J. Barrett and V. Hol- 
mer s.n. (CDA): NOTES: This is the florist’s chrysanthe- 
mum. Original determination as Chrysanthemum morifol- 
ium Ramat. 

Dittrichia graveolens (L.) Greuter: DIST: CCo, SCo, 
ScV, SnFrB: CS: NW: DOC: Alameda Co.: San Francis- 
co Bay Wildlife Refuge at end of Cushing Road, forming 
dense masses on levee. Nov. 3, 1995, B. Ertter 14542 
(UC); Shadow Cliffs Recreation Area, Livermore Valley, 
abundant in riparian woodland. Sept. 28, 1998, B. Ertter 
16412 (UC); Merritt College, common in parking lot on 
west side of campus. Oct. 8, 2000, Ertter 17540 (UC): 
Gravelly sidewalk strip at corner of Monaco and Mission 
Drs. in Pleasanton. 37°39'08"N, 121°52’45”W. Nov. 22, 
2000, D. Petersen 00-54. (CDA); Niles Cyn. Rd betw. 
Sunol and Fremont, mile marker 16. In Alameda Cr., sand/ 
eravel. bat,.37° 35.43" N,/121°54'22"W.. Oct: 27, 2001,-D. 
Petersen s.n. (CDA); Site M.5 storage area in Camp Parks, 
Dublin. Disturbed, graveled parking area. 37°43'28’N, 
121°52'30"W. Oct. 17, 2001, D. Petersen 240 (CDA); Ar- 
royo de la Laguna at Verona Bridge (Pleasanton) on gravel 
bate” beside Stream. © 37757/35"N,, 121°52'55" W.- Nov:: 14, 
2001, D. Petersen s.n. (CDA); Alameda/Contra Costa 
Co. [line]: Redwood Regional Park, West Ridge Road. 
Oct. 15, 2000, Ertter 17541 (UC): Contra Costa Co.: 
Lime Ridge, sterile flat in bowl of quarry area just south 
of Ygnacio Valley Road, in vicinity of Ygnacio Reservoir, 
Concord. Locally common. TOIN, ROIW, Sec. 08, MD. 
Oct. 18, 1998, B. Ertter and W.A. Morosco 16423 (JEPS): 
loc. cit. Dec. 29, 1999, Case and Ertter s.n. (CDA); San 
Mateo Co: About twenty plants on trailside, E end of 
Weeks St. adj. to Bay Lands Nature Preserve, E. Palo 
Alto. Oct. 18, 2001, J. Beall s.n. (CDA); Santa Clara 
Co.: Preston, R.E. (1997, 200-203); abundant, two miles 
north of Alviso Railroad tracks at upper edge of tidal 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA V7 


marsh. Nov. 1, 1984, H.T. Harvey s.n. (CDA, SJSU, UC). 
Det. by C.W. Sharsmith, 5-88.; Overflow channel east of 
Coyote Creek, 25 yards W of Milpitas Sewage Treatment 
Plant. % acre. Oct. 16, 2000, J. Beall s.n. (CDA, UCR): 
roadside and in pasture at 4010 Calaveras Rd., Milpitas. 
Nov. 15, 2000, N. Garrison s.n. (CDA); Solano Co.: Sui- 
sun City, parking lot opposite wildlife center at Peytonia 
Slough, also near Civic Marina. Oct. 2, 2000, A.M. Sha- 
piro s.n. (DAV); Yolo Co.: City of West Sacramento, 
about 0.5 km W of Harbor Blvd., N of West Capitol Ave., 
along a jeep trail on the S side of the railroad embank- 
ment. Oct. 29, 1999, A.M. Shapiro s.n. (DAV): NOTES: 
Also noted by Ertter at Coyote Hills Recreation Area, and 
Lake Del Valle; by K. Hintsa at Rock City, Mount Diablo, 
all Alameda Co. D. Petersen (pers. comm. to Hrusa, 10- 
2001), reports it is becoming common at Camp Parks, 
Alameda Co., where it occupies creekbanks as well as 
roadcuts and roadsides. A much-branched, densely glan- 
dular, odoriferous, fall-blooming annual, superficially 
reminiscent of a tarweed but with narrow overlapping 
phyllaries, or of Conyza but with yellow flowers. Ray 
flowers are reduced, and leaves are alternate. A rapidly 
spreading invasive weed. R. Preston (personal communi- 
cation to B. Ertter, 1-2000) reports it is now moving into 
the Central Valley as scattered individuals on most major 
highways leading inland from the San Francisco Bay 
Area. 

Emilia fosbergii D.H. Nicolson: DIST: SCo: CS: GH/ 
C: DOC: San Diego Co.: Weed in container grown nurs- 
ery stock. Pleasant Knoll Rd., Valley Center. Feb. 14, 
2000, P. Nolan s.n. (CDA): NOTES: Has also been found 
as a rare nursery weed in Sacramento Co. (ScV) probably 
in imported soil. Potential garden weed. 

Emilia sonchifolia (L.) DC. ex Wight: DIST: SnJV: 
CS: GH/C: DOC: Tulare Co.: adventive under green- 
house bench; Terra Bella. T23S, R27E, Sec. 12, MD. May 
28, 1987. R.D. Harris s.n. (CDA): NOTES: Potential gar- 
den weed. 

Grindelia papposa Nesom & Suh: DIST: SCo: CS: 
NCI: DOC: Ventura Co.: along RR in Ventura. 1962, 
H.M. Pollard s.n. (CAS, SBBG): NOTES: Reported in 
Smith, C.F, (1976, pg. 291) as Haplopappus ciliatus 
(Nutt.) DC., but excluded from the 2nd ed. (Smith, C.F, 
1998) and perhaps not persisting. 

Helianthus tuberosus L.: DIST: CCo: CS: C: DOC: 
Alameda/Contra Costa Cos.: single individuals on both 
north and south banks of Cerrito Creek near end of Yo- 
semite Avenue at foot of Albany Hill. Sept. 30, 1995, B. 
Ertter 14526 (UC): NOTES: Source is presumably an up- 
stream creekside planting. 

Helichrysum petiolare Hilliard & B. L. Burtt.: DIST: 
CCo: CS: NW: DOC: Marin Co.: Matt Davis Trail above 
Stinson Beach, S side of Mt. Tamalpais, shade of forest, 
locally common in patches. July 25, 1992, B. Ertter and 
L. Fujii 11260 (JEPS); 200 m N of Panoramic Hwy, E of 
Stinson Beach, 200 m from the SW border of Mt. Tam- 
alpais State Park, growing near the remains of an old 
homestead, nr a larger population of plants 1 km distant. 
July 2, 1997, J. Randall s.n. (DAV); Monterey Co.: Open, 
sunny, sandy soil. Del Monte Forest on the edge of the 
1959 burn, Monterey Peninsula. Growing through and up 
above manzanitas and other shrubs. Jan. 3, 1970, B. F. 
Howitt 3117 (CAS, CDA). 

Heliomeris multiflora Nutt. var. multiflora: DIST: 
SNE: CS: TEN: DOC: Mono Co.: Mammoth, vacant lot 
near the Post Office. Aug. 2, 1998, D.W. Taylor 16936 
(UC). Determined by John Strother: NOTES: Apparently 
escaping from nearby areas seeded for ‘wildflowers’. The 


78 MADRONO 


plants reseed in unmanaged, ruderal vegetation in the de- 
veloped portion of town, and in this setting reseed as an- 
nuals. Heliomeris multiflora var. nevadensis (Nelson) Ya- 
tes, a native perennial, occurs in the White Mountains to 
the east. Brought to our attention by D.W. Taylor. 

Helipterum roseum (Hook.) Benth.: DIST: SnFrB: 
CS: NCI: DOC: San Mateo Co.: McClintock, E., et al. 
(1990, pg. 79). 

Lasiospermum bipinnatum (Thunb.) Druce: DIST: 
SCo: CS: NW: DOC: Santa Barbara Co.: Ross, T. and 
S. Boyd (1996, 433-434). 

Leontodon muelleri (Schultz-Bip.) Fiori: DIST: ScV: 
CS: NCI: DOC: Glenn Co.: in alfalfa field, 6th and Wy- 
oming Aves, NE of Orland. T22N, RO2W, Sec. 08, MD. 
May 2, 1982, G. Stenlund s.n. (CDA). 

Oncosiphon piluliferum (L.f.) Kallersjo: DIST: SCo: 
CS: N: DOC: Riverside Co.: Sanders, A.C. (1996, pg. 
528); Moreno Valley, E of Lake Perris along JFK Blvd. 
in disked field. March 23, 1998, R. Noll s.n. (OBI, SD). 
Det. by D. Keil, 11-2001; San Jacinto Wildlife Area, at 
headquarters off Davis Rd., north of Lakeview. Lakeview 
7.5’ quad. 33°52'N, 117°07'W; T03S, RO2W Sec. 32, SB). 
Elev. 442 m/1450 ft. July 15, 2001, O.F. Clarke s.n. 
(UCR); San Jacinto Wildlife Area, Lovell Unit approxi- 
mately 885 m east of Davis Rd and 76 m north of the San 
Jacinto River levee. Lakeview 7.5’ quad, TO4S, RO2W, 
Sec. 05, SB. El. 433 m. Alkali playa with Rumex, Crypsis 
schoenoides, Phalaris minor, Atriplex argentea, etc. Trav- 
er loamy fine sand, saline alkali. June 18, 1995, D. Bram- 
let 2434 (UCR); San Jacinto Wildlife Area, approximately 
920 m NW of Lakeview and 487 m west of the Davis Rd, 
46 m N of Marvin Rd. Perris 7.5’ quad. TO4S, RO2W, Sec 
06, SB. Elev. 433 m. Alkali playa with Plagiobothrys lep- 
tocladus, Crypsis schoenoides, Cressa truxillensis, etc. 
Willows silty clay. May 6, 1992, D. Bramlet 2265 (UCR); 
Romoland, on ramp to Hwy 215 from Hwy 74, just east 
of Hwy 215. Perris 7.5’ quad. 33°45’08"N, 117°11'06"W. 
Elev. 434 m/1425 ft. Uncommon on disturbed roadside, 
common in abandoned factory yard across 74 to the south. 
Also scattered along Hwy 215 all the way to Riverside. 
May 7, 2001, A.C. Sanders 24176, with Mitch Provance 
and T.B. Salvato (UCR); Val Verde, between Moreno Val- 
ley and Perris, along Hwy 215, 0.9 mi S of Oleander Ave., 
at S end of Patterson Ave. Steele Peak 7.5’ quad. 
33°50'47"N, 117°15’05"W, TO04S, RO4W, Sec. 01, SB. El. 
457 m/1500 ft. Weedy roadside on disturbed agricultural 
plains. Locally common on side of freeway, conspicuous 
for ca. 100 m. May 8, 2001, A.C. Sanders 24209, with 
Mitch Provance and T.B. Salvato (UCR); Moreno Valley, 
along I-215 just south of the March Field Museum, 0.7 
mi S of Van Buren Blvd. Riverside East 7.5’ quad. 
33°52'35"N, 117°15'52”W; TO3S, RO4W, Sec. 26, SB. EI. 
465 m/1525 ft. Weedy roadside on disturbed plains. Lo- 
cally common and conspicuous for ca. 1 km. May 8, 2001, 
A.C. Sanders 24210, with Mitch Provance and T.B. Sal- 
vato (UCR); Lakeview Mtns., Pulsar View Rd, ca. 1 air- 
mile NE of Juniper Flats Rd. and ca. %4—1 mi by road from 
the base of the hills. Lakeview 7.5’ quad, 33°49'06’N, 
117°05'W; T04S, RO2W, Sec. 15, SB. Elev. 610 m/2000 
ft. Chaparral, burned within the past few years. Many na- 
tive wildflowers and some non-natives, all on NW-facing 
slope. Solitary ind. growing in open spot on the burn. Apr. 
11, 1997, B. Pitzer 3121 (UCR): San Diego Co.: San 
Diego Wild Animal Park, off Hwy 78, E of Escondido in 
San Pasqual Valley. 33°06'N, 116°59’'W. Elev. 300 m/984 
ft. Weedy area near back entrance to park. Few plants this 
year, previously common. Apr. 1998, Robert Noll, s.n. 
(UCR); San Diego Wild Animal Park, along back road 


[Vol. 49 


behind exhibits (N of African Plains area); also seen on 
E edge of park and in S African section of Park Botanical 
Garden. 33°06’N, 116°59'W. Scattered to fairly common 
at edges of park; abundant among Aloes in S African sect. 
of Bot. Gdn. Elev. 300 m/984 ft. Mar 25, 1997, Jan Beyers 
s.n. (UCR): NOTES: Original report by Sanders was as 
the synonym Matricaria globifera (Thunb.) Fenzl in Harv. 
and Sond. Comment on the label of the Noll specimen 
above; “‘reported also from Orange Co., Haul Cyn. Rd., 
Irvine Ranch.”’ The current status of this population is not 
known. 

Osteospermum fruticosum (L.) Norl.: DIST: SCo: CS: 
NW: DOC: Los Angeles Co.: Zuma Beach area, mouth 
of Zuma Creek, E end of the County Beach. major veg- 
etation/habitats are small Salix lasiolepis stand w/under- 
story of escaped ornamentals; Typha/Scirpus marsh; rem- 
nant coastal dunes; disturbed roadsides, former parking 
area, and rubble dumping area. Site proposed for habitat 
enhancement. Occasional nr. stream. Point Dume 7.5’ 
quad. 34°01’N, 118°49'W. Elev. <25 ft. Mar. 12, 1997, 
S.D. White 4738 (UCR); Riverside Co.: Mockingbird 
Cyn. area, south of Van Buren Blvd. and ca. % air mi E 
of Mockingbird Cyn. Rd Riverside West 7.5’ quad. T03S, 
ROSW, Sec. 27, SB; 33°52'N, 117°23'W. Willow riparian 
with disturbed coastal sage scrub adjacent. May 8, 1989, 
Ed LaRue s.n. (UCR); NW Palomar Mountains, Agua Tib- 
ia Mountains; NW foothills of Dorland Mtn.: S end of 
Los Caballos Road, UC Emerson Oaks Reserve, at the 
Emerson Cottage. TO8S, RO2W, Sec. 24, SB. Elev. 1720 
ft. Localized escape in Carpobrotus plantings from culti- 
vated plants around the Emerson Cabin. May 2, 1996, 
Darin L. Banks 0953, with E.H. Banks (UCR); San Luis 
Obispo Co.: Toro Canyon Rd., E of U.S. Hwy 101, just 
north of Cayucos. Disturbed soil in farmland. El. 400 ft. 
Aug. 6, 1989, G. Helmkamo s.n. (UCR); Santa Barbara 
Co.: Ferren, W. R., Jr. (1985, pg. 236): NOTES: The 
“freeway daisy”’ of commerce. Similar to O. ecklonis, but 
growth spreading rather than upright. 

Pericallis cruenta (DC. non Roth) Webb. & Berth.: 
DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, 
J.T. et al. (1958, p. 149); Thomas, H. (1961, p. 374); San 
Mateo Co.: McClintock, E., et al. (1990, pg. 83): 
NOTES: Cited reports are as Senecio cruentus DC. non 
Roth. 

Porophyllum ruderale (Jacq.) Cass.: DIST: SCo: CS: 
C: DOC: Orange Co.: Weedy areas along roads surround- 
ing vegetable crop field adjacent to Seal Beach Naval 
Weapons Station. Oct. 17, 2000. P. Guerrero s.n. (CDA): 
NOTES: First report for California of this widespread 
weedy plant, but probably casual as an escape from cul- 
tivation. Native (apparently) from Arizona, New Mexico, 
Texas to South America. Our material referable to the var. 
macrocephalum (DC.) Crong. (P. macrocephalum DC.). 
Apparently used medicinally and perhaps cultivated in 
southern California for that purpose. 

Ratibida columnifera (Nutt.) Wooton & Standl.: 
DIST: CaRH, SCoRI: CS: NCI: DOC: Lassen Co.: 
Growing in dry, rocky soil on the west side of Eagle Lake, 
two miles N of the Eagle Lake Resort on service road 
33NO1. July 21, 1969, N. Santamaria 721 (CDA, CHSC); 
In re-growth vegetation in a cleared area at the north end 
of the Glenn-Eagle airstrip, southwest corner of Eagle 
Lake, 0.8 mi north of Eagle Lake Resort. T31N, RIOE, 
Sec. 04, MD. Aug. 20, 1974, G.D. Barbe 1939 (CDA, 
DAV); Riverside Co.: Murrieta, Warm Springs Creek, ca. 
¥% mile south of Murrieta Hot Springs. 33°35’N, 117°08’'W, 
TO7S RO3W, SB. Elev. 1500 ft. June 8, 1995, G.R. Ball- 
mer and K.T. Stockwell s.n. (UCR), det. by A.C. Sanders, 


2002] 


2002.; June 5, 1996, K.T. Stockwell s.n (UCR); Jurupa 
Mountains: Glen Avon Heights, on the N side of Conning 
St. between Campbell and Lindsay Sts. Fontana 7.5’ quad. 
TO2S, RO6W, Sec. 03, SB. 34°01'35’N, 117°29'45’W, 
Elev. 860 ft Sandy alluvial slopes. Organic crust which 
was observed at this location two years ago is no longer 
present. Solitary perennial. July 30, 2000, M. Provance 
2173 (UCR): San Benito Co.: San Benito. July, 1925, 
Mrs. J.A. Bettys s.n. (JEPS). 

Senecio squalidus L.: DIST: SnFrB: CS: NCI: DOC: 
Alameda Co.: volunteer plants abundant in an unculti- 
vated corner of the experimental growing grounds, U.C. 
Botanic Garden, Strawberry Canyon, Berkeley. Aug. 20, 
1971, T.C. Fuller 19927 (CDA, DAV); north of corpora- 
tion/greenhouse area, UC Botanic Garden, Strawberry 
Canyon, Berkeley. May 17, 1979, G.D. Barbe 2511 
(CDA). 

Tragopogon hybridus L.: DIST: CCo: CS: NCI: DOC: 
Monterey Co.: few plants as waifs, vacant field, Schilling 
Company, Salinas. T15S, RO3E, Sec. 3, MD. May 31, 
1978, J. Lyons and B. Oliver s.n. (CDA). 

Tripleurospermum maritimum (L.) W. D. J. Koch: 
DIST: NCo: CS: TEN: DOC: Mendocino County: Man- 
chester, at the west end of town on gravel road to the 
beach, near north edge of Lagoon Creek T13N R17W, Sec 
25, MD. Pt. Arena 7.5’ quad. El. 80 ft, 39°58'02’N; 
123°42'08"W. Dominating a storage area for dairy silage. 
June 30, 2000, D. W. Taylor 17473 (JEPS): NOTES: 
=Matricaria maritima L. Brought to our attention by 
D.W. Taylor, det. by J. Strother (UC). 


Balsaminaceae 


Impatiens balfouri J.D. Hook.: DIST: n CCo, NCo, ne 
sCo, SakrB: CS: N: DOC: Wilken, D- (1993, pg. 2): 
NOTES: Included in supplement to Munz but accidentally 
deleted from Jepson Manual manuscript. 

Impatiens noli-tangere L.: DIST: n NCo, SnFrB: CS: 
N: DOC: Wilken, D. (1993, pg. 2) also: Alameda Co.: 
Strawberry Creek just above Haas recreation facility be- 
hind UC-Berkeley, local colony in moist shade, Aug. 29, 
1996, B. Ertter 15271, (UC): NOTES: Included in sup- 
plement to Munz (as /. occidentalis) but accidentally de- 
leted from Jepson Manual manuscript. 


Berberidaceae 


Berberis darwinii Hook.: DIST: CCo: CS: NCI: DOC: 
San Francisco Co.: Howell, J.T. et al. (1958, pg. 73): 
Thomas, H. (1961, pg. 172): NOTES: Local escape from 
cultivation. 


Betulaceae 


Alnus cordata (Loisel.) Duby: DIST: NCoRO: CS: C: 
DOC: Sonoma Co.: Best, C., et al. (1996, pg. 87): 
NOTES: Establishing from landscape planting. 


Bignoniaceae 


Campsis radicans (L.) Bureau: DIST: deltaic GV: CS: 
TEN: DOC: Contra Costa Co: Antioch Marina, edge of 
marsh along roadside. July 11, 1998, B. Ertter and W.A. 
Morosco 16370 (UC); Sacramento Co.: brushy roadside 
along Hwy 160 approx. 1 mile S of Freeport and immed. 
N of Freeport Marina. Twining in Vitis californica, Rubus 
discolor. Not obviously persisting from cultivation. June 
9, 2000, G.F. Hrusa 15440 (CDA, DAV): NOTES: A 
scattered escape from cultivation, possibly marginally nat- 
uralized? 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 79 


Catalpa bignonioides Walter: DIST: n SNE SCo. ScV. 
SnJV: CS: NW: DOC: Amador Co.: Forster Ranch, along 
Dry Creek, just east of the Sacramento and San Joaquin 
Co. lines, 38°19’N; 121°00'W, elev. 50 ft, naturalized. 
June 6, 1990, L. LaPré s.n. (UCR), det. by A.C. Sanders: 
Fresno Co.: Kings River area, east of Centerville, just S 
of State Hwy 180 (Kings Cyn Rd.), along river, riparian 
forest and adjacent annual grassland, solitary tree ca. 4 m 
tall at forest margin near road, perhaps a persisting orna- 
mental planting, though there is no sign of a historic build- 
ing, 36°43'30"N; 119°28'00’W, T14S, R23E, Sec. 09, MD, 
elev. 350—400 ft. Apr. 3, 1997, S. White 4871 (UCR), det. 
by A.C. Sanders; Kern Co.: Bakersfield, E side of the 
canal at E end of Hart Memorial Park, in a dense thicket 
of willows, obviously spontaneous. April 25, 1959, E. 
Twisselman 5085 (DAV): Ventura Co.: Spontaneous in 
ditch on SPRR ROW at Loma Dr. crossing, Ojai Valley. 
May 29, 1971, H.M. Pollard s.n. (CAS, CDA): NOTES: 
Plants of uncertain status (probably persisting from culti- 
vation) are occasionally found in moist areas in western 
Riverside Co. (e.g., French Valley S of Winchester and 
San Jacinto River above Cranston Guard Station): Plants 
of uncertain status observed by Hrusa in Sacramento Co. 
along the American River at American River Parkway and 
in Yolo Co. along Sacramento River 8 mi S of Woodland. 
Expected elsewhere in low elevation riparian habitats. 


Boraginaceae 


Echium lusitanicum L.: DIST: CCo, NCo: CS: TEN: 
DOC: Monterey Co.: Rt 68, 1.5 miles south of Pacific 
Grove, occasional on roadside, May 22, 2000, G. Leppig 
1380 (CDA, HSC): San Mateo Co.: common on Rt. 101 
roadside and coastal scrub between Montara and Linda 
Mar. May 24, 2000, G. Leppig 1384 (CDA, HSC): 
NOTES: Also observed by Leppig as occasional on Rte. 
1 in Mendocino Co. and on banks of the Gualala River 
in Gualala (Sonoma Co.). 

Pentaglottis sempervirens (L.) Tausch ex Bailey: 
DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell. 
iE ctoak (1958; po: 11S): Thomas, Hoole ps 2388): 
NOTES: Reported as Anchusa sempervirens L. 


Brassicaceae 


Brassica fruticulosa Cyrillo: DIST: SCo, SnFrB: CS: 
N: DOC: Los Angeles, Riverside, San Bernardino Cos.: 
Sanders, A.C. (1996, pp. 523-524); San Mateo Co.: 
McClintock, E., et al. (1990, pg. 90). 

Cardamine flexuosa With.: DIST: CCo, SCo: CS: N: 
DOC: San Diego Co.: Vincent, M.A. (1997, pp. 305— 
306): San Francisco Co.: In plantings nr. Fisherman’s 
Wharf. April 8, 1998, M.A. Vincent and E.H. Fried 8186 
(DAV): NOTES: Most common as a greenhouse/nursery 
weed. 

Coincya monensis (L.) Greuter & Burdet: DIST: 
NCo: CS: NW: DOC: Humboldt Co.: Lot ESE of Manila 
Community Services District sewage pump station: dune 
sand mixed with gravel fill. Disturbed empty lot with Se- 
necio vulgaris, Lupinus arboreus, Holcus lanatus, Ra- 
phanus sativus, Eriogonum latifolium. Eley. 20’. 40°51'N; 
124°10'W. Feb. 13, 1997, J. Belsher 2 (CDA, DAV, HSC): 
NOTES: Under eradication. Confirmed still present in 
1999 (A. Pickart, personal communication). 

Iberis umbellata L.: DIST: CCo: CS: C: DOC: San 
Francisco Co.: Howell, J.T. et al. (1958, pg. 77). Thomas, 
H. (1961, pg. 185). 

Rorippa sylvestris (L.) Besser: DIST: SCo, SnJV: CS: 
TEN: DOC: San Diego Co.: Nursery property, Sidonia 


80 MADRONO 


Rd., Encinitas Ca. Present for several years in this site. 
May 22, 1998, C. Elmore s.n. (CDA, DAY); loc. cit. June 
28, 1998, J. Blasius s.n. (CDA): NOTES: All known sites 
under eradication. Also confirmed from commercial nurs- 
ery properties in San Joaquin Co. (Lodi), June, 1998; 
Ventura Co. (Oxnard), Jan., 2000. but submitted material 
not adequate for vouchering. Reproduction by root sprouts 
only but plants highly persistent; introduced and spread 
via contaminated nursery stock. 


Cabombaceae 


Cabomba caroliniana A. Gray: DIST: SnJV: CS: NW: 
DOC: Contra Costa Co.: Screen trap at Clifton Court 
Forebay, head of California Water Project. TOIS, RO4E, 
Sec. 35, MD. Oct. 18, 2000, R. Gage s.n. (CDA); San 
Joaquin Co.: Disappointment Slough NW of Stockton, 
TO2N, ROSE, Sec. 09, MD. Sept. 28, 1988; Griffin et al. 
s.n. (CDA); loc. cit. Oct. 6, 1995, Griffin, Finley s.n. 
(CDA); S edge 14 Mile Slough, abundant, with Egeria 
densa, Myriophyllum spicatum. TO2N, ROSE, Sec. 23, 
MD. Sept. 19, 1991, E. Finley, R. Villareal s.n. (CDA); 
Middle River about Bullfrog Marina. July 24, 2001, F. 
Maly s.n. (CDA): NOTES: Reported as common in Lew- 
iston Lake, Trinity Co. (DiTomaso personal communi- 
cation), but no confirming specimen or other plant mate- 
rial has been seen. Recognized by its submerged, deeply 
divided fan-shaped foliage leaves on distinct petioles. 
Emersed leaf-like bracts subtending inflorescences at the 
water surface are oval-perfoliate. Can fill at least a 3 meter 
water column. A purple-foliaged form is known and al- 
though not yet reported for California it is sold in the 
aquarium trade and is expected. Present in Disappointment 
Slough since at least 1980 [L. Anderson (USDA) personal 
communication]. Apparently spreading rapidly in the Sac- 
ramento Delta and ultimately to be expected widely. Vi- 
sual reports need verification as this species has been con- 
fused on cursory observation with Ranunculus aquatilis 
from which it differs in its perennial habit and petiolate 
fan-shaped leaves. 


Campanulaceae 


Campanula medium L.: DIST: SnBR: CS: C: DOC: 
San Bernardino Co.: San Bernardino Mtns, N side of 
Baldwin Lake, at Big Bear Landfill. Former pinyon-juni- 
per woodland, now disturbed and cleared; vegetation 
weedy except at lower edge, where it is Chrysothamnus 
nauseosus scrub. Solitary plant. Big Bear City 7.5’ quad., 
34°18'40’N; 116°49'00"W; TO3N, RO2E, Sec. 30 and Sec. 
31, SB. July 3, 2000, J. Wear s.n. (UCR), det. by A.C. 
Sanders. 

Lobelia erinus L.: DIST: SCo: CS: C: DOC: Marin 
Co.: Howell, J.T. (1970, p. 357); Santa Barbara Co.: 
Escape from cultivation on Cold Spring Road south of 
and near La Paz Road intersection, Montecito. June 16, 
1965, H.M. Pollard s.n. (CDA, RSA, SBBG). 


Caprifoliaceae 


Leycesteria formosa Wallich.: DIST: CCo: CS: NCI: 
DOC: San Francisco Co.: Howell, J.T. et al. (1958, p. 
130-131); Thomas, H. (1961, p. 327): NOTES: Reported 
as a local escape from cultivation. 

Viburnum tinus L.: DIST: CCo: CS: NCI: DOC: San 
Francisco Co.: Howell, J.T. et al. (1958, p. 131); Thomas, 
H. (1961, p. 327): NOTES: Local escape from cultivation. 


[Vol. 49 


Caryophyllaceae 


Silene pseudatocion Desf.: DIST: CCo: CS: NCI: 
DOC: Monterey Co.: Howitt, B.E and J.T. Howell, 
(1973, pg. 14); road shoulder, old road of Hwy 1, Marina, 
Calif. March 13, 1970, A. Allison s.n. (CAS, CDA), det. 
by J.T. Howell, 6-4-1970. San Mateo Co.: Pacifica prop- 
erty, Redwood City. Mar. 9, 1961, A. Jillson s.n. (CDA), 
det. by N. van Kleeck, 3-1971; San Francisco Co.: Single 
plant, obviously spontaneous, in neglected weedy garden, 
San Francisco. April 19, 1972, J.T. Howell 48673 (CAS, 
CDA, DAY). 


Celastraceae 


Maytenus boaria Molina: DIST: SnFrB: CS: NW: 
DOC: Alameda Co.: Gwinn Canyon in Oakland Hills, 
north side of Marlborough Drive, regrowth of scrub after 
major fire. June 21, 1994, B. Ertter 12838a (UC); loc. cit. 
Aug. 29, 1997, E. Leong s.n. (UC): NOTES: A well- 
known ornamental small tree with drooping branches, nar- 
rowly rhombic, toothed leaves, and inconspicuous flowers. 
Considered a pest plant locally, with control efforts un- 
derway. Germinates profusely following fire. 


Chenopodiaceae 


Atriplex muelleri Benth.: DIST: DSon: CS: NCI: 
DOC: Riverside Co.: 20 miles west of Blythe, a single 
plant on roadside. Oct. 17, 1965, J.C. Roos s.n. (COLO, 
UC, UCR): NOTES: Perhaps extirpated, sporadic search- 
es over past 10+ years by Sanders have not found this. 

Chenopodium watsonii A. Nels.: DIST: SnGB: CS: 
NCI: DOC: Los Angeles Co.: Big Tujunga at Colby 
Ranch Rd. Elev. 3200 ft. Dry sunny sandy roadside. Oct. 
1, 1966, L.C. Wheeler 8941 (CDA, RSA): NOTES: De- 
termination by S.E. Clemants (BKL) and S. Mosyakin 
(KW), 3-2001. Original determination as C. album L., dis- 
tinguished readily by the whitened reticulate seed coat of 
C. watsonii. Native in the Great Basin region, further col- 
lections and distribution data are necessary to determine 
if this is introduced or an overlooked California native. 

Salsola kali L. subsp. pontica (Pallas) Mosyakin: 
DIST: DMoj, s ChI: CS: NCI: DOC: Kern Co.: Mouth 
of Red Rock Cyn. Plant yellow-green, growing in shady 
probably sub-alkaline soil in dry wash. Occasional—com- 
mon in such places. Elev. 2300 ft. Creosote bush associ- 
ation. Aug. 22, 1961, E.C. Twisselmann 6468 (DAV). Det. 
by S. Mosyakin 3-2001. Ventura Co.: Mosyakin, S.L. 
(1996, pg. 389): NOTES: Ventura County record is a sin- 
gle collection from San Nicolas Island. U.S. Naval Radio- 
logical Defense Laboratory, near road above sand spit at 
100 feet elevation, R. E. Foreman 42 (US). Label data 
provided by S. Mosyakin (KW). 


Cistaceae 


Halimium lasianthum (Lam.) Spach: DIST: SCo: CS: 
NCI: DOC: Ventura Co.: escaped in yards in Oxnard. 
June, 1949, V. Holmer s.n. (CDA): NOTES: Det. by M.K. 
Bellue. 


Convolvulaceae 


Calystegia silvatica (Kit.) Griseb. subsp. disjuncta 
Brummitt: DIST: NCo, SnFrB: CS: N: DOC: Alameda 
Co.: Codornices Creek at southwest corner of Albany Vil- 
lage. May 22, 2000, M. Hurlbert s.n. (JEPS); Humboldt 
Co.: Growing in waste area at the intersection of 11th and 
B Sts, Arcata. Moist gulch. Aug. 10, 1976, T. Nelson 3166 


2002] 


(CDA); Arcata, 11th and B. St., empty lot. July 8, 2001, 
G. Leppig 1577 (CDA, HSC); Eureka, Waterfront Dr. 
waste area, 100 m N Humboldt County Library. July 10, 
2001, G. Leppig 1578 (CDA, HSC); Arcata, South G St. 
adj. to city of Arcata Corporation Yd. Waste area, road- 
side. July 11, 2001, G. Leppig 1579 (HSC, UC); Marin 
Co.: Brummitt, R.K. (Madrono, in press): NOTES: Pre- 
viously confused with C. sepium subsp. limnophila 
(Greene) Brummitt, but readily distinguished by its larger 
flowers (5—7.5 cm), inflated bracteoles that hide the calyx, 
larger leaves, and glabrous vestiture. Sporadic but persis- 
tent in urban waste areas around Humboldt Bay. 

Convolvulus tricolor L.: DIST: CCo: CS: NCI: DOC: 
Monterey Co.: adventive, vacant field adjacent to Schil- 
ling Company processing plant, Salinas. T15S, RO3E, Sec. 
03, MD. April 24, 1979, J.L. Johnson and B. Oliver s.n. 
(CDA), det. by G.D. Barbe, April 1979 at CAS. 

Dichondra micrantha Urb.: DIST: n SNE SCo, Sn- 
FrB: CS: NCI: DOC: Butte Co.: Heavy infestation, 1 
acre, | mi E of Quincy Rd. and | mi S of Middle Fork 
Feather River on W side of Bidwell Mtn. Private property. 
June 7, 1963, W. Hansell s.n. (CDA), det. by T.C. Fuller; 
Los Angeles Co.: El Segundo Dunes, W of Los Angeles 
Int. Airport and Pershing Dr. 33°56'N, 118°26’W. Elev. 
125 ft. Sept. 18, 1987, A.C. Sanders 7367 (UCR); Riv- 
erside Co.: Hemet, SE corner of State St. and Bibbel, 
1712 ft hill at east end of Diamond Valley. Hemet 7.5’ 
quad. 33°42'N, 116°58'’W; TO5S, RO1W, Secs. 26, 27, 34, 
35, SB, common corner. Elev. 494-518 m/1620—1700 ft. 
Scarce at margins of dried pool and in disturbed soil. May 
3, 2001, A.C. Sanders 24113 (UCR); San Bernardino 
Co.: San Bernardino Mtns., E of Yucaipa, Water Cyn., 
trib. of Wildwood Cyn. from the N. Vicinity of old Hunt 
Ranch, ca. % mi. N of Wildwood Canyon Rd. (T02S 
ROIW Sec. 04, SB. 34°01'30"N, 116°59'30’W) Elev. 3300 
ft. Fairly common on roadside at edge of barren cattle 
corral where clearly naturalized. June 5, 1992, A.C. Sand- 
ers 12365, with E.J. Lott and D. Pendleton (UCR); Ven- 
tura Co.: Flood plain of Coyote Creek nr. confluence with 
Ventura River, Foster Park. Oct. 15, 1969, H.M. Pollard 
s.n. (DAV): NOTES: Also noted on embankment of Cer- 
rito Cr., spreading beyond lawn of Creekside Park (Con- 
tra Costa Co.). Expected elsewhere. Cited in Munz 
(1974, pg. 379) as Dichondra repens Forst. & Forst.f. 
which is misapplied in California to D. micrantha, the 
common cultivated lawn substitute. Dichondra repens has 
also been misapplied in California to the native Dichondra 
donnelliana Tharp & Johnston, thus care must be taken, 
specifically a specimen observed, when applying an epi- 
thet to a reported occurrence identified as D. repens. 
Again, the importance of documenting voucher specimens 
is clear; in this case the holotype of Dichondra donnelli- 
ana was originally determined as D. repens! 

Ipomoea aquatica Forssk.: DIST: ScV: CS: C: DOC: 
Sutter Co: Weed in cultivated, diverse, vegetable row 
crop. Wet. Mung garden N of Yuba City. Oct. 1, 2001, 
G.F. Hrusa 15989 (CDA): NOTES: Becoming widely 
(and illegally) cultivated as a greenhouse crop in Califor- 
nia, less commonly as a row crop in the Central Valley, 
with weedy occurrences thus expected to increase in fre- 
quency. A potential aquatic pest in warm areas. Federal 
Noxious Weed. 

Ipomoea lacunosa L.: DIST: CCo: CS: C: DOC: San- 
ta Cruz Co: Aptos, watershed of Aptos Creek adjoining 
Nisene Marks State Park, weed along 3000 block of Red- 
wood Drive; elev. 550 ft, 37°00’45’”N; 121°54’00"W. Dis- 
turbed opening of residential yard in Sequoia sempervi- 
rens-Lithocarpus densiflorus-Quercus parvula vat. shrevei 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 81 


dominated forest. Waif from birdseed originating from 
bird feeder. With waif Panicum sp. and Helianthus an- 
nuus. Sept. 6, 2000, D.W. Taylor 17585 (JEPS): NOTES: 
Perhaps trivial, but to be expected elsewhere. Originated 
from ‘National Aububon Society Superior Wild Bird 
Food’ [4.53 kg—Wagner Bros. Feed Corp.] purchased ca. 
spring, 2000. Native to eastern North America. The iden- 
tity of the Panicum sp. is unknown because it does not 
[has not] flowered. 

Ipomoea quamoclit L.: DIST: GV: CS: C: DOC: Sac- 
ramento Co: volunteer in residential garden, climbing on 
roses, Sacramento. April 17, 1992, Joe Bandi s.n. (CDA). 


Crassulaceae 


Crassula multicava Lem.: DIST: CCo: CS: NW: 
DOC: Monterey Co.: Pt. Lobos State Reserve. Rock 
crevices along the trail at Big Dome Cove, granite rock 
outcrops in Pinus radiata forest, with abundant Polypo- 
dium calirhiza and lichens, 36°32'01"N; 120°56'50’W, 
Feb. 2, 1998, D.W. Taylor 16282 (JEPS, UC), det. by 
Dean Kelch (UC); San Mateo Co.: McClintock, E., et al. 
(1990, pg. 102): NOTES: Reproducing by bulbils. This 
species was reported for Monterey County by Yadon 
(1995) but was not included in Matthews, M.A. (1997). 
Report and data provided by D.W. Taylor. 

Sedum album L.: DIST: c SNE SnFrB: CS: N: DOC: 
Tuolumne Co.: Yosemite Valley, Yosemite National Park, 
ruderal disturbed margin of paths and parking area in em- 
ployee housing tents on southeast edge of Yosemite Vil- 
lage area, disturbed margin of parking areas and paths in 
open sunny locations, 3900 ft, 37°44'35’N; 119°34'48"W. 
Sept. 9, 1997, D.W. Taylor 16266 (JEPS, UC): NOTES: 
Report and data provided by D.W. Taylor. Also observed 
spreading beyond cultivation into overflow basin in 
Creekside Park, El Cerrito (Contra Costa Co.). 

Sedum dendroideum Sesse & Moc. ex DC.: DIST: 
CCo: CS: NCI: DOC: San Francisco Co.: Howell, J.T. 
et al. (1958, p. 80); Thomas, H. (1961, p. 188): NOTES: 
Local escape from cultivation. 


Cucurbitaceae 


Cucumis anguria L.: DIST: SnJV: CS: NCI: DOC: 
Madera Co.: in highway right-of-way, Madera. July 
1939, M. Bellue s.n. (UC). 

Cucurbita ficifolia Bouche: DIST: SCo: CS: NCI: 
DOC: Ventura Co: Adventive on the waste ground of a 
once inhabited site betw. Ventura Ave. and So. Pac. RR 
nr. Wadstrom, Ventura Oil Fields. Nov. 27, 1968, H.M. 
Pollard s.n. (CDA). 

Cucurbita pepo L. var. medullosa Alef.: DIST: 
NCoRO: CS: C: DOC: Sonoma Co.: Best, C., et al. 
(1996, pg. 118): NOTES: The zucchini of commerce es- 
caping locally. 


Cuscutaceae 


Cuscuta reflexa Roxb.: DIST: SCo: CS: EXT: DOC: 
Los Angeles Co.: Abundant on Hedera canariensis, N 
side Biology Bldg., Cal State Los Angeles. October 23, 
1969, 7.C. Fuller 19021 (CDA): NOTES: State listed 
Noxious Weed. Now eradicated, this was the only known 
occurrence in North America north of Mexico. 


Droseraceae 


Drosera aliciae Hamet: DIST: NCoRO: CS: NCI: 
DOC: Mendocino Co.: Meyers-Rice (Madrono, in press): 
restricted to its introduction site in a wet depression in a 


82 MADRONO 


pygmy forest, 39°15'N; 123°45'W. Nov. 2, 1997, B. Mey- 
ers-Rice MR971101 (DAV): NOTES: Intentionally intro- 
duced. 

Drosera capensis L.: DIST: NCoRO: CS: NCI: DOC: 
Mendocino Co.: Meyers-Rice (Madrono, in press); sev- 
eral scattered colonies of naturalized plants spreading 
through wet depressions and seeps in a pygmy forest, 
39°15'N; 123°45'W. Nov. 2, 1997, B. Meyers-Rice 
MR971103 (DAV): NOTES: Intentionally introduced. Ac- 
cording to B. Meyers-Rice (DAV) this species is probably 
D. linearis auct. non Goldie, as per Smith, G. and C. 
Wheeler (1990-1991, pg. 170) and Hickman, J.C. ed. 
(1993, pg. 541). 

Drosera tracyi MacFarlane: DIST: NCoRO: CS: NW: 
DOC: Mendocino Co.: Smith, G. and C. Wheeler (1990— 
1991, pg. 170): NOTES: Intentionally introduced. Some- 
times treated within D. filiformis Raf. According to B. 
Meyers-Rice (DAV) both typical D. filiformis (as men- 
tioned in Hickman, ed. 1993) and D. tracyi are found at 
this site. 


Ebenaceae 


Diospyros virginiana L. var. virginiana: DIST: SnBr: 
CS: NCI Gn Mendocino Co.). NW (as clonal colonies in 
San Bernardino Co.): DOC: Mendocino Co.: Hopland 
Field Station, headquarters nr. office. June 14, 1959, AHM 
s.n. (AHUC); San Bernardino Co.: San Bernardino Mtns. 
Mill Creek Canyon, on S side of Hwy 38, 1.2 mi above 
Mountain Home Creek at Mountain Home Village, Forest 
Falls 7.5’ quad., 34°06'N; 116°58’30"W, TO1S, ROIW, 
Sec. 10, SB. Elev. 4040 ft/1232 m. Dry meadow w/scat- 
tered trees on alluvial bench in canyon bottom, sandy 
loam w/rocks. Single trees of Pyrus communis and Prunus 
cerasifera also present. Could be old orchard site, but veg- 
etation looked natural, w/no sign of former occupation; 
possibly a few spp. escaped from cultivation at Mountain 
Home. Grove of 74 trees, 2—9 m tall; oldest (75—100 yr) 
dead but 45 cm dbh, live trees 6-12 cm; corollas pale 
yellow, mostly w/4 lobes, but occasionally 5. Grove all 
female?, possibly from sprouts of old tree. Fruits pro- 
duced; but seeds? Discovered ca. 1983 by Goodman. Bark 
dark and deeply fissured into blocks, covered w/lichens. 
June 26, 1999, A.C. Sanders 22903 with John Goodman 
(ARIZ, CAS, MO, RSA, SD, UCR, UNLV, UTEP); loc. 
cit. Nov. 11, 1999, A.C. Sanders 23252 with Mihai Cos- 
tea, T. B. Salvato (UCR); Near Old Mormon Road Mon- 
ument on old loop off Hwy 18 below Crestline, San Ber- 
nardino North 7.5’ quad., TO2N, RO4W, Sec 27, SB. 
34°13'30"N; 117°17'30"W. Elev. 4200 ft/1280 m, canyon 
woodland. Scarce tree ca. 20 m tall and 38 cm dbh. Road- 
side, presumably originally planted (possibly from dis- 
carded seed?), but with a number of saplings (10—15) de- 
rived from root-sprouts surrounding parent tree. Straight 
central trunk with small angled branches, square-checked 
bark. July 21, 2000, A.C. Sanders 23591 with N. Diep 
(UCR): NOTES: The Mendocino Co. specimen at AHUC 
may or may not document a spontaneous occurrence. Oth- 
erwise only two known populations but both are repro- 
ducing vegetatively and have been present for decades. 
Plants are vigorous and obviously successful under natural 
conditions. A population with both sexes present might be 
even more successful. 


Elaeocarpaceae 


Aristotelia chilensis (Molina) Stuntz [A. macqui 
LHer.]: DIST: SnFrB: CS: TEN: DOC: Alameda Co.: 
Strawberry Creek near Life Sciences complex on UC- 


[Vol. 49 


Berkeley Campus, common shrub in understory along 
creek. May 23, 2000, B. Ertter s.n. (UC). 

Muntingia calabura L.: DIST: SnJV: CS: GH/C: 
DOC: Stanislaus Co.: Spontaneous in coco fiber imported 
from Sri Lanka, greenhouse hydroponic operation. Nov. 
11, 1997, T. Watson s.n. (CDA): NOTES: Worldwide 
weed of the tropics and wet subtropics, indigenous to S. 
America. In California known only as a greenhouse weed 
(seedlings). May be expected to volunteer and persist un- 
der mild, moist conditions, esp. cultivated sites. 


Escalloniaceae 


Escallonia macrantha Hook. & Arn.: DIST: CCo: 
CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. 
(1958, p. 81); Thomas, H. (1961, p. 192). 


Euphorbiaceae 


Euphorbia characias L.: DIST: CCo: CS: NCI: DOC: 
Alameda Co.: Albany waterfront, on former landfill, local 
colony (probably extirpated by subsequent park develop- 
ment). Aug. 19, 1994, B. Ertter 13076 (UC): NOTES: 
Single plant also noted on adjacent Albany Hill. Com- 
monly grown as an ornamental. 

Euphorbia cyathophora Murr.: DIST: SCo, SnJV: CS: 
NCI: DOC: Fresno Co.: to six feet, in roses, etc., Fresno. 
Oct. 28, 1959, T.C. Fuller 3213 (CDA); Ventura Co.: 
light infestation on roadside fill, Olsen Rd. betw. Thou- 
sand Oaks and Simi. Feb. 10, 1971, Hannah s.n. (CDA), 
det. by T-C. Fuller. 

Euphorbia dendroides L.: DIST: SCo: CS: NW: DOC: 
Los Angeles Co.: Sanders, A.C. (1997a, 203); Foothills, 
San Gabriel Mtns., Eaton Cyn nr. Kinneloa Mesa, Pasa- 
dena. TOIN, R12W, Sec. 13, SB. Mar. 30, 1981, S. Grang- 
er s.n. (CDA, RSA); Chantry Flats Ranger St., Big Santa 
Anita Cyn. Angeles Nat. Forest. May 21, 2002, J. Hart- 
man, s.n. (CDA, UC/JEPS, UCR); Santa Barbara Co.: 
Smith, C.E (1976, pg. 184); Santa Barbara, Franceschi 
Park, naturalized along rocky slope of Mission Ridge Rd. 
Sept. 5, 1950, C.F. Smith 2859 (DAV, SBBG); loc: cit 
Feb. 13, 1952, C.F. Smith 3224 (DAV, SBBG); Escaped 
on hillside, S of Franceschi Park, Santa Barbara. May 7, 
1959, T.C. Fuller 2433. (CDA, DAV); Vacant field NE of 
Old Mission Santa Barbara. June 11, 1970, C.F. Smith 
10229 (DAV, SBBG); Ventura Co.: Few scattered plants, 
Peto Seed Co. Ranch, Saticoy. May 20, 1959, 7.C. Fuller 
2435 (CDA): NOTES: Several additional collections at 
DAV from the Franceschi Park locality, all from the 
1950s, are not detailed here. 

Euphorbia heterophylla L.: DIST: ScV: CS: NCI: 
DOC: Sutter Co.: Weedy in 4 acre mung bean crop, pre- 
sent for past 2 seasons, 0.15 mi S of Nuestro Rd, W side 
Terra Buena Rd. NW Yuba City. Sept. 26, 1984, G.D. 
Barbe 4104 (CDA, DAV): NOTES: Reported also as an 
uncommon weed in the UCR (Riverside Co.) Botanic 
Garden. 

Euphorbia hirta L.: DIST: SCo: CS: N: DOC: Riv- 
erside Co.: Sanders, A.C. (1997a, 203-4): NOTES: 
Sometimes treated as Chamaesyce hirta (L.) Millsp. Has 
been intercepted as a weed of nursery stock from the 
southeastern states, esp. Florida. 

Euphorbia hypericifolia L.: DIST: CCo, ScV: CS: GH/ 
C: DOC: Monterey Co.: Town of Aromas, Blue Pacific 
Greenhouses at the corner of Carpenteria Rd. and San 
Juan Rd. Aug. 21, 2000, M. Inaba s.n. (DAV); Sacra- 
mento Co.: 12676 Stockton Blvd., uncommon ascending 
weed in greenhouse, presumably imported from Florida or 
Hawaii with foliage plants. May 23, 1988, D. Koutnik s.n. 


2002] 


(DAV): NOTES: All original determinations as Chamae- 
syce hypericifolia (L.) Millsp. 

Euphorbia marginata Pursh: DIST: KR, n SNE ScV, 
SCo: CS: NCI: DOC: El Dorado Co.: S of Camino on 
rd to Pleasant Valley, adventive in roadfill. Aug. 7, 1977, 
G.L. Stebbins 77136 (DAV); Placer Co.: Limited light 
infestation, SPRR yards, Roseville. TION, RO6E, Sec. 11, 
MD. Aug. 26, 1971, Henderson s.n. (CDA); Shasta Co.: 
Limited heavy infestation, roadside, 100 ft N of Calif. 
Forestry Station, French Gulch, Clear Ck. Cyn. T33N, 
RO7W, Sec. 02, MD. Elev. 1500’. Sept. 8, 1970, P. Whipp 
s.n. (CDA); Ventura Co.: Spontanous in waste ground, 
Junipero St. betw. Santa Clara and Main Sts. Ventura. July 
12, 1966, H.M. Pollard s.n. (CAS, CDA); spontaneous on 
ground cleared for waterfront development. Front and 
Palm Sts, Ventura. July 11, 20, 28, 1967, H.M. Pollard 
s.n. (CAS, CDA). 

Euphorbia myrsinites L.: DIST: TR: CS: NCI: DOC: 
Kern Co.: Single plant on stream bank, Vine Street, Fra- 
zier Park. TOON, R20W, Sec. 35, SB. May 14, 1981, J. 
Marks s.n. (CDA), det. by T.C. Fuller. 

Euphorbia rigida M. Bieb.: DIST: SnJV, SCo: CS: 
NCI: DOC: Tulare Co.: escaped from cultivation, RR 
ROW SE of Porterville. T22S, R28E, Sec. 06, MD. 
March, 1993, Ahrendes s.n. (CDA); Ventura Co.: 3—4 
large plants established in grassy, weedy area, Erbs Rd., 
Thousand Oaks. March 30, 1967, Schall s.n. (CDA); in 
ice plant groundcover, median highway strip along US 
Hwy. 101 ca. 3 mi E of Ventura. Feb. 27, 1976, C. Elmore 
s.n. (DAV). 

Euphorbia terracina L.: DIST: SCo: CS: NW: DOC: 
Los Angeles Co.: Sanders, A.C. (1997a, 205): Volunteer 
in UCLA Botanic Garden. Oct. 23, 1967, T.C. Fuller 
16495 (CDA); El Segundo Dunes, immed. W of LAX. 
May 18, 1988, A.C. Sanders 7832 (UCR, CDA); Solstice 
Canyon, Santa Monica National Recreation Area. TO1S, 
R18W, Sec. 16, SB. Mar. 22, 2001, S. Williams s.n. 
(CDA); Monterey Park. Garvey Reservoir, dry slope 
above dam. TO1S, R12W, Sec. 26, SB. June 26, 2001, J. 
Hartman and M. Adams s.n. (CDA); Palos Verdes Pen- 
insula, Rancho Palos Verdes, Ocean Trails development, 
between Palos Verdes Dr. South and the ocean. San Pedro 
7.5’ quad. 33°43'37"N, 118°20'30’W. Elev. 2—10 ft. Base 
of coastal bluffs. July 17, 2001, Jeremiah George s.n. 
(UCR), det. by A.C. Sanders; Zuma Beach area, mouth of 
Zuma Creek, E end of the County Beach, ca. 100—200 
plants on W bank of the stream, in sand, and on adjacent 
remnant dunes. Point Dume 7.5’ quad. 34°O1'N, 
118°49’W. Elev. <25 ft. Mar. 12, 1997, S.D. White 4750 
(UCR), det. A.C. Sanders: Santa Monica Mtns., Malibu 
Lagoon, Malibu Beach State Park, mouth of Malibu 
Creek. Malibu Beach 7.5’ quad. 34°02’N, 118°41'W; 
TO1S, R17W Sec. 32, SB. Elev. 8 m/25 ft. Fairly common 
perennial at edges of cultivated areas at Adamson house. 
Oct. 10, 1998, A.C. Sanders 22259 (UCR); same as pre- 
vious, but: uncommon at edge of road (Cross Creek) on 
W side of lagoon. Oct. 10, 1998, A.C. Sanders 22260 
(UCR): NOTES: Monterey Park form may be at least 
facultatively annual, warrants further study. 

Sapium sebiferum (L.) Roxb.: DIST: ScV: CS: NW: 
DOC: Sacramento Co.: On the N bank of American Riv- 
er in the American River Parkway, few hundred meters 
downstream of the Estates Drive entrance. 38°33'N; 
121°22’W. June 28, 1998, B. Meyers-Rice MR980603 
(CDA, DAV): NOTES: Ornamental, commonly cultivated 
in the residential areas surrounding the American River 
Parkway. Has the potential to naturalize locally in Cali- 
fornia. Reported from SnFrB, but no confirming speci- 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 83 


mens have been seen. A serious pest in the summer wet 
southeastern U.S. 


Fabaceae 


Astragalus cicer L.: DIST: c SNF: CS: TEN: DOC: 
Tuolumne Co.: Disturbed waste area nr. Standard. July 4, 
1998, M. Chambers s.n. (CDA): NOTES: Persisting in 
this site for several years previous to 1998 and observed 
inel999.- 

Cassia nemophila A. Cunn.: DIST: DSon: CS: TEN: 
DOC: Riverside Co.: Coachella Valley, between La 
Quinta and Indio, along Jefferson St. between Ave 54 and 
I-10, 33°41'N; 116°16’W, TO6S, RO7E, Sec. 04, elev. 10 
m. Fairly common 1—2 m shrub, scattered on roadside and 
in adjacent old fields, cultivated nearby in center divider 
of Jefferson St. and spreading by seed into adjacent dry 
lands. May 28, 1999, A.C. Sanders 22794 (UCR, and to 
be distributed): NOTES: Reproducing without care in one 
of the driest parts of the state. [This belongs in Senna, but 
as of 1986 the appropriate combination had not been pub- 
lished (R. Barneby, personal communication; Royal Hort. 
Dict. Gardening, 1992, says the same, but uses the Senna 
nemophila combination anyway “awaiting publication’’)]. 

Ceratonia siliqua L.: DIST: SCo: CS: NW: DOC: Los 
Angeles, Riverside, San Bernardino Cos.: Sanders, A.C. 
(1996, pg. 526). 

Coronilla valentina L.: DIST: SCo, s ChI: CS: NW: 
DOC: Los Angeles Co.: Ross, T. and S. Boyd (1996, pg. 
435). 

Dolichos lignosus Pers.: DIST: SCo: CS: NCI: DOC: 
San Diego Co.: Beauchamp, R. M. (1986, pg. 157). 

Genista monosperma (L.) Lam, non Link, nec Del.: 
DIST: SCo: CS: NW: DOC: Los Angeles Co.: San Ga- 
briel Mtns, base of range at Padua Hills, W of mouth San 
Antonio Cyn. Disturbed alluvial fan and adjacent slopes 
with chaparral. Feb. 3, 1990, S. Boyd 3828 (CDA, RSA): 
San Diego Co.: South of Fallbrook. 100 yds W of Olive 
Hill Rd, approx. %4 mi S of Mission Rd. (rd $13), S side 
of Color Spot nursery. June 19, 2000, J. Giessow s.n. 
(CDA and to be distributed): NOTES: Original det. by E. 
McClintock (CAS); label det. of Boyd 3828 as Genista 
aetnensis (Biv.) DC. Treated in Flora Europaea as Lygos 
monosperma (L.) Heywood and listed by CalEPPC as Re- 
tama monosperma (L.) Boiss. We find the generic distinc- 
tions dubious. Under eradication on adjacent Camp Pen- 
dleton federal lands (personal communication from E. 
Johnson, May 2000). Specimen cited in Rejmanek and 
Randall (1994) as at DAV apparently was never deposited. 

Gleditsia triacanthos L.: DIST: GV: CS: NW: DOC: 
Sacramento Co.: Randall, J.M. and B. Meyers-Rice 
(1997, pp. 399-400). 

Lathyrus sativus L.: DIST: CCo, NCoRO: CS: C: 
DOC: San Luis Obispo Co.: Spontaneous in garbanzo 
bean field, opposite Los Osos Valley Memorial Park Cem- 
etery, W end of Los Oso Valley. July 22, 1971, J.H. Foott 
s.n. (CDA, DAV); Sonoma Co.: Best, C., et al. (1996, pg. 
ta): 

Ononis alopecuroides L.: DIST: SCoRO: CS: NW: 
DOC: San Luis Obispo Co.: Hrusa, G.F (2000, pg. 139): 
NOTES: Known from a single large population, currently 
under eradication by San Luis Obispo Co. Agricultural 
Commissioner’s Office. 

Robinia hispida L.: DIST: ScV: CS: N: DOC: Sac- 
ramento Co.: Colony on bank of Sacramento River at 
divergence of Steamboat Slough, SW corner of Steamboat 
Bridge. Root-sprouting, spreading from planted plants 
down bank, forming a thicket beneath Robinia pseudo- 


84 MADRONO 


acacia. Apr. 9, 2000, G.F. Hrusa 15318 (CDA and to be 
distributed): NOTES: Locally naturalized, but apparently 
spreading only vegetatively. Robinia hispida consists of a 
series of clones, reproducing facultatively by rootsprouts 
and agamospermic seeds (Isely, 1998). Several forms are 
cultivated. 

Senna artemisioides (Gaudich. ex DC.) Randell: 
DIST: SnBr, SnGb: CS: N: DOC: Los Angeles Co.: San 
Gabriel Mts, north of Claremont, along Burbank Fire 
Road in Burbank Canyon, west of Palmer Canyon; grow- 
ing near edge of gravel road, naturalized in area. Jan. 24, 
1993, 7.S. Elias 12445 (UC); San Bernardino Co.: North 
of San Bernardino along Hwy 18, 0.2 miles above the 
lower end of old Waterman Canyon Road, SW side of 
highway on road fill, growing wild. Mar. 9, 1984, F.C. 
Vasek s.n. (UCR): NOTES: Population still extant ca. 
1995-1998 in lower Waterman Canyon just above Hwy 
18. All specimens labeled as Cassia artemisioides Gau- 
dich. ex DC. 

Senna obtusifolia (L.) H.S. Irwin & Barneby: DIST: 
DSon, SnJV: CS: N in DSon; TEN in SnJV: DOC: Riv- 
erside Co.: Sanders, A.C. (1996, pg. 531): NOTES: A 
single individual also found on a roadside in Fresno Co., 
sent without additional data to CDA for confirmation; ma- 
terial in condition too poor for vouchering. 

Sesbania punicea (Cav.) Benth.: DIST: CaRE GV, 
NCoRO: CS: N: DOC: Butte Co.: Shrub on wet sand, 
margin of small pond, % mile W of Pacific Heights Rd., 
Oroville Wildlife Area. T18N, RO3E, Sec. 03, MD, elev. 
140 ft. Aug. 23, 2000, L. Ahart 8660 (CDA, CHSC). Fres- 
no Co.: gravel pit ponds, Pinedale. June, 1988, J. Dun- 
nicliff s.n. (CDA); Sacramento Co.: On the margins of 
William Pond in the American River Parkway, Arden Rd. 
entrance. 38°33’N; 121°22’W. June 28, 1998, B. Meyers- 
Rice MR980604 (CDA, DAV); Shasta Co.: E side Hwy 
273 in Redding, immed. S of Breslauer Rd. Shrubs in 
wash bet. Hwy and RR tracks. Aug. 21, 2000, K. Martyn 
s.n. (CDA); Riverview Country Club, Bechelli Lane. Ex- 
tensive lakeshore infestation. Dec. 11, 2001. K. Martyn 
s.n. (CDA); Sonoma Co.: Best, C., et al. (1996, pg. 140): 
NOTES: Sonoma County report as S. tripletii Host. 

Trifolium alexandrinum L.: DIST: CCo, ScV: CS: 
NCI: DOC: Butte Co.: South side of Evans-Reimer Rd., 
ca. 1 mi E of Pennington Rd., Gray Lodge Waterfowl 
Management Area. May 10, 2001, L. Ahart 8738 (CDA, 
CHSC, UC); Monterey Co: few waifs, vacant field. Schil- 
ling Co., Salinas. T15S, RO3E, Sec. 03, MD. May 31, 
1978, J. Lyons and B. Oliver s.n. (CDA). 

Trifolium cernuum Brot.: DIST: ScV: CS: N: DOC: 
Butte Co.: Near boat ramp off Larkin Rd., Thermalito 
Afterbay, Lake Oroville. May 13, 2000, L. Ahart 8343 
(CHSC, MU, UC/JEPS): NOTES: Determination con- 
firmed by M. Vincent (MU). Reported in Oswald, V. 
(2000). Naturalization local. 

Trifolium gemellum Poir. ex Willd.: DIST: CCo, 
SnFrB: CS: N: DOC: Napa Co.: Henry Road 1.7 miles 
northwest of Dealy Lane [SW of city of Napa]. Common 
on open, grassy, southwest-facing slope in Quercus agri- 
folia woodland, elev. 450 ft, TOSN, ROSW, Sec 12, MD. 
May 12, 2000, J. Ruygt 4248 (UC): NOTES: Naturali- 
zation local. Report and data provided by J. Ruygt. 

Trifolium resupinatum L.: DIST: NCo, CCo, ScvV, 
SCo: CS: NCI: DOC: Humboldt Co.: immediate vicinity 
of Eureka, a single plant. May 14, 1896 and July 8, 1897, 
J.P. Tracy 105 (UC); Monterey Co.: few scattered plants 
in vacant field, flowers pinkish; Schilling company, Sali- 
nas. T15S, RO3E, Sec. 03, MD. May 11, 1978, J. Bunch 
and B. Oliver s.n. (CDA); Santa Barbara Co.: Howell, 


[Vol. 49 


J.T. (1972, pg. 103); Smith, C.E (1976, pg. 179); Sutter 
Co.: Edge of field, 3 miles S of Oswald. June 11, 1967, 
J.T. Howell 42556 and G.H. True (CAS, CDA); Ventura 
Co.: Howell, J.T. (1972, pg. 103): NOTES: Probably only 
casual. 

Trifolium retusum L.: DIST: CaRF: CS: NW: DOC: 
Tehama Co: Calif. Dept. of Fish and Game parcel on 
west side of Manton Road ca. 2 miles north of Dales Sta- 
tion on Hwy 36, ca. 14 miles northeast of Red Bluff, 
T29N, RO2W, Sec 26, MD. Elev. 740 ft, locally abundant 
in gravelly soil. May 20, 1996, V.H. Oswald and L. Ahart 
7613 (JEPS); loc. cit. May 21, 1998, V.H. Oswald 9087 
(JEPS): NOTES: Determination by M. Vincent (MU). 

Trifolium stellatum L.: DIST: CCo: CS: NCI: DOC: 
Monterey Co.: waif, vacant field, Schilling Company, Sa- 
linas. T15S, RO3E, Sec. 03, MD. May 31, 1978, J. Lyens 
and B. Oliver s.n. (CDA). 

Trifolium striatum L.: DIST: n SN: CS: NCI: DOC: 
Nevada Co.: Grass Valley, East Main Street at Dorsey 
Drive, waste ground at entrance to Litten Industries com- 
plex, TI6N, RO8E, Sec. 23, MD. June 3, 1980, G.D. 
Barbe and P. Hiatt 2791 (CDA, JEPS); Spring Hill be- 
tween Grass Valley and Nevada City. Elev. 2700 ft. June 
19, 1973, G.H. True 7584 (CAS, CDA); loc. cit. May 11, 
1973, G.H. True 7469 and J.T. Howell (CAS, CDA); Son- 
oma Co.: Best, C., et al. (1996, pg. 144); Sugarloaf Ridge 
State Park, ca. 1 air mi S of Red Mtn, Adobe Cyn. TO7N, 
RO6OW, Sec. 22, MD. May 21, 1996, F. Bowcutt 2141 
(DAV). 

Trifolium tomentosum Willk. ex Nyman: DIST: 
NCoRO, ScV, SnFrB: CS: NW: DOC: Contra Costa Co.: 
Mount Diablo State Park, connector between Barbecue 
Terrace Road and Wall Point Road, localized patch at edge 
of path through open oak woodland/grassland. May 3, 
1998, B. Ertter 16063 (JEPS); Napa Co.: Imola Ave. 0.2 
mile west of Suscol Ave., Napa. Grasslands, elev. 20 ft, 
TOSN, RO4W, Sec. 14, MD. April 11, 1979, J. Ruygt 501 
(JEPS); Sacramento Co.: Pasture adj. to underpass at Rio 
Linda and SW Elverta, betw. Natomas E Main drain and 
W end of ‘N’ St. TION, ROSE, Sec. 19, MD. Elev. 40 ft. 
May 12, 1992, R. York 92-002 (CDA); Overflow parking 
area, E of Cal Expo at Ethan Way, Sacramento. June 7, 
1987, N. Wymer s.n. (CDA, DAV); Sonoma Co.: Best, 
C., et al. (1996, pg. 144); Annadel State Park, east of 
Ledson Marsh, several dense patches in grassland, both 
sides of Marsh Trail, midway between intersection with 
Lawndale Trail and marsh spillway. April 19, 2000, A. 
Howald 2037 (CDA): NOTES: Resembling T. fragiferum 
in having the calyx inflated in fruit, but annual and with 
only a vestigial involucre. All specimens at CDA appear 
referable to the var. tomentosum sensu Zohary and Heller 
(1984). Easily overlooked and probably more widespread 
than the collections above indicate. 

Trifolium vesiculosum Savi: DIST: CCo, NCoRO, 
ScV: CS: N: DOC: Humboldt Co.: Disturbed road repair 
area at roadside, Hwy 101 nr Orick. TOON, ROIE, Sec. 
06, H. Sept. 14, 1998, P. Haggard s.n. (CDA); Santa 
Cruz Co.: Common on limestone mine tailings above 
Davenport. Nov. 26, 2000, G.F. Hrusa 15725 (CDA); So- 
lano Co.: SW of Davis, abandoned fields between Hwy 
113 and Pedrick Rd. July 1, 1998, M. Rejmanek s.n. 
(CDA, DAV): NOTES: A distinctive species among na- 
tive and naturalized California clovers, readily distin- 
guished by its chartaceous, inflated calyces having 25+ 
prominent longitudinal veins and similarly prominent lat- 
eral venation. Apparently an occasional component of clo- 
ver-containing hydro-seed mixtures; Humboldt Co. and 


2002] 


Santa Cruz Co. occurrences may have originated via this 
pathway. 

Trigonella corniculata L.: DIST: NCoR: CS: NCI: 
DOC: Mendocino Co.: Ukiah, in cover crop. May 5, 
1938, G.T. Nordstrom s.n. (UC). 

Trigonella foenum-graecum L.: DIST: CCo, ScV: CS: 
NCI: DOC: Monterey Co.: Volunteer in vacant field next 
to Schilling plant, Salinas. T15S, RO4E, Sec. 03, MD. 
April 24, 1979, B. Oliver s.n. (CDA); Yolo Co.: north 
edge of Davis just west of B St., weed in barley field. 
Mar. 27, 1951, J.M. Tucker 2058 (DAV, UC). 

Vicia bithynica (L.) L.: DIST: CCo: CS: NCI: DOC: 
Monterey Co.: waste area, Schilling Co., Salinas. T15S, 
RO4E, Sec. 03, MD. May 15, 1978, B. Oliver s.n. (CDA). 


Fagaceae 


Quercus ilex L.: DIST: SCo: CS: N: DOC: Los An- 
geles Co.: Claremont, adventive in plantings of chaparral 
shrubs on grounds of Rancho Santa Ana Botanic Garden, 
15 Nov. 1990, S. Boyd and T. Ross 5305 (RSA, UCR); 
Orange Co.: City of Orange, N end of Yorba St. at San- 
tiago Creek, near Chapman Ave. crossing. Orange 7.5’ 
quad. 33°47'24’N; 117°50'24’W, elev. 260 ft/79 m. Creek- 
bed, several trees to 30 ft. Feb. 1, 2000, Y. Moore s.n. 
(UCR): Riverside Co.: Weed tree in a citrus orchard in 
Rubidoux, area being cleared for houses, but this tree be- 
ing retained, Oct. 23, 1994, Donald E. Peck s.n. (UCR); 
Riverside, east side of UCR campus nr. parking lot 13, 
assoc. with Salix lasiolepis, Populus fremontii, Baccharis 
salicifolia, etc. Elev. 335 m/1100 ft. Solitary 5 m tree at 
edge of wash and parking lot. Clearly spontaneous—no 
cultivated plants in immediate vicinity. May 12, 1997, 
A.C. Sanders 20711 (UCR); Riverside, Watkins Dr. east 
of Blaine St., at Lemona Siding. Riverside East 7.5’ quad. 
33°58'30"N, 117°19'W; TO2S RO4W Sec. 20, SB. Elev. 
335 m/1100 ft. In hedge of oleander and Brachychiton 
along N side of Watkins. At least 6 young trees, mostly 
3—5 m tall, growing as weeds in hedge; plainly sponta- 
neous. May 3, 2001, A.C. Sanders 24123 (UCR). 


Geraniaceae 


Geranium columbinum L.: DIST: NCo, ScV: CS: 
NCI: DOC: Humboldt Co.: ca 7.5 road miles south of 
Ferndale, along Wildcat Road near Green Pond ranch. 
May 20, 1987, Pykala and Norris 751 (MO); Solano Co.: 
5.6 miles W of Winters. April 16, 1968, Ishizuka 19 (MA): 
NOTES: Acc. to Aedo (2000) this is a relative of G. car- 
olinianum L., native to the Old World but widely distrib- 
uted in the northeast US; also in Oregon and Washington. 

Geranium lucidum L.: DIST: SnFrB: CS: N: DOC: 
Alameda Co.: lower end Strawberry Canyon firetrail be- 
hind UC-Berkeley Botanical Garden, locally abundant in 
wet ground at edge of forest. Apr. 11, 1998, B. Ertter 
15979 (UC); loc. cit. Apr. 29, 1998, Ertter 16029 (UC). 

Geranium purpureum Vill.: DIST: NCoR, SnFrB: CS: 
NW: DOC: Alameda Co.: lower end of Strawberry Can- 
yon firetrail, behind UC-Berkeley campus. Nov. 23, 1991, 
B. Ertter and B. Olson 10891 (UC); loc. cit. Apr. 25, 1974, 
L.R. Heckard 3679 (JEPS); Albany Hill, uncommon at 
time of collection but rapidly becoming more abundant. 
May 20, 1995, B. Ertter 14216 (UC); corner of Hearst 
Ave. and Gayley Road, UC-Berkeley campus. Apr. 4, 
2001, B. Ertter and D. Norris 17574 (UC and to be dis- 
tributed); Napa Co.: Kroeber Ranch west of Rutherford, 
in weed-filled meadow. Apr. 25, 1996, B. Ertter and J. 
Ruygt 14601 (UC): NOTES: Similar to G. robertianum, 
but witn consistently smaller (<< 1 cm long), more uni- 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 85 


formly bright pink petals, yellow (vs. orange) anthers, and 
less anthocyanic foliage overall. 

Geranium pyrenaicum Burm. f.: DIST: CCo: CS: C: 
DOC: Alameda Co.: Berkeley Campus, east side of Bot- 
any building. June 15, 1914, W.L. Jepson s.n. (JEPS): 
NOTES: Current determination by Ertter, 1996; previous- 
ly determined as (and possibly basis for CCo record of) 
G. pusillum Burm.f. in Hickman (1993). 

Geranium rotundifolium L.: DIST: CCo, SCo, SnFrB: 
CS: NW: DOC: Alameda Co.: UC-Berkeley campus, in 
Grinnell Natural Area, locally abundant. June 28, 1996, 
B. Ertter 14910 (UC); Albany Hill. July 13, 1996, B. Ert- 
ter 14967 (UC); Contra Costa Co.: Tilden Regional Park 
east of Grizzly Peak Boulevard, scattered colonies along 
dirt road in weedy hillside. May 14, 1996, B. Ertter 14684 
(UC); Los Angeles Co.: Ross, T. and S. Boyd (1996, 435— 
436); San Luis Obispo Co.: Huasna Rd., 1.5 rd miles E 
of bridge over Huasna River; foothill woodland and chap- 
arral patches, area grazed. Occasional on roadside. April 
1, 1995, D. Keil 24714 (OBI); US 101 1.3 miles NW of 
Cuesta Pass, ca. 0.4 miles S of Tassajara Creek Rd.. Coast- 
al live oak woodland and adjacent roadside zone, locally 
common in roadside zone and edge of woodland under- 
story. May 27, 1999, D. Keil 28354 (OBI); Cuesta Grade 
East along Mt. Lowe Rd., ca. 2 road miles from Highway 
101, locally common under shade of Quercus agrifolia on 
bank above road. May 9, 2000, D. Keil 28648 (OBI): 
NOTES: Somewhat reminiscent of Geranium molle, but 
with entire (vs. apically notched) petals, a short awn (< 
1 mm long) on the sepals, fruits that are finely hairy rather 
than wrinkled, and less deeply lobed leaves. Noted as 
weed elsewhere in Berkeley; possibly seen by Ertter on 
Fremont Peak, San Benito Co. 

Geranium texanum (Trel.) A. Heller: DIST: CCo: CS: 
NCI: DOC: Marin Co.: Aedo (2000); Olema. June 7, 
1936, Howell s.n. (NY): NOTES: This species, another 
relative of G. carolinianum L., otherwise occurs in east 
Texas and Louisiana (and the Azores). According to Aedo 
(2000), ““Its presence in California (not previously record- 
ed) probably constitutes a[n] occasional introduction’’. 


Hamamelidaceae 


Liquidambar styraciflua L.: DIST: ScV: CS: TEN: 
DOC: Sacramento Co.: North bank of American River, 
American River Parkway, a few hundred meters downriv- 
er of the Estates Dr. entrance. 38°33’N; 121°22’W. June 
16, 1998, J. Randall s.n. (CDA, DAY). 


Hydrophyllaceae 


Wigandia caracasana HBK.: DIST: SCo: CS: NCI: 
DOC: Munz, PA. (1974, pg. 519): NOTES: No evidence 
for naturalization. Labeled specimens at CDA collected in 
the locations cited by Munz (1974), state that the plants 
were cultivated but without mention of spread from root- 
sprouts or other persistence mechanisms. Because this 
plant has an extensive root system it is a likely candidate 
to remain persistent from cultivation and spread locally. 


Hypericaceae 


Hypericum androsaemum L.: DIST: SnFrB: CS: N: 
DOC: Alameda Co.: Strawberry Creek behind UC- 
Berkeley Botanical Garden, June 16, 1993, B. Ertter 
11898 (UC). 

Hypericum calycinum L.: DIST: CCo, SnFrB: CS: 
TEN: DOC: Contra Costa Co.: Berkeley-Oakland Hills, 
Grizzly Peak Boulevard ca. %4 miles south of Lomas Can- 


86 MADRONO 


tadas junction, dense patch on east side of road at edge 
of mixed scrub, elev. ca. 100 ft. June 25, 1993, B. Ertter 
11907 (UC); San Francisco Co.: Howell, J.T. et al. (1958, 
p. 102); Thomas, H. (1961, p. 239): NOTES: Gen. spread- 
ing vegetatively in California where it is widely planted 
and often persistent. Fully naturalized and spreading by 
seed in Oregon and Washington States, but similar behav- 
ior not confirmed in California. 

Hypericum hookerianum Wight & Arn.: DIST: NCo: 
CS: TEN: DOC: Mendocino Co.: forest road near Little 
North Fork Gualala River and Doty Creek, UTM Zone 
10, 4298N, 4550E, elev. 100 ft, one large patch in second 
growth redwood forest, shrub 1—2 m tall. July 8, 2000, G. 
Leppig 1453 (CDA, HSC); Santa Barbara Co.: Monte- 
cito, Montecito School for Girls, escape from cultivation. 
June 9, 1951, H.M. Pollard s.n. (DAV, SBBG), det. by 
J.T. Howell. 


Lamiaceae 


Calamintha sylvatica Bromf. subsp. ascendens (Jor- 
dan) P.W. Ball: DIST: SnFrB: CS: TEN: DOC: Ala- 
meda Co.: Strawberry Canyon northeast of Panoramic 
Place in Oakland-Berkeley Hills, roadcut below oak for- 
est, small persisting patches at two sites along fire road. 
Sept. 2, 2000, B. Ertter 17518 (UC): NOTES: Nomen- 
clature as in Flora Europaea, which uses a narrow circum- 
scription of Satureja. If the same generic circumscriptions 
were applied in North America, no native Satureja would 
occur in California. An alternate name for this taxon is 
Satureja calamintha (L.) Scheele subsp. ascendens (Jor- 
dan) Brig. This is apparently the first record of the species 
occurring spontaneously in North America, although the 
closely related C. nepeta (L.) Savi is widely established 
in eastern North America. 

Cedronella canariensis (L.) Willd. ex Webb & Berth.: 
DIST: CCo: CS: NCI: DOC: San Francisco Co.: Howell, 
J.T. et al. (1958, p. 119); Thomas, H. (1961, p. 299); Jep- 
son, W.L. (1943, pg. 400); Mud Lake, vicinity of San 
Francisco Bay. July 1914, A. King s.n. (JEPS). 

Galeopsis tetrahit L.: DIST: MP: CS: NCI: DOC: Mo- 
doc Co.: Corporation Ranch, Likely. July 8, 1958, 7.C. 
Fuller 1973 (CDA); Pit River Valley south of Alturas, 
irrigation ditch near S end of west side road, roadside. 
July 12, 1947, H.L. Mason and V. Grant 13417 (DAV, 
UC). 

Lamiastrum galeobdolon (L.) Ehrend. & Polatsch.: 
DIST: NCo: CS: TEN: DOC: Humboldt Co.: Arcata, 
Arcata Community Forest, 100 m S of trails 5 and 10 
intersection. Second growth redwood forest. Large stolon- 
iferous mat. April 13, 2000, G. Leppig 1292, K. Neander 
(CDA, HSC): NOTES: Single 1/10 ha. patch. Represented 
here by a variegated cultivar, also sold as Lamium gal- 
eobdolon or Lamium variegatum. 

Lavandula stoechas L.: DIST: CCo, NCo: CS: C: 
DOC: Alameda Co.: sidewalk crack on Shattuck Ave. nr. 
Lincoln St. north Berkeley. June 25, 2001, B. Ertter 17699 
(UC); Sonoma Co.: Fuller Mountain Rd., nr summit of 
Fuller Mtn., occasional on roadside in mixed conifer for- 
est. April 22, 2000, G. Leppig 1311 (CDA, HSC). 

Mentha xX villosa Huds.: DIST: SnFrB: CS: NCI: 
DOC: Napa Co.: Napa River riparian zone on gravel bar 
with willows, ca. % mile north of Trancas Street, city of 
Napa. El. 10 ft, TO6N, RO4W, Sec. 34, MD. Napa 7.5’ 
quad. Sept. 25, 1989, J. Ruygt 2408 (UC). 

Monarda citriodora Cerv.: DIST: ScV: CS: N: DOC: 
Sacramento Co.: roadside, Jackson Rd. (Hwy 16) W of 
Eagles Nest Rd. TO8N, RO6E, Sec. 25, MD. Elev. 125 ft. 


[Vol. 49 


July 17, 1997, F. Carl s.n. (CDA); loc. cit. August 15, 
1997, G.D. Barbe 4478 (CDA). 

Rosmarinus officinalis L.: DIST: CCo, SCo: CS: C: 
DOC: Alameda Co.: Albany waterfront, on former land- 
fill, several local shrubs. Aug. 19, 1994, B. Ertter 13077 
(UC); Orange Co.: Newport Bay, North Star County 
Beach area on the SW end of the bay, ca. 2 km inland of 
Hwy 1, clearly spontaneous, not planted. Oct. 6, 1990, 
A.C. Sanders 10202 (DAV, UCR): NOTES: Single mature 
shrub also persisting near Huckleberry Regional Botanical 
Area, Alameda Co., spontaneous? 

Salvia longistyla Benth.: DIST: CCo: CS: N: DOC: 
Monterey Co.: Locally but abundantly spontaneous on 
bank of Big Sur River at Big Sur. Apr. 13, 1961, J.T. 
Howell 36487 (CAS, CDA); Munz, P.A. (1968, pg. 103); 
Howitt, B.E and J.T. Howell, (1973. pg. 29): NOTES: 
Cited on pg. 1343 of the Jepson Manual as “not natural- 
ized’”’; however field observation reports it extant and vig- 
orous in 2000, acc. to G. Norman via M.A. Matthews 
(personal communication to Hrusa, 4-2000). 

Salvia microphylla Benth.: DIST: CCo, NCoRO, SCo: 
CS: NCI: DOC: Marin Co.: Howell, J.T. (1970, pg. 358); 
Monterey Co.: Matthews, M.A. (1997, pg. 179).; Coal 
Chute Pt., dry sunny loam, originally cult. Aug. 14, 1936, 
L.B. Wheeler 4369. (Point Lobos State Reserve Herbari- 
um); Santa Barbara Co.: Munz, P.A. (1968, pg. 704); 
Sonoma Co.: Best, C., et al. (1996, pg. 164): NOTES: 
Cited on pg. 1343 of the Jepson Manual as “not natural- 
ized”’ under the name S. grahamii Benth. 

Salvia reflexa Hornem.: DIST: CaRE SNE: CS: GH/ 
C: DOC: Inyo Co.: garden in Independence, Rosedale 
Dr., elev. 4000 ft. July 24, 1996, M. DeDecker 6559 
(CDA, RSA); Shasta Co.: garden of residence at 3657 
Encanto Way northeast of Redding, surrounded by blue 
oak woodland. May 16, 1993, B. Ertter 11837 (UC). 

Salvia virgata Jacq.: DIST: MP, SBr., n SN: CS: NCI: 
DOC: Nevada Co.: just east of Grass Valley, on Empire 
Mine property. July, 1972, L. Mott. s.n. (JEPS); Empire 
Mine property, end of Stacy Lane off Highway 49, south 
side of Grass Valley, TI6N, RO8E, Sec. 34, MD. Weedy 
in 2-acre meadow. June 8, 1972, 7.C. Fuller and G.D. 
Barbe 964 (CDA, UC), det. by Ian C. Hedge (RGBE), 
Feb. 1987; San Bernardino Co.: Lake Arrowhead, garden 
escape. Aug., 1931, Braunton 1056 (DS); Siskiyou Co.: 
scattered plants on 600 sq. ft of drainage way in dry 
rangeland, adjacent to wet slough, Greenhorn Valley, ca. 
2 mi W of Yreka. June 24, 1964, 7.C. Fuller 12244 
(CDA), det. by E. McClintock; loc. cit. July 29, 1968, 
C.S. Giebner s.n. (CDA); Ager Beswick Road, very dry 
roadside. Aug., 1998, L. Parsons s.n. (SEPS): NOTES: 
Extirpated in Nevada Co; status of Siskiyou Co. plants 
currently under investigation. San Bernardino Co. speci- 
men originally identified as Salvia pratensis L. (sensu 
stricto). Acc. to R. Breckenridge (CDFA, Integrated Pest 
Control Branch), Salvia virgata is readily distinguished 
from S. pratensis s.s. by the foetid odor of its foliage. 

Scutellaria caerulea M. & S.: DIST: CCo: CS: C: 
DOC: Santa Clara Co: Weed in commercial field herb 
crop in Gilroy. Rocket Farms. Sept. 20, 1999, K. Meyer 
s.n. (CDA), det. by G.E Hrusa (UC). 

Stachys floridana Shuttlew.: DIST: ScV: CS: GH/C: 
DOC: Sacramento Co.: Abundant in garden, 2424 Park 
Estates Dr., Sacramento. May 21, 1963, K.S. Buchanan 
s.n. (CDA); loc. cit. June 7, 2000, L. Manger s.n. (CDA): 
NOTES: Tuberous perennial, spreading by rootsprouts. 
Present at this locality for at least 37 years. 


2002] 


Lauraceae 


Cinnamomum camphora (L.) J. Presl: DIST: deltaic 
GV: CS: C: DOC: Contra Costa Co.: Antioch National 
Wildlife Refuge, Stamm Unit, two juvenile individuals on 
riparian margin, apparently spontaneous but source not 
evident. May 26, 2001, B. Ertter et al. 17563 (UC): 
NOTES: A fairly common landscape volunteer, but seed- 
lings generally restricted to irrigated sites and seldom al- 
lowed to mature. 

Laurus nobilis L.: DIST: CCo, NCo: CS: TEN: DOC: 
Humboldt Co.: Arcata, occasional in disturbed empty lot 
in redwood forest. Trees 2—4 m tall. April 12, 2000, G. 
Leppig 1289 (CDA, HSC); San Francisco Co.: Howell, 
leletedlen( los Saipo 7) hbomas: slies(l96L, spe. 173): 
NOTES: Single plant persisting on Albany Hill, Alameda 
Co., from unknown source and origin. 


Lentibulariaceae 


Utricularia subulata L.: DIST: NCoRO: CS: NCI: 
DOC: Mendocino Co.: Meyers-Rice (Madrono, in press); 
spreading through wet depressions and seeps in a pygmy 
forest, 39°15'’N; 123°45’W. Nov. 2, 1997, B. Meyers-Rice 
#MR971102 (DAV): NOTES: Intentionally introduced. 


Limnanthaceae 


Limnanthes macounii Trel.: DIST: CCo: CS: N: 
DOC: San Mateo Co.: Buxton (1998, pg. 184); Along 
east side of Hwy | south of Moss Beach, directly opposite 
Half Moon Bay airport, in cultivated field. Elev. ca. 10 
m. Feb. 24, 2000, R. Schmid 2000-2 (UC): NOTES: Un- 
clear whether a locally naturalized alien or a previously 
overlooked native. 


Linaceae 


Linum trigynum L.: DIST: NCo: CS: NCI: DOC: Son- 
oma Co.: Best, C., et al. (1996, pg. 168). 


Malvaceae 


Anisodontea capensis (L.) Bates: DIST: NCoRO, ScV: 
CS: C: DOC: Sonoma Co.: Best, C., et al. (1996, pg. 
171); Sacramento Co.: Among landscape shrubs along 
sidewalk in Sacramento. July 10, 2000, R. Gill s.n. 
(CDA): NOTES: In the nursery trade as Malvastrum ca- 
pense (L.) Gray & Harv. and so reported for Sonoma 
County (Best et al., 1996). 

Anoda pentaschista A. Gray: DIST: DSon: CS: NCI: 
DOC: Imperial Co.: Weed in citrus, nr Bard. Sept. 9, 
1983, L. Pineda s.n. (CDA). 

Gossypium hirsutum L.: DIST: DSon, ScV: CS: NCI: 
DOC: Imperial Co.: Two ruderal plants betw. rd and base 
of canal bank, N side Hwy 98 to Mt. Signal 1.7 mi W of 
Calexico. Oct. 18, 1962, 7.C. Fuller 9804 (CDA); Sac- 
ramento Co.: City of Sacramento, Tahoe Park neighbor- 
hood, near the old fairgrounds, volunteer. Sept. 18, 1995, 
D. Goosen s.n. (DAV). 

Hoheria populnea A. Cunn.: DIST: CCo: CS: NCI: 
DOC: San Francisco Co.: Voluntary in dwarf conifer 
area, Strybing Arboretum, Golden Gate Park, San Fran- 
cisco. Sept. 20, 1973, G. Beutler s.n. (CDA). 

Lavatera olbia L.: DIST: CCo: CS: NCI: DOC: San 
Francisco Co.: Shrubs to 8 ft tall, commonly naturalized 
on non-irrigated waste ground of formerly cultivated gar- 
den, Stanyon St., San Francisco. Aug. 4, 1970, 7.C. Fuller 
se (EDA): 

Lavatera trimestris L.: DIST: SCo: CS: NCI: DOC: 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 87 


Santa Barbara Co.: Smith, C.F (1976. pg. 192); Edge of 
water, Lauro Canyon Reservoir nr. San Roque Rd., Santa 
Barbara. June 25, 1975, C. Smith and J.L. Johnson s.n. 
(CDA). 

Malva verticillata L.: DIST: SCo: CS: NCI: DOC: 
Santa Barbara Co., Ventura Co.: Smith, C.F (1976, pg. 
LOS): 

Sida spinosa L.: DIST: SnJV: CS: NCI: DOC: Fresno 
Co.: near Sanger by the Ciba-Geigy Research Station on 
Annadale Ave., ca. 4% mi W of Reed Ave. Aug., 1996, B. 
Fischer s.n. (DAV). 


Moraceae 


Fatoua villosa (Thunb.) Nakai: DIST: SCo, SnJV: CS: 
GH/C: DOC: Kern Co.: Greenhouse weed, Arvin. Oct. 
19, 1998, Lapp et al. s.n. (CDA); Riverside Co.: Sanders, 
A.C. (1996, pg. 527); San Bernardino Co.: Ontario. 
Weedy throughout nursery, under benches, in walkways. 
Oct. 21, 1983, Cohen s.n. (CDA); San Diego Co.: Nurs- 
ery containers, greenhouse, Fallbrook. Feb. 14, 1985, F. 
McCutcheon s.n. (CDA): NOTES: Also confirmed, but 
not vouchered, from Tehama Co., in commercial green- 
houses. Similar vegetatively to species of Laportea (Ur- 
ticaceae) and has been reported as that genus. Seed form 
readily distinguishes Fatoua from Laportea. 

Ficus palmata Forssk.: DIST: SCo: CS: NCI: DOC: 
Santa Barbara Co.: Spontaneous in creekbed, W Fk. 
Cold Spring Cyn., Santa Barbara. Dec. 23, 1958, H.M. 
Pollard s.n. (CAS, CDA): NOTES: Cited in Munz (1974) 
Smith, C.E (1976) and Rejmanek and Randall (1994) as 
F. pseudocarica Miq. 


Nymphaeaceae 


Nymphaea alba L.: DIST: NCo: CS: NCI: DOC: Men- 
docino Co.: Smith, G. and C. Wheeler (1990-1991, pg. 
150). 


Oleaceae 


Fraxinus uhdei (Wenz.) Lingel.: DIST: SCo: CS: NW: 
DOC: Los Angeles Co.: ““Cottonwood Swamp’’, conflu- 
ence of San Francisquito Canyon stream and two tribu- 
taries draining off the southeast slopes of Red Mountain. 
Warm Springs Mtn. 7.5’ quad., TOSN, R16W, Sec. O01, 
elev. 1680-1690 ft [possible hybrid]. 7.S. Ross 7835 
((RSA? UCR, UC); San Bernardino Co.: Colton, S Pel- 
lisier Rd., near the corner of W Center St. and Orange St., 
S40 ily 20's W. LOZSROSW, SB? Bley, 262 m. 
Ephemeral creek with sandy bottom and the shaded grassy 
slopes above it. Nr. historical settlkement. Associated with 
Populus fremontii, Juglans californica and Rubus discol- 
or. Solitary sapling. Apr. 4, 1999, Mitch Provance 1763 
(UCR): NOTES: Widespread and sometimes locally com- 
mon in coastal southern California riparian zones. For ex- 
ample, along the Santa Ana River near Riverside it forms 
a conspicuous element of the tree canopy near the conflu- 
ence of Spring Brook. Where street runoff flows into per- 
manently moist riparian areas, this species usually ap- 
pears. Identification is confused by the fact that it seems 
to hybridize with the F. velutina Torr. (F. pennsylvanica 
Marsh) complex, including both native plants and cv. 
‘Modesto.’ Easiest to identify in winter because it is ev- 
ergreen, unlike the others. Much was cut last year in the 
area around Haskell Creek in the Sepulveda Basin during 
an effort to control exotics. Acc. to John Eckhoff (person- 
al communication to Sanders), they are “‘finding this tree 
in many of the riparian areas we visit or find ourselves 


88 MADRONO 


working in, like San Gabriel River and Big Tujunga Wash 
at the base of the San Gabriel Mts.”’. Voucher requested 
but not received. 

Ligustrum lucidum W.T. Aiton: DIST: NCoRO, ScV, 
SnBr, SCo: CS: NW: DOC: Riverside Co.: Riverside, 
weed tree in landscaping on UCR campus, elev. 1100 ft. 
Dec. 16, 2000, A.C.Sanders 23728 (UCR); Sacramento 
Co.: Quinn, J., et al. (1991); San Bernardino Co.: San 
Bernardino Mtns., Thurman Flats Picnic Area, below 
Mountain Home Village, Yucaipa 7.5’ quad., TO1S, 
ROIW, Sec. 08, SB, 34°06'30"N; 117°00'05’W, elev. 3480 
ft/1061 m, dense alder forest along the stream; solitary 
small tree ca. 3 m high in forest understory. May 27, 2000, 
A.C. Sanders 23432 (UCR): NOTES: This species is scat- 
tered around as an urban weed tree in the Riverside area; 
seeds apparently dispersed by birds. Also observed by 
Hrusa as seedlings and young trees in riparian zone along 
Arcade Cr., N side Interstate 80, Sacramento Co.; also by 
J. Ruygt (pers. comm., 3-2001), as seedlings on Redwood 
Rd., ca. 0.5 mi W of Hwy 29, and as seedlings and adult 
trees along bank of Camille Cr. At Polley Drive, both near 
Napa in Napa Co. Seldom collected, but apparently wide- 
spread in moist habitats. 

Ligustrum ovalifolium Hassk.: DIST: CCo, NCo, ScV: 
CS: NW: DOC: Mendocino Co.: Sinkyone Wilderness 
State Park, old homesite 2.7 miles south of Needle Rock 
Ranch House. Lost Creek trailhead in red alder woodland. 
June 25, 1995, F. Bowcutt 2009 (DAV, HSC); Rt. 1 near 
entrance to McKerricher State Park. Self-sustaining for 
over 50 years. July 30, 1981, G. L. Smith and C. R. Wheel- 
er 7205 (HSC). Monterey Co.: Elkhorn Slough National 
Estuarine Research Reserve, disturbed fields near South 
Marsh. May 22, 2000, G. Leppig 1382 (HSC); Sacra- 
mento Co.: Quinn, J., et al. (1991): NOTES: Known sites 
highly localized. 

Olea africana Mill.: DIST: SCo: CS: TEN: DOC: Riv- 
erside Co.: Riverside, Mt. Rubidoux, NE foot of the 
mountain above the end of 9th St., 33°59’N; 117°23'W, 
TO2S, RO2W, Sec. 22, SB. Elev. 1000 ft/305 m, E-facing 
decomposed granite slopes at edge between landscaped 
(residential) areas and coastal sage scrub. Disturbed and 
weedy with some no longer tended ornamentals. A locally 
common shrub or tree to 10 m tall. Oct. 23, 1996, A.C. 
Sanders 19643 (UCR): NOTES: Some individuals prob- 
ably originally planted (persisting ornamentals), but others 
growing in cracks of boulders, etc. and plainly spontane- 
ous. This is so scarce as a cultivated plant that this might 
easily be the only naturalized locality, where it is doing 
well. Reproduction is apparently by seed. 


Onagraceae 


Fuchsia magellanica Lam.: DIST: CCo, NCo: CS: 
NW: DOC: Contra Costa Co.: Cerrito Creek west of San 
Pablo Avenue, several reproducing shrubs on stream bank. 
Sept. 17, 1999, B. Ertter 16845 (UC); Humboldt Co.: 
Low shrub naturalized at base of Alnus sp., loop trail 
above Fern Canyon, Prairie Ck. Redwood State Park, 7.1 
mi W of Highway 101 on Fern Canyon Rd. June 18, 1974, 
G.D. Barbe 1872 (CDA); Mendocino Co.: Mendocino, 
edge of headland behind Presbyterian church, uncommon 
shrub mixed in willow thicket. Sept. 29, 1992, B. Ertter 
11449 (UC); Monterey Co.: Carmel Highlands, Fern 
Canyon east of Highway 1, shade of pine forest along 
creek. June 13, 1993, B. Ertter, V. Yadon, and M.A. Mat- 
thews 11890 (UC); Gibson Canyon, near Carmel High- 
lands. Growing in moist cyn. bottom, 6—10 pls. over sev- 
eral hundred yds, 600 ft elev. June 13, 1994, D. Kelch s.n. 


[Vol. 49 


(DAV); San Francisco Co.: Lobos Creek between Lin- 
coln Blvd. and Baker Beach, deep shade. Nov. 15, 1992, 
B. Ertter 11459 (UC): NOTES: Some individual sites 
may be considered tenuous (TEN). Label data for second 
Mendocino Co. site in Sinkyone Wilderness State Park, 
[old home sites. T24N, R19W, Sec. 26, MD. May 23, 
1989, F. Bowcutt 1325 (DAYV)], are not clear that location 
iS Spontaneous. 

Fuchsia X hybrida Voss.: DIST: SCo: CS: NCI: DOC: 
Ventura Co.: Persisting or spontaneous (?) in a clump of 
Ricinus shrubs on SPRR right of way, east Ventura. June 
1, 1961, H.M. Pollard s.n. (CDA). 


Orobanchaceae 


Orobanche hederae Duby: DIST: CCo: CS: TEN: 
DOC: Alameda Co.: UC-Berkeley campus, small per- 
sisting colony in Hedera groundcover next to Koshland 
Hall. June 7, 2000, B. Ertter 17310 (UC); between student 
center and Alumni House. May 8, 2001, B. Ertter 17626 
(UC). 


Papaveraceae 


Fumaria capreolata L.: DIST: CCo, SnFrB: CS: N: 
DOC: Alameda Co: Landscape weed in Livermore area. 
March 9, 1994, C. Elmore s.n. (DAV); Contra Costa Co.: 
Miller Knox Regional Park, at edge of excavation on 
west-facing hillside, localized but dense colony at base of 
coast live oak. Mar. 21, 1999, B. Ertter 16486 (UC); Ma- 
rin Co.: Muir Beach, 5.5 miles west of Hwy 1. Open area, 
slightly sandy soil, coastal strand with partial moisture, 
full sun. Not abundant. Elev. 50 ft. May 16, 1992, J.N. 
Le 17 (DAV); San Francisco Co.: Spontaneous along 
path betw. the Conservatory and Fuchsia garden, Golden 
Gate Park, San Francisco. Sept. 25, 1980, J.T. Howell 
53901 (DAV); San Mateo Co: McClintock, E., et al. 
(1990, pg. 135): NOTES: Known sites widely scattered. 
Differing from the other naturalized species of Fumaria 
in California in the larger flowers (ca. 12 mm long) and 
broadly ovate sepals. Apparently becoming more com- 
mon. Similar to F. macrosepala Boiss. which may also be 
represented in California (Ertter 16486). 

Papaver X hybridum L.: DIST: CCo, ScV, SnJV: CS: 
NCI: DOC: Kern Co.: Point of Rocks, western (Kern) 
County. Apr. 27, 1950, E. McMillan and C. Smith 2684 
(DAV); Antelope Valley, 3 mi W of Point of Rocks, the 
probable origin of the plants, not previously observed here 
by local farmers. Apr. 8, 1962, E. Twisselmann 6770 
(CDA, DAV); Madera Co.: E side of Rd 26, 0.1 mi S of 
Ave. 12, 1 mi W of Madera, dominant weed on one acre 
of newly planted vineyard. Apr. 6, 1967, J.S. Davis s.n. 
(CDA); loc. cit. Apr. 19, 1967, T.C. Fuller 15609 (CDA, 
DAV); Sacramento Co.: SW corner of Metropolitan Air- 
port property near Garden Highway. Apr. 4, 1992, K. Mill- 
er s.n. (CDA, DAV); San Luis Obispo Co.: Choice Val- 
ley Hills, SE side of Sinsheimer Flat, dense colony in a 
bare area in a dry-farm wheat field (barley field in 7027). 
Apr. 29, 1962, E. Twisselmann 7025, 7027 (CDA, DAV): 
NOTES: The Twisselmann and J.S. Davis specimens 
were originally determined as Papaver apulum Ten. var. 
micranthum (Bor.) Fedde. Comment by Twisselmann in- 
dicated it is recent in San Luis Obispo Co. 


Passifloraceae 


Passiflora caerulea L.: DIST: NCoR, SnGb, SCo, s 
SN: CS: N: DOC: Los Angeles Co., Riverside Co. and 
San Bernardino Co:. Sanders, A.C. (Madrono, in press); 


2002] 


Fresno Co.: Sequoia Mills, no date, K. Brandegee s.n., 
(UC); Napa Co.: Calistoga, July 11, 1910, K. Brandegee 
s.n. (UC): NOTES: Long persisting and difficult to erad- 
icate in garden situation, indicating strong potential to nat- 
uralize. Serves as host plant for non-native gulf fritillary 
caterpillars. Reported as non-spontaneous in Ventura Co. 
by H.M. Pollard (specimen at DAV, SBBG). 

Passiflora manicata (Juss.) Pers.: DIST: SCo: CS: 
NCI: DOC: Santa Barbara Co.: Smith, C.F (1976, pg. 
197). 

Passiflora mixta L. f.: DIST: CCo: CS: N: DOC: San 
Francisco Co.: Golden Gate Park, local patch at junction 
of Crossover Drive and John EK Kennedy Drive. Sept. 11, 
1993, B. Ertter 12269 (UC): NOTES: Passiflora mollis- 
sima auct. non (Kunth) L.H. Bailey as discussed under P. 
tarminiana. Freely reseeding in garden situation on Cat- 
alina Ave., Berkeley (Alameda Co.). 

Passiflora tarminiana Coppens & Barney: DIST: 
CCo: CS: N: DOC: Contra Costa Co.: edge of vacant 
lot at SW corner of San Pablo Avenue and Carlson Ave., 
climbing on Sambucus at edge of Cerrito Creek. Sept. 17, 
1999, B. Ertter 16846 (UC): NOTES: Observed by Ertter 
to be also established upstream near BART path. Seed- 
lings occasionally encountered (and eliminated) by Ertter 
on Albany Hill, near Cerrito Creek. population; Reported 
also from Riverside Co. All original determinations as 
Passiflora mollissima auct. non (Kunth) L.H. Bailey, cur- 
rent determinations by D. Goldman. See Novon 11(1): pg. 
9, 2001, for more information. Potentially a noxious pest, 
as in Hawaii (as P. mollissima sensu auct.), where capable 
of smothering native forests. 


Pedaliaceae 


Sesamum indicum L.: DIST: ScV, SNF: CS: C: DOC: 
Mariposa Co.: Few plants along roadside, Hwy 140 nr. 
Catheys Valley. Aug. 30, 1978, K.A. Parker s.n. (CDA): 
Sacramento Co: Single plant in asphalt divider, Kiefer 
Blvd, nr. Bradshaw Rd. Aug. 30, 1977, K. Miller s.n. 
(CDA): NOTES: Occurrences probably originating via 
commercial bird seed. 


Plumbaginaceae 


Limonium ramosissimum (Poir.) Maire subsp. prov- 
inciale (Pignatti) Pignatti: DIST: SCo: CS: NW: DOC: 
Santa Barbara Co., Ventura Co.: From Rick Burgess 
garden in Oxnard; originally collected in Carpenteria Salt 
Marsh, where it has naturalized about its mouth and is a 
very serious problem. 1994, Rick Burgess s.n. (SBBG): 
NOTES: Apparently from garden plants in an adjacent 
subdivision. Data provided by Dieter Wilken (SBBG). 


Polygalaceae 


Polygala myrtifolia L.: DIST: SCo: CS: NCI: DOC: 
Santa Barbara Co.: Smith, C.F (1976, pg. 183). 


Polygonaceae 


Polygonum multiflorum Thunb.: DIST: SnFrB: CS: 
GH/C: DOC: Marin Co.: Rampant weed in garden of 
Margadant Hayakawa, Eldridge Ave, Mill Valley. Vigor- 
ously spreading, but does not flower. Jan. 15, 1976, M. 
Hayakawa s.n (CDA), det. by E. McClintock, 9/1977: 
Cultivated in greenhouse [from roots dug at site of pre- 
vious collection], 3294 Meadowview Rd., Sacramento. 
Nov. 21, 1980, G.D. Barbe 3023 (CDA): NOTES: Dried 
tubers of this species (“Fo-ti” or ‘Ho Shou-wu’) are used 
as a folk remedy (Tyler 1982). 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 89 


Polygonum orientale L.: DIST: SCo, ScV: CS: C: 
DOC: Sacramento Co.: spontaneous in residential yard, 
2458 Catalina Dr. Sacramento. Aug. 11, 1975, F. Hine 
s.n. (CDA); Santa Barbara Co.: Smith, C.F (1976 pg. 
122): NOTES: A specimen labeled only as “Mendocino 
Co., July 25, 1958, T. Erickson, Jr. s.n” is at CDA, but 
spontanaiety unknown. Other known occurrences are fu- 
gutives from cultivation. 


Ranunculaceae 


Caltha palustris L.: DIST: CCo, NCoR, NCo: CS: 
NW: DOC: Alameda Co.: Oakland Hills, Canyon east of 
Skyline Blvd, growing in stream that separates Huckle- 
berry Preserve from Sibley Volcano Park, along Skyline 
trail, nr. old homestead site. May 1, 1993, E.A. Dean 359 
(DAV): Mendocino Co.: Smith, G. and C. Wheeler. 
(1990-1991, pg 151-152). Sonoma Co.: Rubtzoff. P. 
(1959, pp. 31-32): NOTES: Rubtzoff record also reported 
in Best, C., et al. (1996, pg. 201). 

Clematis terniflora DC.: DIST: ScV: CS: TEN: DOC: 
Sacramento Co.: Invasive in residental landscape and ad- 
jacent field on Larkspur Lane in Citrus Heights. Both sites 
relatively moist, both with sunny and shady situations. 
Climbing in Populus fremontii to 40+ ft, also in lower 
borders. Apr. 28, 2000, G.F. Hrusa 15389 (CDA): 
NOTES: Origin of infestation uncertain, but may have 
originated as garden ornamental. First noted as a pest in 
this site in 1992. Owners are attempting extirpation. 

Clematis vitalba L.: DIST: CCo: CS: NCI: DOC: San 
Francisco Co.: Howell, J.T. et al. (1958, pg. 72): Thomas, 
H. (1961, pg. 169): NOTES: Volunteer from planted ma- 
terial in Strybing Arboretum, Golden Gate Park, San Fran- 
cisco. 

Nigella damascena L.: DIST: SnFrB,. NCoRO: CS: N: 
DOC: Contra Costa Co.: waste ground, Brentwood Road 
at SPRR tracks, Brentwood. TOIN, RO2E, Sec. 18, MD. 
June 18, 1974, J. deFremery and C. Butler s.n. (CDA): 
Sonoma Co: Best, C., et al. (1996, pg. 203): NOTES: 
Garden escape, naturalized as a weed about habitations. 

Ranunculus cortusifolius L.: DIST: CCo: CS: TEN: 
DOC: Alameda Co.: Berkeley, University of California 
campus, untended garden plots at NW corner of Valley 
Life Sciences Building. May 8, 2001, B. Ertter 17625 
(UC): NOTES: Possibly originating from deliberately 
strewn seeds, but now self-sustraining and spreading to 
adjacent untended plots. Determination by R. Ornduff, 
2000. 


Rhamnaceae 


Ziziphus jujuba L.: DIST: ScV: CS: C: DOC: Sacra- 
mento Co.: Ditch at corner Sheldon Rd and Hwy 99, sap- 
ling. July 7, 1998, N. Wymer s.n. (CDA): Yolo Co.: 15— 
20 plants up to 1 meter tall on W side of Hwy 113 appr. 
1 mi N of Covell exit in Davis. Obviously spontaneous, 
some inside ROW. Aug. 21, 2001, D. Adams s.n. (CDA): 
NOTES: Escape from cultivation, perhaps from garden 
trash. 


Rosaceae 


Cotoneaster lacteus W.W. Smith: DIST: CCo, SnFrB: 
CS: NW: DOC: Alameda Co.: Albany Hill, scattered col- 
ony. Feb. 9, 1997, Ertter 15449 (UC): Contra Costa Co.: 
Ygnacia Valley Road cut through Lime Ridge, single 
shrub. Jan. 5, 1997, Ertter 15432 (JEPS): San Mateo Co.: 
McClintock, E., et al. (1990, pg. 144): NOTES: Noted 
elsewhere in the east San Francisco Bay Area. Flowers 


90 MADRONO 


like C. pannosus but leaves larger, 3.5—7 cm long, obvi- 
ously depressed-veiny, obovate-elliptic with obtuse apex; 
inflorescence often more floriferous as well. 

Crataegus monogyna Jacquin: DIST: NCoRO, SnFrB: 
CS: NW: DOC: Alameda Co.: Dry Creek/Pioneer Re- 
gional Park east of Hayward. Jan. 19, 1992, B. Ertter 
10894 (UC); Anthony Chabot Regional Park, Cascade 
Trail. April 29, 1981, Jack Stratford s.n. (JEPS); also ob- 
served by Ertter as well-established in Sibley Volcanic 
Regional Preserve, Oakland Hills; San Mateo Co.: 
McClintock, E., et al. (1990, pg. 102): NOTES: Also ob- 
served as widely scattered plants on the Palisades, south 
shoulder of Mt. St. Helena in Napa Co. Thorny shrub to 
small tree, leaves 3—5-lobed halfway or more to midvein. 

Cydonia oblonga Mill.: DIST: ScV, SCo, SnJV: CS: 
NCI: DOC: Sacramento Co.: bush 10 ft tall and across, 
one of a number of plants persistant from cult. in a fence- 
row, W side of Elk Grove-Florin Rd., 0.1 mi N of Sheldon 
Rd., Elk Grove. Nov. 19, 1969, 7.C. Fuller 19101 (CDA); 
Santa Barbara Co.: Smith, C.E (1976, p. 160); Stanis- 
laus Co.: % mi W of La Grange, Nof Hwy 132, woodland 
at edge of dredge tailings, occasional escape in this area. 
April 3, 1969, P. Allen 123 (DAV): NOTES: Sacramento 
County site now in developed area, probably extirpated. 

Cydonia sinensis Thouin: DIST: NCo: CS: NCI: 
DOC: Mendocino Co.: Smith, G. and C. Wheeler, (1990— 
1991, pg. 181): NOTES: Reported as Chaenomeles si- 
nensis (Thouin) Koehne. 

Eriobotrya japonica Lindl.: DIST: SnFrB, SCo: CS: 
NCI: DOC: Alameda Co.: Oakland, Joaquin Miller Park, 
Sunset Trail. Occasional in redwood forest. Dec. 4, 2000, 
G. Leppig 1521 (HSC); Santa Barbara Co.: Smith, C.F 
(1976, p. 160): NOTES: Seedling observed on Albany 
Hill (Alameda Co.) by Ertter. Expected elsewhere. 

Filipendula vulgaris Moench: DIST: SnFrB: CS: NCI: 
DOC: Alameda Co.: few scattered plants uphill from 
Australasian beds, Univ. of California Botanic Garden, 
Strawberry Canyon, Berkeley. Oct. 12, 1978, G.D. Barbe 
2459 (CDA). 

Photinia davidsoniae Rehd. & Wilson: DIST: CCo: 
CS: NCI: DOC: San Francisco Co.: Howell, J.T. et al. 
(1958, p. 84): NOTES: Probably no more than a casual 
escape from cultivation. 

Potentilla anglica WLaicharding: DIST: NCo, SCo, 
ScV, SnFrB: CS: NW: DOC: Contra Costa Co.: Ertter, 
B. (1997, pg. 78); Bishop Ranch Regional Open Space 
southwest of San Ramon, shaded streambed. Aug. 8, 
1993, B. Ertter and B. Olson 12198 (UC); Mendocino 
Co.: Smith, G. and C. Wheeler, (1990-1991, pg. 184); 
Sacramento Co.: Wet seep behind house, S end of Blue 
Oak Flat at summit of American River Bluffs, Folsom 
Lake State Park, Natoma Unit. Perhaps not persisting. 
May 21, 1990, G.F. Hrusa 7930 (CDA); San Diego Co.: 
Poway, Blue Sky Ecological Reserve, Oak Grove area of 
park, stream edge. Aug. 22, 1995, M. Hanson s.n. (UC): 
NOTES: Included in Munz and Keck (1959) but left out 
of Jepson Manual due to lack of sufficient evidence avail- 
able at the time that this species was truly naturalized in 
California. 

Potentilla reptans L.: DIST: SnJV: CS: TEN: DOC: 
Fresno Co.: Town of Fresno, a lawn weed. June, 1997, 
B. Fischer s.n. (CDA, DAV). 

Prunus laurocerasus L.: DIST: NCo, SCo: CS: NW: 
DOC: Humboldt Co.; Arcata, occasional, naturalized in 
redwood forest behind Humboldt State University, west of 
Fern Lake, April 12, 2000, G. Leppig 1291 (CDA, HSC); 
Santa Barbara Co.: upper Alturas Rd., Santa Barbara. 


[Vol. 49 


Sept. 30, 1939, M. Van Rensselaer 1343 (UC): NOTES: 
Escaping from Humboldt State University landscaping. 

Prunus persica (L.) Batsch: DIST: CaR, SnFrB: CS: 
C: DOC: Contra Costa Co.: south base of Mount Diablo, 
lone tree in Sycamore Canyon, Apr. 25, 1995, B. Ertter 
and C. Thayer s.n. (JEPS); Los Angeles Co.: Whittier 
Hills, (Puente Hills, pro parte): Turnbull Canyon nr. 
mouth. Three trees, escape or persistent from cult.?, no 
habitations nearby. T2S, R11W Sec. 22. Elev. 505 ft. Apt. 
21, 1991, T. Ross 4499 (CDA, RSA, UC); Siskiyou Co.: 
Mount Shasta, along Everett Memorial Hwy, adventive in 
chaparral. Elev. 4300 ft. July 4, 1968, W.B. and V.G. 
Cooke 39264 (UC). 

Prunus serrulata Lindl.: DIST: SnFrB: CS: C: DOC: 
Alameda Co.: Strawberry Canyon firetrail behind UC 
Berkeley Botanical Garden, several small trees in shady 
wooded area, Apr. 15, 1999, B. Ertter 16502 (UC). 

Pyracantha coccinea M. Roem.: DIST: NCoRO, 
SnFrB: CS: TEN: DOC: Alameda Co.: Strawberry Can- 
yon, at Poultry Farm Station, 500 ft, Oct. 6, 1934, C.M. 
Belshaw 218 (UC). Sonoma Co.: Best, C., et al. (1996, 
jog, ZS). 

Pyracantha crenatoserrata (Hance) Rehder: DIST: 
CCo, GV, SCo, SnFrB: CS: N: DOC: Contra Costa Co.: 
Lime Ridge open space, occasional shrub in grassland of 
quarry area, May 10, 1998, B. Ertter 16115 (JEPS); Sac- 
ramento Co.: Moist disturbed areas in vicinity of Willow 
Creek parking area, E side Lake Natoma. Common. 
38°37'N; 121°02'W. Elev. 75 m. May 2, 1990, G.F. Hrusa 
7802 (CDA); loc. cit. July 26, 1990, G.F. Hrusa 8155 
(CDA); San Mateo Co.: McClintock, E., et al. (1990, pg. 
147); Ventura Co.: Seedlings growing in railroad gravel 
ballast at Foster Park, Ventura. Aug. 14, 1964, H.M. Pol- 
lard s.n. (CAS, CDA); Escape from cultivation in brush 
strip under Ventura River bluff N of and nr Santa Ana 
Blvd. crossing, Oak View. Oct. 23, 1963, H.M. Pollard 
s.n. (CAS, CDA): NOTES: Distinguished by the narrowly 
obovate leaves to 2 cm wide, often marginally toothed and 
apically notched. Also observed by Ertter on Albany Hill 
in Alameda Co. Common as seedlings in urban gardens, 
lawns. Expected elsewhere. 

Pyracantha crenulata (D. Don) M. Roem.: DIST: 
NCoRO, SnFrB: CS: TEN: DOC: Sonoma Co.: Best, C., 
et al. (1996, pg. 215): NOTES: Also observed by Ertter 
on Albany Hill, Alameda Co. 

Pyrus communis L.: DIST: SCo, ScV, SNE, SnFrB: 
CS: N: DOC: Contra Costa Co.: EBMUD land east of 
Tilden Park, ca. % mile southeast of Inspiration Point, 
grazed grassland near streamlet, single well browsed large 
shrub. Possibly only persisting from cultivation. May 7, 
1993, B. Ertter and C. Thayer 11700 (UC); Mono Co.: 
Spring 0.7 mile S of the mouth of Milner Creek, 0.6 mile 
N 08° W of Copper Queen Mine, TO4S, R33E, Sec. 21, 
MD. Elev. 5210 ft. Persisting apparently several decades 
after introduction, compact trees ca. 4 m high in silty 
moist alluvial soil with Chrysothamnus, Stanleya, Robi- 
nia. May 29, 1986, James D. Morefield and Douglas H. 
McCarty 3718 (RSA, UC); Orange Co.: Santa Ana 
Mountains, 4 mi NE Lake Irvine, Black Star Canyon, 
Cleveland National Forest, Trabuco District. Oak wood- 
land canyon with riparian woods along the stream. Grassy 
clearings and chaparral on the canyon slopes. April 18, 
1966, E.W. Lathrop 6176 (RSA); Southern Santa Ana 
Mountains, San Mateo Canyon Wilderness Area. Western 
edge of the wilderness in Lucas Canyon along canyon 
floor from crossing of Lucas Canyon Trail, upstream % 
mile to spur trail leading to old mining habitations in side 
canyon, T07S, RO6W, Sec. 10, SB. Elev. 1000—1350 ft. 


2002] 


Low shrub, single individual, apparently adventive about 
old homestead. May 20, 1992, S. Boyd and T. Ross 7468 
(RSA); Santa Barbara Co.: Smith, C.F (1976, pg. 164); 
Santa Clara Co.: Coyote Creek. Mar. 30, 1901, A.P. 
Chandler 911 (UC); Sonoma Co.: Pitkin Marsh, near Fo- 
restville, eastern branch of the Upper Marsh. At the edge 
of a thicket, on damp ground. Oct. 26, 1952, P. Rubtzoff 
1332 (CAS, RSA); W branch upper marsh. Oct. 26, 1952, 
Rubtzoff 1329 (UC); Ventura Co.: Matilija Canyon, off- 
spring of old ranch planting in creek bottom. Oct. 6, 1946, 
H.M. Pollard s.n. (RSA, SBBG); Munz, P.A. (1974, pg. 
758): NOTES: Sacramento Co.: observed as a possible 
escapes adjacent to orchards approx. one mile S of Free- 
port. 

Rosa multiflora Thunb. ex Murray: DIST: CaR, CCo, 
ScV: CS: NCI: DOC: Butte Co.: ca. 5 airmiles southwest 
of Gridley, south side of Evans Reimer Rd. ca. % mile 
east of Pennington Road, T17N, RO2E, Sec. 07, MD. 70 
ft, May 2, 1998, L. Ahart 7958 (JEPS): San Mateo Co.: 
McClintock, E., et al. (1990, pg. 148): Siskiyou Co.: Mt. 
Shasta City, sedge meadow at corner of Lake St. and 
Commercial St. June 9, 2001, B. Ertter 17662 (UC): Shas- 
ta Co.: Squaw Creek Canyon, April 19, 1992, D.W. Tay- 
lor 12602 (UC). Det. by B. Ertter, 11-2001. 

Rubus ulmifolius Schott var. ulmifolius: DIST: CCo, 
NCoR: CS: N: DOC: Napa/Lake Co.: Highway 29 on 
north side of Mount St. Helena, roadside at edge of ser- 
pentine area, locally common, forming large tangles. Nov. 
27, 1997, B. Ertter and L. Constance 15885 (UC): 
NOTES: Also observed in the Presidio of San Francisco 
(San Francisco Co.) by Ertter. 


Rubiaceae 


Coprosma repens A. Rich.: DIST: CCo: CS: N: DOC: 
San Francisco Co.: Howell, J.T. et al. (1958, p. 130); 
Thomas, H. (1961, p. 323); San Mateo Co.: McClintock, 
E., et al. (1990, pg. 149): NOTES: Observed in Monterey 
Co. by Sanders, (10-2000) on seacliffs in Pacific Grove. 
An escape from cultivation. 


Salicaceae 


Populus nigra L. cv. ‘Italica’: DIST: SCo, SNE, 
NCoRO: CS: TEN: DOC: Alameda Co.: Tree NW of 
Botanical Garden, Strawberry Canyon, Berkeley. July 10, 
1943, B. Rodin 245 (DAV): Sonoma Co.: Best, C., et al. 
(1996, pg. 221); Mono Co.: mouth of Milner Creek, 
spring 0.7 mi S; 0.6 mi n 83 degrees w of Copper Queen 
Mine, Owens Valley drainage, White Mountains. Silty 
moist alluvial soil sloping 3% SW _ w/Chrysothamnus, 
Stanleya, Salix, Rosa, Artemisia. Elev. 5210 ft. T04S, 
R33E, Sec. 21, MD. May 29, 1986, James D. Morefield 
and Douglas H. McCarty 37l6e (UC): Ventura Co.: 
Spreading by root-sprouts from parent tree along ditch on 
Loma Dr. S of Ventura Ave, Ojai Valley, Ventura. Nov. 
8, 1967, H.M. Pollard s.n. (CAS, CDA): NOTES: Per- 
haps original plants persistent from cultivation, but often 
spreading aggressively by root sprouts. Best et al. (1996) 
report that only staminate plants are in cultivation in Cal- 
ifornia. 


Sarraceniaceae 


Sarracenia aff. rubra Walter: DIST: NCo: CS: N: 
DOC: Mendocino Co.: Smith, G. and C. Wheeler (1990— 
1991, pg. 171): NOTES: Intentionally introduced. Ac- 
cording to B. Meyers-Rice (DAV) the pitcher plants intro- 
duced in this site are represented by hybrid forms. No 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 91 


clear S. rubra has been as yet found, although some in- 
dividuals may outwardly approach it in appearance. 


Scrophulariaceae 


Anarrhinum bellidifolium (L.) Willd.: DIST: SnFrB: 
CS: C: DOC: Alameda Co.: Strawberry Canyon firetrail 
behind UC Berkeley, 500 m elev., July 18, 1999, N. Hill- 
yard s.n. (JEPS): NOTES: Determination verified by B. 
Ertter and M. Wetherwax. 

Limnophila X ludoviciana Thieret: DIST: ScV: CS: 
TEN: DOC: Butte Co.: Rice field at NE corner of Ham- 
ilton and Riceton Rds, Rice Experiment Station NW of 
Biggs. Scattered plants on moist mud betw. experimental 
rice plots. TION, RO2E, Sec. 35, MD. Elev. ca. 100 ft. 
Oct. 22, 1998, Oswald, Ahart, Ertter 9674 (CDA, CHSC): 
Rice Experiment Station near Biggs, mud of drained ex- 
perimental plots, rare. Oct. 22, 1998, B. Ertter 16446 with 
L. Ahart and V. Oswald (UC), det. by D. Philcox (K), 
1998; Yuba Co.: Ahart, L. (1981, pgs. 7, 8): In shallow 
water in open area in a rice field, 2 miles NE of Marys- 
ville. Sept. 25, 1977, L. Ahart s.n. (CAS, CDA, CHSC) 
det. by D. Philcox (K), Dec., 1977: NOTES: =Limno- 
phila indica (L.) Druce X L. sessiliflora Blume. Reported 
by Ahart (1981) as L. indica. Examination of both Yuba 
and Butte Co. material by Philcox (K) indicates our plants 
are best referred to the hybrid form. Weed of agriculture, 
subject to elimination via drainage of cultivated rice fields. 

Mazus japonicus Kuntze: DIST: SCo: CS: GH/C: 
DOC: Orange Co.: Bordier’s Nursery, Irvine Boulevard, 
Santa Ana, spontaneous in locally prepared soil mix. Aug. 
9, 1973, G.D. Barbe 1694 (CDA, UC). 

Penstemon strictus Benth.: DIST: SNE: CS: TEN: 
DOC: Mono Co.: Witcher Creek, at crossing of jeep road 
(Inyo National Forest road 4854) from Swall Meadows. 
June 29, 1988, D.W. Taylor 9917b (JEPS): NOTES: Ap- 
parently escaping from nearby rural residential areas of 
Swall Meadows, either from cultivation or as seeded for 
‘wildflowers’; common in unmanaged ruderal vegetation 
of vacant lots in the developed portion of the subdivision. 
The Witcher Creek location is in an area of natural veg- 
etation removed some 500 m and over a small hill from 
the most proximal habitations. The occurrence was still 
present in 1998. Report and data provided by D.W. Taylor. 

Penstemon subglaber Rydb.: DIST: SNE: CS: TEN: 
DOC: Mono Co.: Mammoth, vacant lot near the Post Of- 
fice. Aug. 2, 1998, D.W. Taylor 16939 (JEPS): NOTES: 
Apparently escaping from nearby areas seeded for ‘wild- 
flowers.’ The plants reseed in unmanaged, ruderal vege- 
tation in the developed portion of town, but have not yet 
been seen afar. Report and data provided by D.W. Taylor. 

Scrophularia peregrina L.: DIST: SCo: CS: N: DOC: 
Los Angeles Co.: Ross, T. and S. Boyd (1996, pg. 436): 
Claremont, RSABG; SE edge of Indian Hill Mesa, ver- 
nally moist clay embankment. Common winter and spring 
weed in the area. March 24, 1993, T. Ross 6819 (CDA, 
RSA, UC): NOTES: Introduced in 1950s during biosys- 
tematic study of Scrophularia. Robust specimens may 
mimic Scrophularia californica. Native to Mediterranean 
region. 

Verbascum olympicum Boiss. non Bunyard: DIST: 
NCoRI: CS: NW: DOC: Sonoma Co.: Common on 
benches above rocky, dry bed of Leslie Creek from 
approx. Mark West Ck. confluence to first main tributary 
from NE. Mixed with Verbascum thapsus. July 25, 2000, 
G.F. Hrusa 15690, 15691, 15692 (CDA, and to be dis- 
tributed): NOTES: Readily distinguished from other nat- 
uralized Verbascum by its dense white-arachnoid pubes- 


92 MADRONO 


cence, especially in the inflorescence. Flowers are brighter 
yellow and larger than in V. thapsus. First escaping from 
ornamental plantings at residence on CAS Pepperwood 
Preserve in 1976. Population, mixed with Verbascum 
thapsus, is large and apparently increasing. Possible hy- 
brid forms are also scattered at this locality (July 25, 2000, 
Hrusa 15693, CDA). 


Solanaceae 


Atropa belladonna L.: DIST: CCo, ScV: CS: NCI: 
DOC: Sacramento Co.: Hampton Rd., Sacramento. Gar- 
den weed. Aug. 24, 1999, Ken and Mary Brown s.n. 
(CDA); San Francisco Co.: Howell, J.T. et al. (1958, p. 
121); Thomas, H. (1961, p. 304). 

Capsicum annuum L.: DIST: SCo: CS: NCI: DOC: 
Ventura Co.: Spontaneous in fallow field on lower Pier- 
pont Bay, Ventura. Sept. 15, 1959, H.M. Pollard s.n. 
(CDA). 

Cestrum parqui L’Her.: DIST: c SNE SCo, SnFrB: 
CS: NCI: DOC: Amador Co.: S side of Jackson on rd to 
Mokelumne Hill. Aug. 9, 1933, C.B. Wolf 5206 (UC); 
Napa Co.: Napa City. 1893, no collector (UC); Santa 
Barbara Co.: Toro Canyon Creek at Southern Pacific RR 
crossing, Summerland. Aug. 8, 1961, H.M. Pollard s.n. 
(CDA, SBBG). 

Lycium ferocissimum Meirs: DIST: SCo: CS: NCI: 
DOC: Los Angeles Co.: shrub 3 m tall, on canal bank in 
salt marsh, 400 m N of 431 East Culver Blvd, Playa del 
Rey. TO2S, R1I5W, Sec. 27, SB. Jan. 16, 1979, T.C. Fuller 
20255 (CDA): NOTES: Persistent from cultivation? 

Nicotiana X sanderae Hort. ex Wats.: DIST: CCo: 
CS: C: DOC: San Francisco Co.: Howell, J.T. et al. 
(1958, p. 121); Thomas, H. (1961, p. 305). 

Nicotiana tabacum L.: DIST: SnFrB, SCo: CS: C: 
DOC: Contra Costa Co.: | volunteer plant at waterfront 
at E end of building, Golden Gate Fish Co., Point San 
Pablo, Richmond. Dec. 20, 1966, R. DeBoer s.n. (CDA), 
det. by T.-C. Fuller; San Diego Co.: Cholla nr. San Diego. 
Apr. 7, 1885, Cleveland and Greene s.n. (UC); Santa 
Barbara Co.: Smith, C.F (1976, pg. 251). 

Petunia violacea Lindl.: DIST: CCo, SCo: CS: NCI: 
DOC: San Bernardino Co.: Alta Loma, Hellman and 
19th St., roadside. April 2, 1961, G. Pilone 140 (DAV); 
San Luis Obispo Co.: roadside, no dwelling in vicinity, 
Perfumo Canyon Rd, 0.9 mi W of junct. with Los Osos 
Valley Rd., ca. 5 mi SW of San Luis Obispo. July 19, 
1962, 7.C. Fuller 9237 (CDA); Ventura Co.: Naturalized 
locally along San Antonio Creek at Royal Oaks Dairy and 
for some distance downstream, Ojai. Aug. 13, 1966, H.M. 
Pollard s.n. (CDA, SBBG): NOTES: An escape from cul- 
tivation but extent of true naturalization not known. 

Solanum gayanum (Remy) Phil. f.: DIST: CCo: CS: 
NCI: DOC: San Francisco Co.: Howell, J.T. et al. (1958, 
p. 122); Thomas, H. (1961, p. 303): NOTES: Escape from 
cultivation. 

Solanum scabrum Mill.: DIST: ScV, SnJV: CS: C: 
DOC: Butte Co.: Six volunteer plants, city dump, Sterling 
City. T24N, RO4E, Sec. 28, MD. Sept. 11, 1974, Sauer 
and Heinricks s.n. (CDA), det. by T.C. Fuller; Fresno Co.: 
Town of Five Points, growing at the Westside Research 
and Extention Center in mix of seeds of Solanum nigrum 
complex planted in tomato field for herbicide trial. 1999, 
Bill Fisher s.n. (DAV); Sacramento Co.: Vigorous 
growth to 8 dm, vacant lot W side of 1226 D St. Sacra- 
mento. Aug. 26, 1988, G.D. Barbe 4310 (CDA). 

Solanum villosum Mill.: DIST: ScV: CS: C: DOC: 


[Vol. 49 


Yolo Co.: Knights Landing, tomato field near wet area. 
May, 1996, G. Miyou s.n. (DAV). 


Urticaceae 


Boehmeria cylindrica (L.) Sw.: DIST: GV: CS: NW: 
DOC: Sacramento Co.: Hrusa, G.E (2000, pgs. 138-— 
139); NOTES: Observed by Hrusa as becoming increas- 
ingly common about Snodgrass Slough, Sept. 2001; also 
observed by Hrusa as common in Sutter Slough and Elk 
Slough (Yolo Co.) immed. W of Courtland, Aug. 2002. 
Urtica-like foliage but without stinging hairs, the opposite 
leaves and inflorescence of sessile, aggregate clusters su- 
perficially resemble Lamiaceae. Readily distinguished by 
the mostly ebracteate inflorescence (although leafy at the 
apex), scabrous leaves, round stems, absence of a corolla, 
and unisexual flowers. Boehmeria cylindrica may be mon- 
oecious or dioecious, but all California material examined 
to this point is monoecious, with staminate flowers fewer 
and confined to the uppermost parts of the spike. 

Laportea aestuans (L.) Chew: DIST: SCo: CS: GH/C: 
DOC: San Diego Co.: Weed in greenhouses. Commercial 
nursery, Fallbrook. April, 1999, A. Amador s.n. (CDA): 
NOTES: Vegetatively similar to Fatoua villosa (TYhunb.) 
Nakai, (Moraceae). Seed form unequivocally distinguishes 
F. villosa from the genus Laportea. 


Verbenaceae 


Verbena rigida Spreng.: DIST: CCo, NCoR, ScV, 
SnFrB: CS: NCI: DOC: Contra Costa Co: few scattered 
plants in lawn, Elmwood Rd, El Sobrante. Sept. 3, 1965, 
R. DeBoer s.n. (CDA); Marin Co.: Spontaneous in filled 
ground, San Pedro Rd. at Marin Yacht Club, San Rafael. 
Filled ground, edge of Yacht Harbor. Sept. 23, 1965, Gor- 
don True 2652 (CDA); Sacramento Co.: spontaneous in 
sidewalk, 15th and P Sts., Sacramento. Aug. 15, 1961, 
R.M. Hawthorne s.n. (CDA); Sonoma Co.: Howell J.T. 
(1972, pg. 102): NOTES: Sometimes treated as V. venosa 
Gillies and Hooker. 

Vitex agnus-castus L.: DIST: SCo, ScV: CS: NCI: 
DOC: Santa Barbara Co.: naturalized on SPRR tracks 
at Ortega St., Santa Barbara. June 11, 1968, P. Okuye s.n. 
(CDA); Yolo Co.: Creek by apiary, University Farm Cam- 
pus. Oct. 14, 1932. H.A. Barthwick s.n. (DAV). 


Vitaceae 


Cissus antarctica Venten.: DIST: SCo, SnFrB: CS: 
NCI: DOC: Alameda Co.: Escaped ornamental, flood 
channel, Union City. Feb. 2, 1981, Sweigert s.n. (CDA); 
Los Angeles Co.: Weed in CalTrans yard betw 710 and 
Ist. St., East Los Angeles. Oct. 9, 2001, M. Adams s.n. 
(CDA, UCR). 

Vitis aestivalis Michx.: DIST: SCo: CS: NCI: DOC: 
Ventura Co.: Apparently spontaneous, Valley Rd nr. San- 
ta Ana Blvd., Oak View. June 2, 1972, H.M. Pollard s.n. 
(CAS, CDA). 

Vitis rupestris Scheele: DIST: NCoRO: CS: TEN: 
DOC: Sonoma Co.: Best, C., et al. (1996, pg. 247): 
NOTES: Winegrape rootstock. 


Zy gophyllaceae 


Peganum harmala L.: DIST: DMoj: CS: EXT?: DOC: 
Kern Co.: Edwards AFB, roadside % mi E of Lancaster 
Blvd, on Old Hospital Rd. Across from “‘P”’ housing sec- 
tion. Single plant. TOON, R1OW, Sec. 14, SB. June 29, 
1990, D. Charlton 4453 (CDA); San Bernardino Co.: 
Abandoned 15 acre pasture, Minneola and Elkhorn Rds, 


2002] 


Newberry Springs. TOON, RO2E, Sec. 27, SB. July 8, 
1988, J. Hitchcock s.n. (CDA); 50-60 clumps, 3-5 ft 
diam. in abandoned pasture, Minneola and Elkhorn Rds, 
Daggett. T11N, ROLE, Sec. 27, SB. July 1, 1988, J. Hitch- 
cock s.n. (CDA); Newberry Springs, vicinity of Silver 
Valley Rd. N of RR tracks nr the airport, in yards of res- 
idents and vacant lots. Aug. 23, 1988, D. Pendleton s.n. 
(DAV): NOTES: Noxious Weed under eradication by Cal- 
ifornia Dept. of Food and Agriculture. 


ANGIOSPERMS—-MONOCOTS 
Alismataceae 


Sagittaria brevirostra Mackenzie & Bush: DIST: 
CCo: CS: NCI: DOC: Marin Co: Chileno Valley, the 
laguna. June 18, 1947, J.T. Howell 23261 (UC): NOTES: 
Specimen originally determined as S. latifolia; current de- 
termination by K. Rataj, 1968, with confirmation by C.B. 
Hellquist, 1994. Cited in Haynes and Hellquist (2000). 

Sagittaria rigida Pursh: DIST: CaRH, CCo, NCoRI: 
CS: NW: DOC: Oswald, V.A. et al. (1998, p. 185): 
NOTES: Introduced to ‘improve/enhance’ waterfowl hab- 
itat? 


Araceae 


Arum palestinum Boiss.: DIST: NCo: CS: TEN: DOC: 
Humboldt Co.: Arcata Marsh and Wildlife Sanctuary, 
Butchers Slough upper salt marsh. May 8, 1998, G. Lep- 
pig 755 (CDA, HSC); Humboldt Bay, Woodley Island 
near hunting cabin, moist Juncus meadow. UTM Zone 10 
4518N 4020E. July 14, 1976, Peter Sorenson s.n. (HSC): 
NOTES: This species, although locally present in small 
numbers is long-lived and thus highly persistent. Arcata 
Marsh population under eradication by City of Arcata. 

Dracunculus vulgaris Schott: DIST: SCo, SnFrB: CS: 
N: DOC: Alameda Co.: ca. halfway up Claremont Can- 
yon in Oakland Hills, locally established colony on steep 
roadfill. June 16, 1995, B. Ertter and L. Constance 14258 
(UC); loc. cit. June 7, 1992, B. Ertter 11082 (UC): Santa 
Barbara Co.: Santa Barbara, growing in lot probably 
once under cultivation. May 17, 1948, R.S. Beal, Jr. s.n. 
(UC): NOTES: Claremont Canyon colony still present as 
of April 1990, possibly expanding. 

Pinellia ternata (Thunberg) Makino: DIST: SnFrB: 
CS: NCI: DOC: San Francisco Co.: Golden Gate Park, 
“accidentally introduced”’. May, 1935, E. Walther s.n. 
(UC): NOTES: Determined by S.A. Thompson, 1994; 
original determination as P. tripartita (Blume) Schott. 


Cyperaceae 


Bulbostylis barbata Kunth: DIST: CCo: CS: GH/C: 
DOC: Santa Cruz Co: Weed growing in commercial 
greenhouses, Watsonville. Probably originating from Sri 
Lanka. May 14, 1998, T. Watson s.n. (CDA). 

Cyperus flavescens L.: DIST: ScV: CS: NW: DOC: 
Butte Co.: damp sand, E side Sac. R., NW Parrott Land- 
ing, 1 mi SE Ord Ferry, 12 mi SE Chico. Aug. 13, 1999, 
Lowell Ahart and V. Oswald 8143 (CDA, CHSC), det. by 
V. Oswald, 10-99; loc. cit. Oct. 28, 1999, L. Ahart 8303 
(CHSC): Arch Rock tunnel, Feather River Hwy, Elev. 500 
m. Sept. 6, 1981, L. Ahart 3123 (CHSC, DAV): Peter 
Ahart Ranch, 1 mi N and 2% mi E of Honcut. July 19, 
1975, L. Ahart 901 (CHSC); loc. cit. Aug. 21, 1975, L. 
Ahart 957 (CHSC); loc. cit. Sept. 4, 1987, L. Ahart 5877 
(CHSC); loc. cit. Aug. 19, 1995, L. Ahart 7618 (CHSC, 
JEPS); wet sand on N. Fk. Feather River nr Poe Power- 
house, riparian woodland, elev. 890 ft. Aug. 30, 1987, L. 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 93 


Ahart 5865 (CHSC); wet sand on margins of small pond 
in Oroville Wildlife Area. Common. Elev. 140 ft. Aug. 
23, 2000, L. Ahart 8664 (CHSC): moist gravel bar along 
Sacramento R. 0.25 mi downstream from Murphy’s 
Slough. Riparian woodland. July 31, 1983, V. Oswald 94] 
(CHSC); Feenstra’s Riverview Orchard W end of Cana 
Hwy between S half Cana Lake (on Dicus Slough) and 
Sacramento River. Uncommon. Wet gravel near shallow 
pools on large gravel bar along river. Portion of Rancho 
Bosquejo, elev. 160 ft. T23N, RO2W, MD. Aug. 13, 1987, 
V. Oswald 3215 (CHSC): Tehama Co.: Hog Lake Plateau 
along Hwy. 36 NE of Red Bluff, T28N, RO3W, Sec. 14, 
MD, elev. 430 ft, moist soil along Paynes Creek ca. 1 mi 
upstream from the old bridge site. Sept. 4, 1996, V. Os- 
wald and L. Ahart 8270 (CHSC), det. L. Janeway; Yuba 
Co.: Wet soil in irrigated pasture on Eugene Ahart Ranch 
in Marysville, elev. 100 ft. Sept. 11, 1975, L. Ahart s.n. 
(CHSC), det. V. Oswald; shallow water on disturbed rocky 
soil, S side Scott Forbes Rd, 0.25 mi E of intersection 
Scott Forbes Rd and Peoria Rd. E of Gray Dr. E of 
Browns Valley. Uncommon, el. 300 ft. Aug. 13, 1997, L. 
Ahart 7865 (CHSC), det. V. Oswald: NOTES: Additional 
collections from some of these same localities are at 
CHSC, the earliest from 1961. 

Cyperus flavicomus Michx.: DIST: ScV, SNF: CS: N: 
DOC: Butte Co.: University of California Biggs Rice Re- 
search Station, near the corner of Riceton and Hamilton 
Roads, in rice field, Aug. 30, 2000, DiTomaso s.n. (DAV): 
Edge of Middle Honcut Road, about *% mile E of Hwy 70, 
ca. 12 mi S of Oroville, common, valley grassland, L. 
Ahart 4810 (CHSC); 8 mi NE of Butte City, ditch bank. 
Oct. 15, 1946, J.E. Chattin and C. Ferrell s.n. (UC); Tu- 
lare Co.: 3 mi below Three Rivers. Sept. 25, 1920, L. 
Abrams 7715 (CAS): NOTES: Determination of Di- 
Tomaso s.n. by G.C. Tucker. The Ahart and Abrams spec- 
imens were originally determined as Cyperus albomargin- 
atus Mart. & Schrad., a synonym of C. flavicomus; how- 
ever, the original determinations as C. albomarginatus 
have not yet been confirmed. 

Cyperus gracilis R. Br.: DIST: SCo: CS: NCI: DOC: 
Los Angeles Co: volunteer in residential garden, Santa 
Monica. Aug. 20, 1983, Tom Yutani s.n. (CDA). 

Cyperus iria L.: DIST: SCo, ScV: CS: TEN: DOC: 
Santa Barbara Co.: Germinating in coco fiber from Sri 
Lanka. Commercial greenhouse in Carpenteria. Grown to 
maturity in CDFA greenhouse, Sacramento. Oct. 14, 1998, 
T. Watson s.n. (CDA); Yuba Co.: Dry to damp soil, dis- 
turbed area S side Woodruff Lane, just W Kimball Lane, 
5 mi NE Marysville. Elev. 80 ft. Sept. 5, 1999, L. Ahart 
8197 (CDA, CHSC). 

Cyperus owanii Boeck: DIST: CCo, SCo: CS: NCI: 
DOC: Los Angeles Co.: Spontaneous, UCLA Bot. Gar- 
den, Westwood. April 19, 1972, T.C. Fuller 1994] (CDA): 
San Diego Co.: Barranca, Balboa Park, San Diego. April 
22, 1969, T.C. Fuller 18244 (CDA); loc. cit. Oct. 21, 
1969, T.C. Fuller 19012 (CDA, DAV): San Francisco 
Co.: Golden Gate Park, California Academy of Sciences, 
San Francisco. Oct. 18, 1967, T.C. Fuller 16748; loc. cit. 
Nov. 21, 1968, 7.C. Fuller 18013 (CDA); Santa Barbara 
Co.: Smith, C.F (1976, pg. 94): NOTES: Only known 
occurrences in North America N of Mexico. 

Cyperus papyrus L.: DIST: SCo: CS: NCI: DOC: San 
Diego Co.: Beauchamp, R.M., (1986, pg. 48). 

Fimbristylis autumnalis (L.) Roem. & Schult.: DIST: 
ScV: CS: NW: DOC: Butte Co.: Disturbed area E side 
Sacramento River NW Parrott Landing, 1 mi SE Ord Fer- 
ry, 12 mi SW Chico. Riparian woodland. Aug. 13, 1999, 
L. Ahart 8145 and V. Oswald (CDA, CHSC). 


94 MADRONO 


Scirpus cyperinus (L.) Kunth: DIST: c SNF: CS: NW: 
DOC: Mariposa Co.: Moist banks and alluvial flats, Mer- 
ced River near mouth Yosemite Creek, Yosemite Valley. 
Aug. 13, 1976, J.T. Howell 52233 (CAS, CDA); loc. cit. 
Sept. 28, 1978, G.D. Barbe 2447, 2444 (CDA, CHSC): 
NOTES: Determination confirmed by L. Janeway 
(CHSC) 2-2000. 

Scirpus prolifer Rottb.: DIST: SCo: CS: NCI: DOC: 
San Diego Co.: Persisting in sandy soil of lathhouse, 19 
km NW of Fallbrook. Nov. 14, 1978, G.D. Barbe 2472 
(CDA, DAV). 


Eriocaulaceae 


Eriocaulon cinereum R. Br.: DIST: SnJV: CS: EXT: 
DOC: Stanislaus Co.: Krause rice fields, Modesto. Sept. 
18, 1947, B.G. Markos s.n. (CDA): NOTES: Extirpation 
likely as rice fields are no longer maintained in the Mo- 
desto region. Cited from California in Flora North Amer- 
ica (Kral 2000). 


Hydrocharitaceae 


Limnobium laevigatum (Humb. & Bonpl. ex Willd.) 
Heine: DIST: SCo, SnFrB: CS: NW: DOC: Alameda 
Co.: Jordan Pond in Garin Park; East Bay Regional Park 
District, Hayward. Floating and rooted in bottom mud. 
Osis Sig IDLO, Seo Zl, IMs, OCs ZZ, OMS Ik. Jaa 
s.n. (CDA); Garin Regional Park east of Hayward, silted- 
in pond below Jordan Pond, well-established colony both 
floating and stranded. Sept. 4, 1998, B. Ertter 16408 (UC); 
rimming Jordan pond, some in bloom, Oct. 24, 1998, B. 
Ertter 16458 (UC); Riverside Co.: Covering % acre of 
outdoor pond at Desert Lawn Cemetery, Desert Lawn Dr., 
Calimesa. TO2S, ROIW, Sec. 31. SB. Aug. 23, 1999, J. 
Chandler s.n. (CDA): NOTES: Santa Cruz Co.: in a 
backwater of the San Lorenzo River. Voucher specimen 
not yet received, but scrap material confirmed by B. Ertter 
in fall 2000. Not relocated in spring 2001. Floating or 
stranded rosettes spreading by stolons; leaves + round, 1— 
3.5 cm across, sessile to long-pediceled, convexly spongy 
on underside. Sometimes treated as L. spongia subsp. lae- 
vigatum (Humb. & Bonpl. ex Willd.) Lowden. As ob- 
served by Ertter, eradication effort in Jordan Pond by East 
Bay Regional Parks District (EBRPD) has not yet suc- 
ceeded in eliminating the Limnobium, but has concurrent- 
ly severely impacted the originally diverse aquatic flora, 
underscoring just one of the major problems inherent in 
the control of aquatic pests. This plant is currently pro- 
hibited from being sold commercially in California; how- 
ever, as in the case of Salvinia molesta, it remains for sale 
by aquatic plant nurseries and dealers. The closely related 
Limnobium spongia (Bosc) Steud. (sensu stricto) is cur- 
rently offered for mail-order sale by midwest nurseries 
and is thus eventually expected to escape in California. 
Both taxa are a threat to California wild wetlands, irri- 
gation ditches, canals, sloughs, farm ponds or private 
lakes, and if either should become established in navigable 
waterways they are especially likely to spread rapidly and 
widely. The two established and reproducing colonies de- 
scribed above are in non-navigable waters. It should also 
be noted that any number of aquatic taxa sold unlabeled 
in California nurseries could be Limnobium or other un- 
identified taxa which could display the same degree of 
aggressiveness as does Limnobium should they be intro- 
duced into aquatic habitats where there are no natural con- 
trols. 


[Vol. 49 


Iridaceae 


Tris foetidissima L.: DIST: NCoRO, SnFrB: CS: N: 
DOC: Alameda Co.: well established along a stretch of 
the Golden Spike Trail in Redwood Regional Park, Oak- 
land Hills. Oct. 15, 2000, Ertter 17542 (UC); Contra 
Costa Co.: Sibley Volcanic Preserve in Oakland Hills, 
single clump in understory of oak-pine forest behind in- 
terpretive center. May 20, 2000, B. Ertter 17025 (UC); 
Sonoma Co.: Best, C., et al. (1996, pg. 261): NOTES: 
Spreads readily in garden situations. Seeds with red arils 
indicate a high potential for bird dispersal into wildlands. 

Tris germanica L.: DIST: NCoRO, SnFrB: CS: TEN: 
DOC: Santa Barbara Co.: Munz, P.A. (1974, pg. 907); 
Sonoma Co.: Best, C., et al. (1996, pg. 261): NOTES: 
Seen often about old habitations and sometimes persistent 
long after signs of human infestation have disappeared. 
Citations here both imply local naturalization. Also ob- 
served as a casual in Alameda and Contra Costa Cos. 

Tris orientalis L.: DIST: SCo: CS: NCI: DOC: Santa 
Barbara Co.: Smith, C.F (1976, pg. 106). 

Ixia polystachya L.: DIST: NCo: CS: NCI: DOC: 
Mendocino Co.: persisting in scattered locations among 
the headstones of Evergreen Cemetery, Hwy | at Moun- 
tain View Rd, | mi S of Manchester. T13N, R17W, Sec. 
06, MD. May 27, 1982, G.D. Barbe 3628 (CAS, CDA): 
NOTHES: dupl. det. by J.-M. Mullin, 1983 (BM). 

Ixia speciosa Andrews: DIST: NCo: CS: NCI: DOC: 
Mendocino Co.: Smith, G. and C. Wheeler (1990-1991, 
Pearls): 

Moraea collina Thunb.: DIST: CCo: CS: N: DOC: 
Santa Cruz Co.: remnant native coastal prairie site on 
marine terrace lying between Rodeo Gulch and Hidden 
Valley Road, ca. 75 meters SSE of pole 14/117 of the PG 
and E Paul Sweet-Green Valley 115 kv double-wood pole 
transmission line (site is 1.5 air mi NNE of Dominican 
Hospital); elev. 370 ft, grassland on edge of thickets of 
Quercus agrifolia. May 2, 2000, D.W. Taylor 17446 
(JEPS): NOTES: Naturalization localized at present. 
Goldblatt (1998) treats this as Moraea collina Thunb., not 
as Homeria (Thunb.) Salisb. which is the name in use by 
the horticultural trade and by Australian authors. The en- 
tire genus Homeria is on the Federal Noxious Weed Act 
quarantine list (see Federal Register, May 25, 2000, p. 
33741-33743), and is thus illegally sold, although com- 
monly available in California nurseries. Native to South 
Africa. Related species are aggressive invaders of pastures 
in New Zealand, Australia. All Homeria spp. are toxic to 
livestock. 

Moraea polystachya Ker Gawl.: DIST: SCo: CS: NCI: 
DOC: Santa Barbara Co.: Plants abundantly naturalized 
on roadside, N side of Cabrillo Blvd., SW corner of Bird 
Lagoon, Montecito. TO4N, R27W, Sec. 24, SB. Dec. 11, 
1968, 7.C. Fuller 18041; loc. cit. Mar. 23, 1971, T.C. 
Fuller 19820 (CDA). 


Juncaceae 


Juncus nodatus Cov.: DIST: SnFrB: CS: N: DOC: 
Alameda Co.: UC-Berkeley Botanical Garden, edge of 
path near vernal pool section. July 16, 1998, H. Forbes 
s.n. (UC): NOTES: Well-established local colony, not de- 
rived from cultivated material. Determination by B. Ertter. 


Liliaceae (sensu lato) 


Agapanthus praecox Willd.: DIST: CCo: CS: NCI: 
DOC: San Mateo Co.: McClintock, E., et al. (1990, pg. 
167): NOTES: Clearly originating as a garden escape. 


2002] 


Allium cepa L.: DIST: NCoRO: CS: NCI: DOC: Son- 
oma Co.: Best, C., et al. (1996, pg. 269): NOTES: A local 
garden escape in 1963, probably not persistent. 

Allium sativum L.: DIST: NCoR: CS: C: DOC: Son- 
oma Co.: Best, C., et al. (1996, pg. 269): NOTES: A local 
garden escape. 

Amaryllis belladonna L.: DIST: CCo, SCo, SnFrB, 
SnBr: CS: TEN: DOC: Contra Costa Co.: observed by 
Ertter near Donner Cabin, Mount Diablo State Park 
[voucher prepared but lost]; observed elsewhere in San 
Francisco Bay Area.; Los Angeles Co.: Malibu, upslope 
from Winter Canyon, near Pepperdine Univ., N of Pacific 
Coast Hwy, E of Malibu Cyn Rd., SW of Malibu Civic 
Center Way, elev. ca. 300 ft, 34°02’N; 118°42’W, TOIS, 
R17W, Sec. 31, SB. Mar. 15, 1995, S. White 2585 (UCR); 
Monterey Co.: Yadon, V. (1995); San Mateo Co.: 
McClintock, E., et al. (1990, pg. 167); Sonoma Co.: Sea 
Ranch, just ESE of Leeward Rd. entrance along State 
Hwy. 1. Aug. 13, 1975, M. Hektner 220 (DAV), voucher 
for Wasmann J. Biol. 35(1):26—-53: NOTES: Also ob- 
served by Sanders apparently naturalized near Yucaipa, 
San Bernardino Co. Cultivated widely. Often long-per- 
sistent from cultivation and known from numerous home- 
stead sites in northern and southern California. 

Chlorophytum capense (L.) Druce: DIST: CCo: CS: 
C: DOC: San Francisco Co.: Howell, J.T. et al. (1958, 
p-53); Thomas, H. (1961, p. 117): NOTES: A garden es- 
cape. 

Hyacinthus orientalis L.: DIST: CCo: CS: C: DOC: 
San Francisco Co.: Howell, J.T. et al. (1958, p. 53); 
Thomas, H. (1961, p. 117): NOTES: A garden escape. 

Kniphofia uvaria (L.) Hooker [or hybrid derivitive]: 
DIST: CCo, NCo, SCo, SnFrB: CS: NW: DOC: Alameda 
Co.: North foot of Albany Hill (eliminated by subsequent 
roadwork). May 25, 1992, B. Ertter 11070 (UC); Contra 
Costa Co.: Miller Knox Regional Park, open hillside 
south of watertank, single large patch, expanding. Mar. 
21, 1999, B. Ertter 16487 (UC); Humboldt Co: Arcata 
Marsh and Wildlife Sanctuary, dike at Klopp Lake, on 
disturbed fill on Humboldt Bay. April 9, 2000, G. Leppig 
1287 (CDA, HSC); Mendocino Co.: Smith, G. and C. 
Wheeler (1990-1991, pg. 107); Hwy. 1 at Navarro Rd, 
7.5 miles S of Elk. Roadside pastures. Common. July 9, 
2000, G. Leppig 1459 (CDA, HSC); Sinkyone Wilderness 
State Park, mouth of Jackass Creek at the end of Wheeler 
Rd. Old home site on the S side of Cyn. T23N, R19W, 
Sec. 01, MD. May 28, 1987, F. Bowcutt 1011 (DAV); 
San Francisco Co.: Howell, J.T. et al. (1958, pg. 53); San 
Mateo Co.: McClintock, E., et al. (1990, pg. 168);. Santa 
Barbara Co.: upper east fork of Cold Spring Canyon 
above Montecito. Jan. 21, 1967, H.M. Pollard s.n. (SBBG, 
UCR); Sonoma Co.: Best, C., et al. (1996, pg. 275): 
NOTES: Expected elsewhere. Can be long persistent from 
cultivation as at Fort Hunter Liggett (Monterey Co.) 
where it is found about old homesteads without any in- 
dication of spontanaety. Other ‘populations’ may originate 
via garden trash, but many locations are obviously spon- 
taneous. Observed in coastal meadows and on roadsides, 
where occasional to common in Monterey Co. but not 
yet vouchered. Also observed in locations other than the 
records above in Sonoma, Mendocino and Humboldt Cos. 

Leucojum aestivum L.: DIST: CCo: CS: NCI: DOC: 
San Luis Obispo Co.: Keil, D.J., et al. (1985, pg. 222). 

Narcissus pseudonarcissus L.: DIST: NCo, SnFrB: 
CS: NW: DOC: Contra Costa Co.: Mount Diablo State 
Park, Donner Cabin site, persisting colony. Feb. 16, 1997, 
B. Ertter 15451] (JEPS); northwest end of Wildcat Canyon 
Park, occasional clumps among brush-grassland mosaic. 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 95 


Mar. 7, 1999, B. Ertter and L. Fujii 16485 (UC); Molate 
Beach Park, coastal prairie. April 7, 1996, B. Ertter 1458] 
(UC); Humboldt Co.: Fickle Hill Road, abundant in open 
areas. TOON, ROIE, Sec. 27, H. Mar. 9, 1963, J.C. Reppas 
18 (HSC); Old Arcata Road at Jacoby Creek Rd., common 
along roadsides, TOSN, ROIE, Sec. 16, H. Apr. 8, 1965, 
N.D. Dennis 15 (HSC); North Bank Rd., 4 mi N of Arcata, 
escaped from cultivation, ROIE, TOON, Sec. 09, H. Apr. 
14, 1965, J. L. Baker 1965 (HSC): NOTES: A diversity 
of hybrid cultivars are also sporadically encountered (e.g., 
Molate Beach Park, Contra Costa Co.) These may be 
persistent from cultivation, the remnants of garden trash, 
or spontaneous. 

Narcissus tazetta L.: DIST: NCo, SnFrB: CS: NW: 
DOC: Contra Costa Co.: Mount Diablo State Park, Back 
Canyon Road at cross road above Donner Canyon trail- 
head, seepage area in grassland, well established spread- 
ing colony. Jan. 25, 1998, B. Ertter and A. Dennis 15910 
(JEPS); Humboldt Co.: Patrick’s Point State Park, near 
Ceremonial Rock. 21 July 1986. E. Mackey s.n. (HSC). 

Ornithogalum umbellatum L.: DIST: CCo, NCoRI: 
CS: GH/C: DOC: Lake Co.: Upper Lake, along Menden- 
hall Rd, off Hwy 20, in walnut orchard. Apr. 4, 2000, R. 
Elkins s.n. (DAV); Santa Cruz Co.: Garden weed, Riv- 
erside Dr., Watsonville. Apr. 12, 1971, D.H. Shaw s.n. 
(CDA). 

Pancratium maritimum L.: DIST: SCo: CS: N: DOC: 
Los Angeles Co.: El] Segundo Dunes, immediately west 
of L.A. International Airport and Pershing Dr., E of Vista 
Del Mar Blvd., nr. 33°56'N; 118°26'W, coastal dunes for- 
merly largely occupied by residential neighborhoods, the 
houses removed ca. 15 years ago. Assoc. with Croton cal- 
ifornicus, Abronia umbellata, Camissonia cheiranthifolia 
and various persisting ornamentals. Forming vegetative 
clumps in sand; also some reproduction by seed. July 22, 
1987, A.C. Sanders 7156 (RSA, SBBG, UCR): NOTES: 
An escaping ornamental; native to beaches in southern 
Europe. Reported also from Ventura Co. at Mugu Lagoon 
and San Buenaventura State Beach, but no specimens yet 
available for confirmation. 

Tulipa clusiana DC. in Redoute: DIST: SCo: CS: 
TEN: DOC: Riverside Co.: City of Riverside, east edge 
near Moreno Valley, spontaneous at the site of the former 
Desert Nursery, N of Hwy 60, E of Day Street, unknown 
origin, possibly accidentally introduced with other plant 
material, now reproducing and increasing in one area with 
no care or watering. Mar. 23, 1994, A. Miller 94-1 (UCR): 
NOTES: At almost the same time as this collection, this 
species was reported to Sanders as a lawn weed in Riv- 
erside, but no voucher materialized. This plant is appar- 
ently a common weed in the Mediterranean basin. It is 
quite showy and might get moved around intentionally as 
an ornamental. 


Poaceae 


Acrachne racemosa (Roem. & Schult.) Ohwi: DIST: 
SCo: CS: TEN: DOC: Riverside Co.: Sanders, A.C., 
(1996, pg. 524). 

Aira caryophyllea L. var. cupaniana (Guss.) Fiori: 
DIST: SnFrB: CS: NCI: DOC: Contra Costa Co.: Mount 
Diablo State Park, Wall Point Road, prescribed burn. May 
28, 1995, M.L. Bowerman s.n. (JEPS), verified by J. 
Wipff, 1998: NOTES: =Aira cupaniana Guss. Not relo- 
cated in 1998—99. 

Amphibromus neesii Steud.: DIST: ScV: CS: NW: 
DOC: Sacramento Co.: grown to maturity in greenhouse 
from plant dug out of large colony in vernal pool at corner 


96 MADRONO 


of Sunrise and Keifer Aves., SE of Sacramento. July 28, 
2000. G.F. Hrusa s.n. (CDA): NOTES: Although known 
from only a single site, this species is of particular interest 
because of its ability to invade and survive in vernal 
pools. Native to and sometimes listed in the Australian 
literature under the synonym Amphibromus nervosus (R. 
Br.) Druce. First seen in this site in 1990 by J. Glazner of 
North Fork Associates, Auburn, CA (personal communi- 
cation to Hrusa). Amphibromus neesii forms cleistoga- 
mous seeds; these first noted and identified at the Cali- 
fornia Dept of Food and Agriculture Analysis and Iden- 
tification (now Plant Pest Diagnostics) Branch Seed Lab- 
oratory in 1967 as a contaminant in Trifolium 
subterraneum seed imported from Australia. 

Aristida dichotoma Michx.: DIST: n SNF: CS: NW: 
DOC: Butte Co.: Dry bare granite soil, Big Bald Rock. 
Uncommon, inconspicuous. Yellow pine forest, elev. 3260 
ft. Aug. 5, 2000, L. Ahart 8623 and V. Oswald (CDA, 
CHSC); Dry bare granite soil above seeps on bare granite. 
N side Bean Ck. Rd., ~ %4 mi. SE Little Bald Rock and 
1% mile NE Big Bald Rock, 4 airmiles NE Madrone Lake 
and 13 mi NE of Oroville. Uncommon. Yellow pine for- 
est, elev. 3500 ft. T21N RO6E, Sec. 29, MD. Aug. 5, 2000, 
L. Ahart 8624 and V. Oswald (CDA, CHSC): NOTES: 
Determinations by K. Allred (NMCR). 

Chloris truncata R. Br.: DIST: DSon, SCo, SnJV: CS: 
N: DOC: Imperial Co.: Weed in alfalfa, 5 mi S of El 
Centro and 3.5 mi. SW of Heber. T16S R13E, Secs. 35, 
36. Aug. 8, 2000, J. Johnson s.n. (CDA, DAV); Merced 
Co.: Almond orchard at corner of Vista Ave and Hwy. 99 
S, N of Chowchilla. July 19, 1999, J. DiTomaso s.n. 
(CDA, DAV); Riverside Co.: Sanders, A.C., (1996, pg. 
526; 1999, pg. 113); Weedy in hybrid bermudagrass turf, 
turfgrass nursery, Leon Rd. Winchester. Nov. 9, 1978, E. 
Storm s.n. (CDA), det. by G.E Hrusa, Oct., 1999. 

Echinochloa crusgalli subsp. spiralis (Vasing.) Tzvel- 
ev: DIST: ScV, SnGB: CS: N: DOC: Butte Co.: Afton 
Rd. nr. Biggs. Aug. 23, 1947, M.K. Bellue s.n. (CDA); In 
milo field. Sept. 16, 1958, J. Harroun s.n. (CDA); Los 
Angeles Co.: San Gabriel Mtns., Angeles National Forest: 
Little Rock Reservoir, dry sunny sandy flat toward upper 
end. Sept. 6, 1966, L.C. Wheeler 8873 (CDA, RSA); Sac- 
ramento Co. sine loc. July, 1943, Bellue s.n. (CDA): 
NOTES: First report of this variety for North America. 
Expected widely. Determinations by S.L. Mosyakin 
(KW), March 26, 2001. 

Echinochloa esculenta (A. Br.) H. Scholz: DIST: 
NCo, SCoRI, SnJV: CS: NCI: DOC: Del Norte Co.: 
Abundant along a slough, Reservation Ranch, Smith Riv- 
er. Oct. 2, 1968, E.J. Garrett s.n. (CDA); Monterey Co.: 
near Chualar. Sept. 1938, McElrath s.n. (CDA); Fresno 
Co.: near Fresno. Sept., 1938, McElrath s.n. (CDA): 
NOTES: Determinations by S.L. Mosyakin (KW), March 
26, 2001. Echinochloa frumentacea Link (E. crusgalli var. 
frumentaceum (Link) Trimen) has been consistently mis- 
applied to this plant in California. It is probably most 
often an escape from cultivation, although populations 
may be locally persistent. 

Ehrharta longiflora Sm.: DIST: SCo: CS: NW: DOC: 
San Diego Co.: Simpson, M.G., et al. (1996, pg. 79): 
NOTES: Known only from a single location. 

Eragrostis curvula (Schrad.) Nees var. conferta Nees: 
DIST: SCo: CS: N: DOC: Riverside Co.: Sanders, A.C. 
CIGRNO. (2s DZ). 

Eremochloa ciliaris (L.) Merr.: DIST: CCo: CS: EXT: 
DOC: San Francisco Co.: Thurber, G. in S. Watson, 
(1880, pg. 261—2): NOTES: Original report as Jschaemum 
leersioides Munro (specimen at CAS). Citation in Howell, 


[Vol. 49 


J.T. et al. (1958), is based on the above report. Extirpation 
is assumed although there has been no thorough investi- 
gation of its presence or absence. 

Gaudinia fragilis (L.) P. Beauv.: DIST: NCoRO: CS: 
NW: DOC: Sonoma Co.: Daniel, T. and C. Best (1992, 
pp. 309-310); Best, C., et al. (1996, pg. 300): NOTES: 
Reported location highly localized, should be sought else- 
where in the region. 

Glyceria fluitans (L.) R. Br.: DIST: CaRH, NCo, 
NCoRI: CS: NW: DOC: Del Norte Co.: Veneer Mill 
pond, Redwood NP. June 5, 1995, G. Leppig 242 (CDA, 
HSC); Humboldt Co.: Arcata Bottom, wet slough. July 
20, 1933, J. T. Tracy 12801 (HSC, RSA); Stone Lagoon 
moist field. Aug. 3, 1924, J. T. Tracy 6749 (RSA); Arcata, 
Alder Grove Pond, on edge of pond. June 4, 1995, G. 
Leppig 242 (CDA, HSC); loc. cit. May 25, 1998, G. and 
S. Leppig and K. Neander 791 (CDA, HSC); loc. cit. April 
24, 1999, G. and S. Leppig 1125 (CDA, HSC); Lake Co.: 
Kelseyville, swampy places. June 8, 1924, J.W. Blankin- 
ship s.n. (RSA); Shasta Co.: Cassel. June 24, 1930, M. 
Kjilsberg s.n. (DAV): NOTES: According to G. Leppig 
(HSC), this may be an overlooked native. 

Hordeum vulgare L. [sensu lato. Including material re- 
ferable to var. trifurcatum (Schlecht.) Alef.]: DIST: CA- 
FP: CS: C: DOC: Alpine Co.: W Fk. Carson River, ca. 1 
mi W from Woodfords along Crystal Springs Rd. June 22, 
1974, D.W. Taylor 3911 (UC); Los Angeles Co.: San Cle- 
mente Island, widely scattered over mesa summit and S 
slopes. May, 1936, N. Murbarger 174 (UC); Marin Co.: 
nr. Inverness. June 3, 1945, J.T. Howell 20918 (UC); 
Mariposa Co.: valley floor, Yosemite Valley, Yosemite 
Natl. Park. Apr. 1, 1934, P.S. Bartholomew s.n. (UC); 
Mendocino Co.: nr. Ukiah, volunteer in oat-field. May 25, 
1899, J. Burtt Davy and W.C. Blasdale 5059 [awned] 
(UC); May 25, 1899, J. Burtt Davy 5058 [awned] (UC); 
Sacramento Co.: Volunteering in residental garden. Mar. 
29, 1979, K. Miller s.n. (CDA); San Diego Co.: Simpson, 
M.G. et al. (1996, pg. 80); San Luis Obispo Co.: ROW 
along Hwy 46 approx. 4 mi E Paso Robles. Weed on 
roadside, solitary. Elev. 200 m. 35°39’N; 120°26’W. Apr. 
4, 1992, G.F. Hrusa 10345 (CDA); Cuyama Valley, weed 
at edge of barley field, immed. W of int. Hwy 33 and 
Hwy 166. Penetrating into planted field along edges. Elev. 
650 m. 34°55'N; 119°33’W. Apr. 26, 1998, G.F. Hrusa 
14276 (CDA); Siskiyou Co.: Volunteer; subsaline flat and 
roadside, betw. Old Highway 99 and RR at Truttman 
Lane, 3.2 km S of Grenada. Elev. 800m. T44N, RO6W, 
Sec. 34, MD. Apr. 28, 1980, G.D. Barbe 2589 (CDA); 
Ventura Co.: San Nicolas Island, N edge of mesa, E end 
of old landfull, E of Living Compound. Apparently used 
for erosion control on inactive landfill site. Apr. 1, 1992, 
S. Junak SN-789 (JEPS, SBBG): NOTES: Although spo- 
radic occurrences are well-documented, these were all de- 
termined to be casuals and accordingly the species was 
not included in The Jepson Manual. This species is also 
commonly used for roadcut erosion control and it may be 
difficult to determine if a local site or population is estab- 
lished or recently planted. Records listed above are rep- 
resentative; the species can be expected in every Califor- 
nia county. Numerous unvouchered records are on file at 
CDA (based on specimens submitted but not retained); 
observed as common on road cutbanks throughout north- 
ern California, especially so along that part of the Inter- 
state 5 corridor. Hordeum vulgare var. trifurcatum is a 
hooded form, reported as a roadside weed almost as often 
as the typical variety, although probably due to its visible 
distinctiveness and not frequency. Annotations at UC by 
N. Jacobsen (1980) do not recognize var. trifurcatum. 


2002] 


Leptochloa dubia (Kunth) Nees: DIST: CaRF/n SNF: 
CS: N: DOC: Butte Co.: west side of Hwy 70 ca. 1% 
miles north of bridge across the West Branch of Lake 
Oroville, just north Rich Gulch Road, T22N, RO4E, Sec 
09, elev. 1300 ft. Common on dry rocky disturbed soil. 
Sept. 6, 1997, L. Ahart 7894 (CHSC, JEPS). 

Nassella tenuissima (Trin.) Barkworth: DIST: CCo: 
CS: C: DOC: Contra Costa Co.: Creekside Park, El Cer- 
rito, overflow basin for Cerrito Creek, single vigorous 
plant. July, 2000, B. Ertter s.n. (UC): NOTES: Also read- 
ily establishing in sidewalk cracks adjacent to cultivated 
plants in gardens throughout Berkeley, this species has a 
high potential for spread into natural areas. A popular hor- 
ticultural species whose weedy behavior should be care- 
fully monitored. 

Panicum maximum Jacq.: DIST: SCo: CS: GH/C: 
DOC: Riverside Co.: Sanders, A.C. (1996, pg. 529). 

Panicum repens L.: DIST: SnFrB: CS: NCI: DOC: 
Alameda Co.: abundant weed of ornamental shrubs, 
climbing to 3 ft high in some shrubs, Oxford Tract, U.C. 
campus, NE corner of Walnut St. and Hears Ave., Berke- 
ley. TOIS, RO4W, Sec. 02, MD. Nov. 21, 1968, T.C. Ful- 
ler 18011 (CDA): NOTES: Rhizomatous perennial intro- 
duced into California from Florida in Melaleuca stock. 

Panicum rigidulum Bosc ex Nees var. rigidulum: 
DIST: ScV, n SnJV: CS: NCI: DOC: Butte Co.: Bank of 
Feather River, Oroville; elev. 175 it. Sept. 22, 1961, J.T. 
Howell 36714 (CAS, CDA), original det. as P. agrostoides 
Spreng.; Sacramento Co.: 1 mi S of Fair Oaks along 
American River. Growing in sandy gravel of dry flood 
banks along river. Oct. 31, 1961, B. Crampton 6072 
GabiW@): Moc: cit: Oct, 116; 1953.4 B Crampton 1706 
(AHUC); American River near Natoma. Moist sandbar. 
Sept. 21, 1950, R. Tofsrud s.n. (AHUC); Stanislaus Co.: 
edge of Tuolumne River 3 miles SW of La Grange, wet 
soil. Aug. 23, 1961, P. Allen s.n. (JEPS); Tuolumne Co.: 
W side of Hetch Hetchy Dam, Swamp Lake Area, Yosem- 
ite NP. Aug. 9, 1958, H.L. Mason 14855 (UC); Yolo Co.: 
Common among rocks, east levee of Merritt Island, at Rd 
142, SE corner of County. July 22, 1972, C. Quick 72-12 
(AHUC): NOTES: Current determination of P. Allen s.n. 
by M. G. LeLong, 1995; previous determinations include 
P. lindheimeri Nash. and P. agrostoides Spreng. Included 
in Oswald and Ahart (1995, pg. 290) as a synonym of P. 
acuminatum var. lindheimeri (Nash) Fern. 

Panicum texanum Buckl.: DIST: SnJV: CS: TEN: 
DOC: Fresno Co.: In a vineyard along Academy Ave. 
near Sanger. August, 1983, B. Fischer s.n. (AHUC), det. 
by B. Crampton, 1983; Kern Co.: Weed in vineyard, De- 
lano. Sept., 1992, H. Kempen s.n. (DAV), det. by G.FE 
Hrusa, 1992: NOTES: An uncommon agricultural weed, 
sometimes treated in Urochloa, and so filed at DAV. 

Pennisetum glaucum (L.) R. Br.: DIST: SnJV: CS: C: 
DOC: Merced County: Highway 33 about 3 miles S of 
southern outskirts of Gustine; moist roadside ditch near 
Main Canal at Pfitzer Road; TO8S, RO9E, Sec 29, MD. 
37°12'28"N; 121°00'37"W; Howard Ranch quad., elev. 
110 ft. With Paspalum dilatatum, Sorghum halepense 
dominant. July 12, 2000, D.W. Taylor 17480 (JEPS): 
NOTES: Escape from cultivation? Large areas were being 
grown (presumably for dairy silage) within about a half 
mile upstream of the site above. Although there was a 
sizable population, the occurrence could be attributed to 
seed washing downslope along the ditch alignment. In the 
Jepson Manual (pg. 1296), this name was misapplied to 
Setaria pumila (Poir.) R. & S. as the synonymized com- 
bination Setaria glauca (L.) P. Beauv. 

Pennisetum latifolium Spreng.: DIST: CCo: CS: NCI: 


HRUSA ET AL.: NON-NATIVE PLANTS IN CALIFORNIA 97 


DOC: Santa Cruz Co.: three clumps of plants to 12 ft, 
naturalized on shoulder of Southern Pacific RR just N of 
30th Ave. crossing, Santa Cruz. Sept. 29, 1969, J. Bauer 
si =(CDA); oc: cit. ‘Oct. 19, 1969, T.C. £uller’ 18980 
(CDA, DAV, UC); Yolo Co.: Davis, UC farm, volunteer 
in row A12 of grass garden. Oct. 25, 1923, P.B. Kennedy 
s.n. (AHUC). 

Phalaris coerulescens Desf.: DIST: SnFrB: CS: C: 
DOC: Contra Costa Co.: Mount Diablo, Black Hawk 
Ridge Road south of Sycamore Creek, single plant on 
roadside. June 12, 1999, B. Ertter 16713 (JEPS): NOTES: 
Like P. paradoxa, but perennial with bulbous based 
culms, sterile spikelets not clublike. Not relocated in 2000, 
possibly due to road grading at the site. 

Phyllostachys aurea A. & C. Riviere: DIST: SCo: CS: 
NCI: DOC: Placer Co.: Open dry slope above wash, 2 
mi W of Auburn. Nov. 13, 1969, B. Crampton 8531 
(AHUC); Stanislaus Co.: Colony established along per- 
manent stream, La Grange Dam Rd | mi E of La Grange. 
June 20, 1969, P.S. Allen 429 (DAV); Ventura Co.: Pop- 
lar grove and thicket at juncture of San Antonio Creek 
and Ventura River near Arnaz Apple Orchards, Oak View. 
Re-collected after 15 yr in same spot. Colony enlarged. 
Dec. 21, 1962, H.M. Pollard s.n. (AHUC, CAS, CDA, 
SBBG). 

Phyllostachys bambusoides Siebold & Zuccarini: 
DIST: CCo: CS: NCI: DOC: San Mateo Co.: Mc- 
Clintock, E., et al. (1990, pp. 181-182). 

Piptochaetium stipoides Hackel ex Arech. sensu lato: 
DIST: NCo: CS: NW: DOC: Marin Co.: About 4 mile 
E of Hwy 1 on trail to Bolinas Ridge, just north of road 
to Alpine Dam. May 9, 1978, C. Best s.n. (CAS, CDA); 
single plants or patch to 3 meters across, S-facing slope 
of canyon on W slope of Bolinas Ridge, 0.5 km E of Hwy 
1 at southern limits of Dogtown. TOIN, RO8W, Sec. 11, 
MD. June 2, 1978, G.D. Barbe 2392 (CDA); loc. cit. G.D. 
Barbe 2391 (cited in correspondence, specimen location 
unknown), determinations by Gladys Perez-Camargo 
(BAA), B. Rosengurtt (MVFA), and L.T. Ellis (K); infre- 
quent patches to 10 meters across, S facing slope W of 
Bolinas Ridge, betw. Coppermine and Wilkins gulches at 
Dogtown (Woodville), 4 km N of Bolinas. Elev. 121 m. 
TOIN, RO8W, Sec. 11, MD. May 16, 1979, G.D. Barbe 
2502 (CDA, DAV, UC); Ridge between Garden Canyon 
and Pike County Gulch on western slope of Bolinas Ridge 
ca. 3.25 miles north of Stinson Beach. Occasional in hard 
packed soil of old road. May 17, 1978, 7.H. Harris s.n. 
(DAV): NOTES: Numerous more recent collections from 
the same localities above are at UTC. It is interesting that 
although the original California specimens of the P. sti- 
poides alliance were determined as var. purpurascens 
(Hackel) Parodi, study of the associated correspondence 
indicated considerable variation among these specimens. 
Intraspecific application within the P. stipoides group is 
currently in dispute; and in addition, Barbe 2392 was orig- 
inally det. at K as P. grisebachii (Speg.) Herter. Subse- 
quent annotations at SI by E.G. Nicora indicate all were 
part of P. stipoides var. purpurascens. More recent col- 
lections determined by E. Sanchez (BA) again indicate 
two taxa are present at this site: P. stipoides var. purpur- 
ascens and var. stipoides. Apparently further field study 
on Bolinas Ridge would be useful. Well-established there 
where it is mixed with P. setosum. The land was private 
with restricted access until two years before the first col- 
lections in 1978, thus the species has possibly been pre- 
sent on the site for a considerable time. Speculated to have 
been introduced with cattle from S. America. 

Polypogon imberbis (Phil.) Bjorkm.: DIST: CCo: CS: 


98 MADRONO 


NCI: DOC: Contra Costa Co.: near Martines. June 7, 
1900, Burtt Davy 6662 (US); San Luis Obispo Co.: 
Oceano Beach in moist spots among sand dunes. July 18, 
1947, R.F. Hoover 7314 (OBI, US): NOTES: Included in 
the Jepson Manual as P. elongatus Kunth, based on the 
two specimens cited above. Both of these recently re-de- 
termined by Robert Soreng (US) as P. imberbis. Hoover 
for San Luis Obispo Co. (1970) and especially Mason 
(1957) were uncomfortable with the P. elongatus deter- 
minations but did not suggest an alternative name. 

Pseudosasa japonica (Sieb. & Zucc. ex Steud.) Ma- 
kino ex Naka: DIST: CCo: CS: NCI: DOC: San Fran- 
cisco Co.: Howell, J.T. et al. (1958, pg. 43); Thomas, H. 
(1961, p. 75): NOTES: Spreading vegetatively from cul- 
tivation. 

Schedonnardus paniculatus (Nutt.) Trel.: DIST: CaR: 
CS: NCI: DOC: Siskiyou Co.: fenceline grass, Ager 
Road, Montague. Aug. 5, 1980, D.H. Shaw s.n. (AHUC, 
CDA). 

Spartina anglica C.K. Hubb.: DIST: CCo: CS: NW: 
DOC: Marin Co.: Creekside Park adjacent to Corte Ma- 
dera Creek, Greenbrae. Growing in low-flow channel of 
coastal salt marsh, assoc. with Distichlis spicata and Spar- 
tina densiflora. May 6, 2000, S. Klohr s.n. (CDA, DAY): 
NOTES: Only known site in California. More likely in- 
troduced to Creekside Park during marsh restoration work 
in 1977 sensu Spicher and Josselyn (1985) than via natural 
dispersal from Washington sensu Bossard et al. (2000). 

Stipa capensis Thunb.: DIST: DSon (Coachella Val- 
ley): CS: NW: DOC: Riverside Co.: Coachella Valley/ 
San Jacinto Mtns. Foothills. Alluvial fan of Chino Canyon 
along the Palm Springs tram road, N of the road 14 tele- 
phone poles above Hwy 111 (Palm Canyon Dr.), elev. 300 
m/950 ft, 33°51’N; 116°34'W, TO4S RO4E Sec. 04, SB. 
Creosote bush scrub on rocky alluvium cut by arroyos; 
with Larrea, Hymenoclea, Hyptis, Ambrosia, Justicia, 
etc., three patches seen, ca. 200—300 individuals. Mar. 19, 
1995, A.C. Sanders 16148, with G. Helmkamp, P. 
MacKay, et al. (UCR), det. by M. Barkworth; loc. cit. Apr. 
6, 1995, A. C. Sanders and M. Skinner 16393 (UCR); 
Coachella Valley, Chino Cyn., foothills of the San Jacinto 
Mtns., along the road to the Palm Springs Tram 1.5 mi 
above Hwy. 111. Palm Springs 7.5’ quad., 33°50'34"N; 
116°34'51”"W, TO04S, RO4E, Sec. 04, SB. Elev. 1200 ft/366 
m, rocky loam on alluvial fan, creosote bush scrub with 
Larrea, Ambrosia dumosa, Opuntia echinocarpa, Kra- 
meria grayi, etc., common annual on roadside and spread- 
ing into desert vegetation. Apr. 15, 2000, A.C. Sanders 
23321, with Giles Waines, Mitch Provance, T.B. Salvato, 
et al. (UCR); Cathedral Canyon, border of Rancho Mirage 
[and] Cathedral City, 33°45’N; 116°30’W. Mar. 11, 1997, 
Denise Woodard and Gilbert Goodlet s.n. (UCR), det. by 
A.C. Sanders; San Jacinto Mtns., S of Chino Canyon, at 
NW end of Palm Springs, 33°50'20"N; 116°33'45"W, 
TO4S, RO4E, Secs. 09 and 10, SB. Elev. 1148 ft/350 m, 
flat areas with Hyptis, Psorothamnus schottii, some creo- 
sote bush and smoke trees, locally abundant in disturbed 
places. Mar. 18, 1997, J. Wear and N. Moorhatch s.n. 
(UCR), det. by A.C. Sanders: NOTES: First records for 
California and North America of this annual Stipa with 
long awns and sharp callus tips. This species will be a 
severe nuisance if it becomes widely established. The 
seeds readily become caught in the fur of dogs and other 
animals and so will probably create veterinary problems, 
and will certainly be subject to ready dispersal. The very 
sharp callus can easily pierce human skin and cause un- 
pleasant sores. This plant is obviously a Stipa in the broad 


[Vol. 49 


sense, but its distinctly annual habit will quickly distin- 
guish it from all other known Californian Stipeae. Acc. to 
M. Barkworth, in the narrow taxonomic sense this plant 
is an Achnatherum, but the published combination in that 
genus by P. Beauv (Essai Agrostogr. 146) is invalid, hav- 
ing as its basionym Milium capense L. and not Stipa ca- 
pensis Thunb. Thus, in Achnatherum there is not currently 
an available epithet. 

Themeda quadrivalvis (L.) Kuntze: DIST: SCo: CS: 
TEN: DOC: San Bernardino Co.: City of Ontario; pri- 
vate residence, southeastern corner of yard beneath power 
lines. Nov. 24, 1991, T7.S. Ross 6026 (CDA, RSA). 

Tribolium obliterum (Hemzl.) Renvoize: DIST: CCo: 
CS: NW: DOC: Monterey Co.: well-established on sand 
dunes in scattered locations about the former Fort Ord 
Army base. June 6, 2000, B. Delgado, E. Finley, B. Oliver 
s.n. (CDA, DAV); Matthews, M.A. (1997, p. 341); loc. 
cit. April 17, 1990, V. Yadon H-3828 (PGM); loc. cit. 
April 14, 1998, V. Yadon H-3695 (PGM): NOTES: First 
collected by V. Yadon (PGM). Sieglingia decumbens (L.) 
Bernh. [=Danthonia decumbens (L.) DC.] misapplied. 
Reported in Matthews (1997) under that name. First re- 
cords for North America. Native to South Africa. 

Triticum aestivum L.: DIST: CA-FP: CS: C: DOC: 
Alameda Co.: Anthony Chabot Regional Park, archery 
range. June 2, 1981, J. Stratford s.n. (JEPS); Fresno Co.: 
Big Creek, 5000 ft. Aug. 9, 1956, H.M. Pollard s.n. (DAV, 
SBBG); Humboldt Co.: Arcata waste area near Com- 
munity center. Apr. 14, 2000, G. Leppig 1293 (CDA, 
HSC); Kern Co.: Twisselmann (1956, pg. 211); Los An- 
geles Co.: San Gabriel Mtns, Glendora Mtn. Rd., int. with 
Upper Monroe Rd. Rd. margin and adjacent vegetation. 
Locally naturalized. May 8, 1992, T. Ross 6305 (UC); 
Marin Co.: Penalosa, J., (1963, pg. 27); Howell, J.T. 
(1970, p. 74); Mendocino Co.: nr. Walkers Valley. May 
25, 1899, J. Burtt Davy and W.C. Blasdale 5062 (UC); 
Monterey Co.: roadside on Dolon Rd at Elkhorn Rd, May 
24, 2000, G. Leppig 1383 (CDA, HSC); San Francisco 
Co.: Howell, J.T. et al. (1958, pg. 45); San Luis Obispo 
Co.: Appearing from straw used to stabilize planted areas 
on sand dunes immed. W of Oso Flaco Lake. Elev. 150 
ft. Apr. 5, 1987, A.P. Griffiths 18086 (CDA, OBI); loc. 
cit. May 18, 1987, A.P. Griffiths 18187 (CDA, OBI); San 
Mateo Co.: McClintock, E., et al. (1990, pg. 184); Santa 
Barbara Co.: Santa Barbara, N of Botanic Garden, Mis- 
sion Cyn. Cr. June 19, 1944, C.F. Smith 945 (DAV, 
SBBG); Sonoma Co.: Best, C., et al. (1996, pg. 312): 
NOTES: The above list is not exhaustive. As a casual, 
this species is relatively commonly encountered and can 
be expected throughout California; it is likely known from 
every county. Probably the most common introduction 
vector is straw bales used for roadside erosion control, but 
also may be a contaminant or a component of seeding 
mixes. It is popular for use as the latter because of its 
usually ephemeral nature. Although individual sites rarely 
persist, new locations appear regularly. Naturalization has 
been reported however, as in Ross 6305 above (Los An- 
geles Co.), and its extent should be investigated further. 


Pontederiaceae 


Heteranthera rotundifolia (Kunth) Griseb.: DIST: 
ScV: CS: N: DOC: Butte Co.: Oswald, V.H. and L. Ahart, 
(1994, pg. 297); Yuba Co.: Damp soil in rice field, S of 
Woodruff Lane, just E of Mathews Rd, E of Sam Shin- 
taffer’s rice dryer, approx. 5 mi NE of Marysville. Com- 
mon. Elev. 80’. TI6N, RO4E, MD. Sept. 17, 1999, L. 
Ahart 8241 (CDA, CHSC). 


MAprRONO, Vol. 49, No. 2, pp. 99-114, 2002 


FIELD ASSESSMENT OF THE CALIFORNIA GAP ANALYSIS PROGRAM 
GIS DATABASE IN CENTRAL CALIFORNIA 


JOHN E KARLIK' EUGENE D. ALBERTSON 
University of California Cooperative Extension, 1031 S. Mt. Vernon, 
Bakersfield, CA 93307 


Y. JAE CHUNG 
U.S. Army Corps of Engineers, Los Angeles District, Regulatory Branch, 
PO. Box 532711, Los Angeles, CA 90053-2325 


ALISTAIR H. MCKAY 
University of California Cooperative Extension, 1031 S. Mt. Vernon, 
Bakersfield, CA 93307 


ARTHUR M. WINER 
Environmental Science and Engineering Program, 650 Young Drive, 
University of California, Los Angeles, CA 90095-1772 


ABSTRACT 


Given the key role played by biogenic volatile organic compounds (BVOCs) in photochemical smog 
formation and atmospheric chemistry, it is critical to generate accurate BVOC emission inventories. 
Assembling such inventories requires reliable characterization of the areal coverage of plant species to 
quantify the leaf mass of BVOC-emitting vegetation. A recent GIS-based description of vegetation cov- 
erage in the natural areas of California is provided by the Gap Analysis Program (GAP) database. We 
conducted an assessment of this database in Central California through ground-based vegetation surveys 
to evaluate the use of GAP for developing a BVOC emission inventory for Central California. A modified 
stratified randomized sampling design was used to select and assess 18 GAP polygons. Quantitative 
vegetation surveys were conducted along belt transects in polygons dominated by trees and along line 
transects in polygons dominated by shrubs to determine percent cover of plant species for comparison to 
GAP data. The species listed by GAP accounted for a range of 0—88% of the relative cover in the 
polygons, with a mean of 43%. Of the 76 species listed by GAP for primary, secondary and tertiary 
assemblages for which data were collected (those species above the survey height), 33 were found to be 
correctly listed within their respective assemblages, 13 were found to be listed for the wrong assemblage, 
and 30 were below percentages of co-dominants of any assemblage. In the 18 polygons, a total of 51 
additional species not listed by GAP were found to be present in amounts sufficient to consider them as 
potential co-dominants. Summed over all 18 polygons, BVOC emission indices based on field data were 
20% less than those based on GAP, but for individual polygons differences ranged from — 100% to more 


than + 100%. 
Key Words: 


BVOC emission inventories require data for 
emission rates, areal coverage, and leaf mass of re- 
spective plant species. With the proposal of a tax- 
onomic methodology for assigning isoprene and 
monoterpene emission rates to unmeasured plant 
species (Benjamin et al. 1996), emission rates can 
in principle be estimated for many of the 6,000 
plant species in California based on measurements 
within respective families and genera (Karlik and 
Winer 2001la; Winer and Karlik 2001). For Cali- 
fornia, vegetation spatial distribution and compo- 
sition has been described for urban and natural ar- 
eas within Orange County and the non-desert por- 
tions of Los Angeles, Riverside, and San Bernar- 
dino Counties (Winer et al. 1983: Miller and Winer 


' jfkarlik @ucdavis.edu 


Vegetation survey, biogenic hydrocarbons, Gap Analysis Program, GAP, BVOC. 


1984; Horie et al. 1991; Benjamin et al. 1997), and 
limited studies of plant composition have also been 
conducted for the Phoenix, AZ, urban area (Karlik 
and Winer 2001b) and the urban areas of Santa Bar- 
bara and Ventura Counties (Chinkin et al. 1996). 
However, a validated inventory of vegetation spe- 
cies composition and spatial distribution, specifi- 
cally to develop a BVOC emissions inventory, has 
not been established for the extensive areas of nat- 
ural vegetation in the San Joaquin Valley air basin. 

A potential source of information concerning 
vegetation in the natural areas of the Central Valley 
and the Sierra Nevada is the Gap Analysis Program 
(GAP) database, which is coordinated by the Unit- 
ed States Geological Service—Biological Resources 
Division (formerly the National Biological Service) 
to identify the distribution and management status 
of plant communities, especially to identify gaps in 


100 


habitats for plant or animal species needing protec- 
tion. GAP compiled a geographic information sys- 
tem (GIS) database (based primarily on remote- 
sensing data) describing vegetation type and dom- 
inance in terms of areal coverage (Davis et al. 
1994, 1995). Unlike other vegetation maps which 
describe plant geography only in terms of plant 
communities, the California GAP database de- 
scribes vegetation in quantitative terms using dom- 
inant plant species and species assemblages. 

Because BVOC emissions inventories rely on 
species-specific measurements of both leaf mass 
and BVOC emission rates (Benjamin et al. 1997), 
GAP offers the advantage of providing species-spe- 
cific vegetation distribution data. Moreover, the 
GAP GIS database is recent for California (Davis 
et al. 1995) and provides seamless coverage of the 
state. Although large-area small-scale GIS databas- 
es based on remote-sensing data, such as GAP, offer 
a potentially inexpensive and relatively simple ap- 
proach to characterizing the distribution and species 
identities of natural vegetation within an airshed, 
use of such GIS databases for BVOC emissions in- 
ventory development requires evaluation of their 
accuracy and reliability for this purpose through 
ground-based observations. 

The present GAP validation study was modeled 
after a previous study conducted in San Diego 
County (Winer et al. 1998; Chung and Winer 
1999). Quantitative vegetation surveys were con- 
ducted along belt transects in polygons dominated 
by trees, and along line transects in polygons dom- 
inated by shrubs, in order to determine percent cov- 
er of major plant species for comparison with GAP 
listings. 

We report here the results of a ground-based as- 
sessment of the GAP database within the Great Val- 
ley and Sierra Nevada ecological regions using 
vegetation surveys of representative GIS polygons. 
The surveys employed a modified stratified random 
sampling approach and a survey protocol based in 
part on the recommendations of the developers of 
the GAP database (Stoms et al. 1994), and refine- 
ments from the preceding study of GAP in San Di- 
ego County (Chung and Winer 1999). Data gath- 
ered from field surveys conducted during the 1999 
and 2000 summers were used to assess the accuracy 
and concomitant utility of the GAP GIS database 
for providing quantitative information of plant spe- 
cies identities and coverages for BVOC emission 
inventories. 


METHODS 
Acquisition and Preparation of the GAP Database 


As noted earlier, the purpose of GAP was to 
identify the distribution and management status of 
selected components of biodiversity. The central 
tool of this program was an ARC/INFO GIS data- 
base with plant species and vegetation class attri- 
butes associated with polygons within a defined 


MADRONO 


[Vol. 49 


geographic region. This database was generated 
from summer 1990 Landsat Thematic Mapper sat- 
ellite imagery, 1990 high altitude color infrared 
photography, vegetation type map surveys based on 
field surveys conducted between 1928 and 1940, 
and miscellaneous vegetation maps and ground sur- 
veys (Davis et al. 1995). Polygons were delimited 
based on climate, physiography, substrate, and dis- 
turbance regime. Landscape boundaries were sub- 
jectively determined through photointerpretation by 
expert personnel so that between-polygon variation 
was greater than within-polygon variation. The final 
result was a vegetation map with a 100 ha mini- 
mum mapping unit and a 1:100,000 mapping scale 
(Davis et al. 1995). 

The GAP database for the southern portion of the 
Sierra Nevada and Great Valley ecological regions 
was obtained at the beginning of the project. Seven 
hundred forty-two polygons in the Great Valley and 
1420 polygons in the Sierra Nevada ecological re- 
gions were found within the counties of Kern, Tu- 
lare, Kings, Fresno, and Madera. For each polygon 
in the database, a primary vegetation assemblage 
was listed. For most polygons a secondary vege- 
tation assemblage was listed, and for some poly- 
gons a tertiary vegetation assemblage was also giv- 
en. The primary assemblage was defined as the as- 
semblage covering the majority of the polygon, and 
the secondary and tertiary assemblages as covering 
relatively smaller areas of the polygon. Each as- 
semblage consisted of up to three co-dominant 
overstory species, each with a minimum of 20% of 
the relative cover of that assemblage. Relative cov- 
er of a given plant species within an assemblage 
was the fraction of total assemblage vegetation cov- 
er occupied by the given species. 


Polygon Selection 


GAP data for each polygon were used to gener- 
ate an isoprene and monoterpene index for the 
polygon (Winer et al. 1998; Chung and Winer 
1999). Although polygons with high emissions in- 
dices may hold more interest for emissions mod- 
elers, after lengthy discussions and review of com- 
ments on a previous study design for GAP (Winer 
et al. 1998; Chung and Winer 1999), we decided to 
use a random selection process rather than focusing 
on “‘high emitting”’ polygons for field validation in 
this study. To remain a candidate for field valida- 
tion, the polygon had to be below the atmospheric 
boundary layer, taken as 1800 m elevation, and 
within the San Joaquin Valley air basin. Further se- 
lection from the remaining polygons involved an 
iterative process accounting for feasibility, includ- 
ing physical access and permission to survey veg- 
etation. A road map was overlaid on the area to see 
if there was access by roadways, and if so a uni- 
versal transmercator (UTM) grid was generated. 
Polygons with a large public land component (e.g., 
within National Forest) were favored due to the rel- 


2002] KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 101 
TABLE 1. POLYGONS FROM THE GREAT VALLEY AND SIERRA NEVADA GAP DATABASE SELECTED FOR FIELD SURVEY. 
robes Approximate Area Elevation Transect Centerpt. Transect 
(no.) GAP No. location (ha) (m) Type (no.) (no.) 
1 16541 Kern National Wildlife oper 66 Line 4 16 
Refuge 
2 17198 Kern River & Hwy 99 284 120 Belt 3 8 
3 15485 Lemoore 256 64 Line 3 11 
4 17023 Buttonwillow 254 82 Line 2 8 
5 16442 Glennville 1383 1520 Belt 3 12 
6 16753 Keysville 195 1570 Belt 3 12 
Hy 16908 Kern River Canyon 640 760 Belt e: 12, 
8 14572 Sequoia National Park 4431 1280 Belt 3 12 
9 16791 Bodfish 309 1200 Belt 3 12 
10 16273 California Hot Springs 2688 1570 Belt 4 14 
11 13790 Sequoia National Park 635 1270 Belt 2 8 
12 13795 Kings Canyon 2473 1340 Belt 1 2 
13 16269 California Hot Springs 341 930 Belt 2 6 
14 16687 Lake Isabella 1019 845 Belt 3 f2 
15 16783 Alta Sierra 476 1060 Belt 2, 8 
16 16776 Alta Sierra 642 1340 Belt 2 4 
17 16756 Alta Sierra 1969 1080 Belt 2 4 
18 16735 Alta Sierra 812 1030 Belt 2 4 


ative ease of gaining permission to conduct surveys 
on such properties compared to privately owned 
properties. 

Based on these criteria and the time and resourc- 
es available for this research, 18 polygons in south 
central California were selected and surveyed for 
the present study, as seen in Table 1. Four polygons 
(nos. 1—4), located on the Central Valley floor, con- 
sisted primarily of shrubs and herbaceous vegeta- 
tion, and fourteen polygons (nos. 5—18), located in 
the Sierra Nevada mountains and foothills, consist- 
ed primarily of woodland and forest vegetation 
thie. ft): 

A subsample of polygons was selected as a test 
for correctness of the geographic location of a spe- 
cific GAP polygon; in other words, a test of the 
registration of the GAP database. Three surround- 
ing polygons adjacent to polygon 15 were selected 
for survey during the summer, 2000, sampling sea- 
son. The plants found in these surrounding poly- 
gons could then be compared with those listed for 
the center polygon to see if plant communities list- 
ed for the center polygon were found instead in a 
surrounding polygon. The three surrounding poly- 
gons were no. 16 located to the southeast, no. 17 
located on the western end, and no. 18 located to 
the northeast of polygon 15. 


Selection of Sample Elements 


If permission was obtained to access most of a 
polygon, sample elements of 500 * 500 m squares 
were selected by overlaying a 500 X 500 m UTM 
grid on the polygon, assigning sequential numbers 
to every grid square within | km of a road, and 
randomly selecting the centerpoint locations for the 
needed number of elements. The number of center- 
points and corresponding elements varied with 


polygon size. For polygon areas of <1000, 1000— 
10,000, and >10,000 ha, two, three, and four cen- 
terpoints were chosen, respectively, although ter- 
rain or accessibility sometimes limited the number 
of centerpoints. This method was similar to the one 
employed in the Utah GAP validation project (Ed- 
wards et al. 1995) and that of Chung and Winer in 
San Diego County (1999). In several cases, suitable 
survey sites were not available within the vicinity 
of a road, so hikes of up to two hours along a trail 
were needed to reach the desired area within the 


polygon. 


Vegetation Survey Protocol 


The specific survey protocol chosen depended on 
the type of vegetation being assessed. Within the 
polygons dominated by trees, surveys were per- 
formed by a team of two along 6 m wide, 500 m 
long belt transects orthogonal at the centerpoint in 
most elements. Six meter wide belt transects make 
the mechanics of sampling easier while not signif- 
icantly compromising accuracy (Lindsey 1955). For 
these belt transects, the surveyors walked 250 m 
north, south, east, and west away from the center- 
point, using a magnetic compass to maintain 
course. 

Within polygons dominated by shrubs, the sur- 
vey for each element consisted of two 300 m line 
transects orthogonal at the centerpoint. Line tran- 
sects have been used to estimate relative cover for 
chaparral (Bauer 1943) and for sage scrub (Kent 
and Coker 1992; Zippin and Vanderwier 1994). The 
minimum square-shaped area needed to encompass 
a sample element within a polygon was therefore 
25 ha for forests and woodlands, and nine ha for 
scrub and chaparral. 

The survey team located the centerpoint of a par- 


102 MADRONO 


Location of 
Great Valley polygons 
Nos. 1-4 


Fic. 1. 


ticular sample element using a global positioning 
receiver (GPS) locked onto UTM coordinates gath- 
ered from the GAP database. A handheld GPS unit 
(Garmin 12XL), with an accuracy of approximately 
22115 im Gr 22D mn, wa ISLS) ancl AVOO, mesoecinvelby, 
was employed. Plant community and site descrip- 
tions were recorded and elevation at the centerpoint 
was determined using a hand-held altimeter (Pretel 
Instruments). 


Data Collection 


For belt transects, data collected included the 
crown radii, crown height, and diameter at breast 
height of trees and the crown dimensions of shrubs. 
For line transects, plant species identity, crown 
height and number of 0.1 m segments along a meter 
tape occupied by that plant species were noted. 
Plants such as grasses and forbs below a height of 
about 0.6 m were not recorded. Additional details 
of plant measurement methods have been reported 
previously (Winer et al. 1998; Chung and Winer 
1999). Plant nomenclature follows Mabberley 
Geom: 


[Vol. 49 


Location of 
Great Valley polygons 
Nos. 5-18 


Locations of the polygons selected for field survey from the GAP database. 


Data Analysis 


Data analysis followed the example of Chung 
and Winer (1999). For each polygon, the GAP da- 
tabase listed primary, secondary, and sometimes 
tertiary species assemblages and the estimated areal 
proportion (p) of each assemblage within a poly- 
gon. Each species in a listed assemblage was a co- 
dominant, providing =20% relative cover within 
the assemblage. Therefore, the expected coverage 
of any species listed in the GAP database for a 
given polygon was =0.2p. For example, in polygon 
5 Quercus kelloggi was listed as a co-dominant in 
a primary assemblage that occupied 60—70% of the 
polygon. Using a mean value of 65% for p, GAP 
predicted Quercus kelloggi would cover 20.2 X 
65%, or 213% of the polygon. 

The polygon coverage of plant species inferred 
from the GAP database by this procedure was com- 
pared with the cover data gathered from the field 
surveys in the 18 selected polygons. First, the cov- 
erage of each species within each sample element 
of a polygon was calculated. Then from the species 
coverage for each sample element, the mean cov- 


2002] 


erage and upper limit of the two standard error (SE) 
confidence intervals for the polygon were calculat- 
ed, corresponding to an 85% confidence interval 
(McClave and Dietrich 1985). 


RESULTS 


Species Composition and Abundance within 
GAP Polygons 


Table 2 summarizes results for the 18 polygons 
surveyed, listing the most abundant species ob- 
served for each polygon, the percent abundance 
predicted from the GAP database, the percent abun- 
dance determined by the field surveys, and the up- 
per limits of a two SE interval of the percent com- 
position. Plant species not listed by GAP or not 
found in surveys at =1% are omitted from this ta- 
ble, but may be found in the report of Winer and 
Karlik (2001). Total plant cover within the polygon 
ranged from as little as 7%, as found in polygons 
2 and 9, up to 82% as found in polygon 12. In 
general, most of the sample cover in a polygon was 
attributable to a few species and many of the most 
abundant species found within the polygon were 
listed as co-dominants by the GAP database. How- 
ever, the percentages of these GAP co-dominant 
species varied greatly. Total cover of GAP co-dom- 
inant species cover ranged from as little as 0% as 
found in polygon 18 up to 66% as found in polygon 
2: 

Relative cover of GAP-listed species compared 
to sampled species can be derived from the per- 
centages listed in Table 2. For the Great Valley 
polygons (nos. 1—4), GAP species listings ranged 
from 29-72% of the plant species found. The sum 
of GAP species percentages for these polygons was 
46, and for all species sampled, 100. Thus, GAP- 
listed plants accounted for 46% of the relative cov- 
er for polygons nos. 1—4 considered together. For 
the Sierra Nevada polygons (nos. 5—18), the rela- 
tive cover of GAP-listed species for each polygon 
ranged from O—88%, and the relative cover was 
43% for polygons nos. 5—18 considered together. If 
all 18 polygons were considered together, GAP-list- 
ed plants accounted for 43% of the relative cover 
overall. To investigate whether GAP listings were 
more accurate for large vs smaller polygons, poly- 
gon size was multiplied by the corresponding per- 
centages found for GAP-listed species vs for all 
species. The results suggested polygon size did not 
influence accuracy of GAP listings, since relative 
cover of GAP species was then found also to be 
43%. 

The observed sample cover of some co-domi- 
nants in GAP polygons often substantially exceed- 
ed the minimum predicted values. For example, in 
polygon 12 Quercus chrysolepis provided 51% of 
the polygon sample cover though =7% and =3% 
were predicted from the GAP listings for the sec- 
ondary and tertiary assemblages, respectively. In 
polygon 5 Quercus kelloggii provided 31% of the 


KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 103 


polygon sample cover when =13% was predicted 
by GAP. 

In contrast, several polygons possessed co-dom- 
inant species that were found to be well under the 
predicted GAP percentages. For example, polygon 
9 was found to have 1% mean sample cover of 
Yucca whipplei when GAP predicted the species to 
have 211% and =7% sample cover for the primary 
and secondary assemblages, respectively. In poly- 
gon 12, Pinus ponderosa was predicted by GAP to 
be found with sample cover =11% when the field 
study found it to be only 0.3% sample cover. 

Because Quercus may be the most important ge- 
nus of native woody plants with regard to BVOC 
emissions in California’s airsheds, we compared the 
percentage of all oak species sampled to that in- 
ferred from GAP listings. For the 14 Sierra Nevada 
polygons (nos. 5—18), the mean coverage per poly- 
gon of oaks calculated from GAP listings was 17%, 
while from our field surveys oaks averaged 29% of 
the total cover. For all 18 polygons, the mean cov- 
erage of oaks calculated from GAP per polygon 
was 13% and from field surveys 22%. Since GAP 
listings give a lower limit for species abundances, 
one might expect to find oaks at the same or higher 
percentages than GAP listings. Considering poly- 
gons individually, for six of the 18 polygons we 
found oak species percentages within a factor of 
two of the respective GAP-predicted percentages. 
For an additional five polygons, oaks were listed as 
comprising zero percent cover, and were not found 
in the field surveys, in agreement with GAP list- 
ings. Thus, for 11 of 18 polygons, GAP listings for 
oaks were in reasonably good agreement with field 
data. For three polygons, nos. 8, 14, and 18, oaks 
were found at 20, 4, and 43% although no oak spe- 
cies were listed by GAP. Thus, overall GAP data 
were in reasonable agreement with field data, and 
the lower limit of oak coverages as given by GAP 
may underestimate oak abundance. 

Since Quercus species vary over almost two or- 
ders of magnitude in isoprene emission rate (Csiky 
and Seufert 1999), the accuracy of oak species list- 
ings for any vegetation database is important for 
BVOC emission inventories. Using the data in Ta- 
ble 2, we considered oak species accuracy for each 
polygon by calculating the sum of absolute values 
of field percentage found for each oak species mi- 
nus the respective percentage inferred from its GAP 
listing. These values for polygons ranged from 0% 
for the polygons where no oaks were listed or 
found to more than 40% for polygons 12, 15, 17, 
and 18 with a mean for all polygons of 19%. The 
accuracy of oak species listings did not appear to 
be related to polygon size. 

We also evaluated the data in Table 2 with regard 
to non-oak genera and species considered to be 
moderate or high BVOC emitters, specifically those 
thought to have emission rates greater or equal to 
10 or 2 pg g-' h'' for isoprene and monoterpenes, 
respectively, as based on measured values or tax- 


104 MADRONO [Vol. 49 


TABLE 2. MEASURED SPECIES COVER COMPOSITION OBSERVED IN SAMPLED GAP POLYGONS LISTED IN ORDER OF OBSERVED 
SAMPLED COVER. Species with mean sampled cover of <1% or not listed in GAP are omitted. N.D. = no data. For 
example, for Avena spp. and Bromus spp. species were observed but below survey height (about 0.6 m), and data were 
not recorded. 


Sampled 
cover 
GAP predicted Mean sampled (s + 2SE) 
Polygon Species cover (%) cover (%) (%) 
1 Cyperus difformis — L/ 30 
Typha spp. =11 & =3 15 38 
Scirpus acutus —- 6 17 
Xanthium strumarium — 4 8 
Brassica nigra — 3 7 
Distichlis spicata — 3 6 
Scirpus californicus — 1 3 
Baccharis salicifolia _ 1 2 
Suaeda ramossissima —_ 1 2 
Atriplex coronata — 1 2 
Allenrolfea occidentalis 27] 1 22 
Salix spp. =] 0.2 0.6 
Carex spp. 22\lil 6 228 0.0 — 
Juncus spp. == 0.0 — 
Tamarix spp. == WSO =3 0.0 — 
Total of sample cover 3S) 
GAP co-dominants 16 
2 Unknown #1 — 3 7 
Populus fremontii = D 3 
Platanus racemosa -— i D; 
Salix spp. — 1 2 
Distichlis spicata 21/3) eG, ei) N.D. — 
Total of sample cover 7 
GAP co-dominants 2 
3 Allenrolfea occidentalis 22117 Go 223 21 43 
Salix sp. — 9 Da 
Populus fremontii = D, 5 
Eucalyptus spp. =3 0.0 = 
Total of sample cover 32 
GAP co-dominants 3 
4 Atriplex polycarpa 22 (9) 5) 9 
Adenostoma fasciculatum — 2 6 
Avena spp. and Bromus spp. 220) N.D. — 
Total of sample cover 8 
GAP co-dominants 5 
5 Quercus kelloggii =13 Syl 4] 
Quercus wislizenii 2213) 6's 227/ 14 19 
Quercus garryana =] 6 sI97/ 
Calocedrus decurrens _- =) 9 
Cercocarpus betuloides =|) 5 8 
Abies concolor 2 5) 
Pinus ponderosa =13 1 2, 
Quercus berberidifolia ] 1 
Total of sample cover 65 
GAP co-dominants Si 
6 Quercus kelloggii NB ce 227) 14 ES) 
Pinus ponderosa 22/3) (6 227) 6 13 
Arctostaphylos spp. — =) 11 
Quercus chrysolepis — 4 10 
Abies magnifica —- 3 6 
Calocedrus decurrens — 2 7 
Abies concolor =13 1 3 
Cercocarpus betuloides — ] 2 
Quercus wislizenii =7 ] 1 
Total of sample cover 38 
GAP co-dominants DD, 


2002] 


Polygon 
I 


10 


11 


KARLIK ET AL 


Species 


Quercus wislizenii 
Quercus douglasii 
Rhamus crocea 
Ceanothus cuneatus 
Quercus garryana 
Platanus racemosa 
Pinus sabiniana 

Quercus dumosa 
Aesculus californica 
Adenostoma fasciculatum 


.. GAP ASSESSMENT IN CENTRAL CALIFORNIA 


TABLE 2. CONTINUED. 


GAP predicted 
cover (%) 


= 7/ 


Avena spp. and Bromus spp. =11 


Total of sample cover 
GAP co-dominants 


Abies concolor 

Quercus douglasii 
Quercus kelloggii 
Calocedrus decurrens 
Ceanothus integerrimus 
Aesculus californica 
Quercus chrysolepis 
Pinus ponderosa 

Pinus lambertiana 
Cornus nuttallii 
Sequoiadendron gigantea 
Umbellularia californica 
Adenostoma fasciculatum 


Total of sample cover 
GAP co-dominants 


Chrysothamnus nauseosus 
Juniperus californica 
Yucca whipplei 
Eriogonum fasciculatum 
Artemisia tridentata 


Total of sample cover 
GAP co-dominants 


Quercus douglasii 
Aesculus californica 
Quercus wislizenii 
Quercus kelloggii 
Quercus garryana 
Quercus chrysolepis 
Cercocarpus betuloides 
Pinus sabiniana 

Rhus diversiloba 
Ceanothus cuneatus 
Ribes sp. 

Total of sample cover 
GAP co-dominants 


Cercocarpus betuloides 
Ceanothus integerrimus 
Ceanothus cuneatus 
Quercus dumosa 
Aesculus californica 
Quercus chrysolepis 
Arctostaphylos sp. 
Umbellularia californica 
Arctostaphylos mewukka 


Total of sample cover 
GAP co-dominants 


Mean sampled 
cover (%) 


io) 


ONFRRFN WW ORR KYPNWWHW HK OW WO 


oo 


~— 


105 


Sampled 
cover 
(S—Z2SB) 
(%) 


Wore BWW BND WW 


© 


106 


Polygon 
12 


13 


IS) 


16 


7 


Species 


Quercus chrysolepis 
Calocedrus decurrens 
Umbellularia californica 
Cercocarpus betuloides 
Ceanothus cuneatus 
Quercus dumosa 
Arctostaphylos sp. 
Salix spp. 

Pinus ponderosa 

Abies concolor 
Aesculus californica 
Pinus contorta 


Total of sample cover 
GAP co-dominants 


Quercus wislizenii 

Aesculus californica 

Quercus douglasii 

Ceanothus cuneatus 

Quercus dumosa 

Umbellularia californica 
Fremontodendron californicum 
Cercocarpus betuloides 

Pinus sabiniana 

Avena spp. and Bromus spp. 


Total of sample cover 
GAP co-dominants 


Ceanothus cuneatus 
Pinus sabiniana 

Quercus douglasii 
Quercus wislizenii 
Quercus dumosa 
Ephedra california 
Mimulus aurantiacus 
Yucca whipplei 
Adenostoma fasciculatum 
Juniperus californica 
Artemisia tridentata 
Avena spp. and Bromus spp. 


Total of sample cover 
GAP co-dominants 
Quercus kelloggi 
Quercus douglasii 
Quercus garryana 
Quercus dumosa 
Aesculus californica 
Quercus wislizenii 
Ceanothus cuneatus 
Pinus sabiniana 
Cercocarpus betuloides 
Pinus ponderosa 
Avena spp. and Bromus spp. 


Total of sample cover 
GAP co-dominants 


Ceanothus cuneatus 
Quercus lobata 
Quercus douglasii 
Aesculus californica 
Quercus wislizenii 
Quercus kelloggii 
Quercus garryana 


MADRONO 


TABLE 2. CONTINUED. 


GAP predicted 


cover (%) 


=7 & =3 
=11 


Mean sampled 
cover (%) 


[Vol. 49 


Sampled 
cover 
(s + 2SE) 
(%) 


2002] 


KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 107 


TABLE 2. CONTINUED. 


Polygon Species 


Quercus dumosa 
Ribes sp. 
Pinus sabiniana 
Cercocarpus betuloides 
Avena spp. and Bromus spp. 
Total of sample cover 
GAP co-dominants 

18 Quercus douglasii 
Quercus wislizenii 
Aesculus californica 
Ceanothus cuneatus 
Quercus dumosa 
Pinus sabiniana 
Cercocarpus betuloides 
Adenostoma fasciculatum 
Unidentified chaparral shrubs 
Bare exposed rocks 
Total of sample cover 
GAP co-dominants 


onomy (Benjamin et al. 1996; Karlik and Winer 
2001a). For the 11 polygons where such genera or 
species were listed, the field surveys gave percent- 
ages lower than those inferred from GAP for seven 
polygons, and higher than GAP for four polygons. 
For the seven polygons where no genera or species 
with moderate or high emission rates were listed, 
the field surveys found an absence of such genera 
or species in two polygons, and for the other five 
polygons percentages of emitting species ranged 
from 1—11% of total area. In considering the mean 
of GAP listings for BVOC-emitting species vs the 
mean of the field survey data for all 18 polygons, 
GAP predicted a mean coverage per polygon of 9% 
for these species, and field data were in good agree- 
ment with a mean of 8% of such species found. 
Thus, on average GAP data were harmonious over- 
all with field observations for non-oak genera and 
species considered to be important BVOC emitters. 


Correctness of GAP Listed Species within 
Species Assemblages 


Species found within the polygons in the field 
were compared to their GAP listings and assessed 
for correct placement based on assemblage data, 
and the results are given in Table 3. A species was 
considered to be correctly listed when the percent- 
age found in the field within two standard errors (s 
+ 2SE, Table 2) exceeded the GAP-predicted per- 
centage. A species was considered listed incorrectly 
when listed by GAP as a co-dominant in a partic- 
ular assemblage (primary, secondary, and tertiary) 
but found at a lower percentage cover so as to place 
it within a different assemblage. Potential co-dom- 
inant species were defined as species not listed by 


Sampled 
cover 
GAP predicted Mean sampled (S)-- 2SB) 
cover (%) cover (%) (%) 
— 4 6 
— 2 6 
— p? 6 
— 1 1 
=>3 N.D. — 
62 
8 
= 25 40 
— 15 40 
— 9 Dal 
— 9 9 
— 3 9 
— 1 BY 
— 1 2 
=13 0.0 — 
=5 N.D — 
>3 N.D — 
63 
0) 


GAP as present in the polygon but found in field 
surveys to have cover percentage large enough to 
at least fall within the tertiary assemblage of a par- 
ticular polygon. When GAP listed no species for 
the secondary or tertiary assemblage, an arbitrary 
value of =7% and =3% up to the next greater listed 
assemblage percentage were assigned, respectively, 
to identify potential species belonging to a partic- 
ular assemblage. We note that GAP assigns mini- 
mum percentages to plant species coverages, but 
does not assign maxima; therefore, species listed in 
two or more coverage classes were considered to 
have a correct listing in these classes if present in 
sufficient quantity for the greater percentage re- 
quirement. For example, in polygon 6 the species 
Quercus kelloggii was listed in both primary and 
secondary assemblages, found to be present at 
14%, and considered to be correctly listed for both 
assemblages. 

The agreement of field results with GAP data 
varied among polygons, as seen in Table 3. There 
were several polygons (nos. 3, 5, 6, 10, 13) in 
which the field results agreed with all or the ma- 
jority of GAP listings. In contrast, there were sev- 
eral instances where species listed by GAP in either 
the primary, secondary, or tertiary assemblage were 
not observed in the polygon in sufficient abundance 
for for their respective assemblage. For example, in 
polygon 6 Abies concolor in the primary assem- 
blage and in polygon 7 Quercus douglasii in the 
secondary assemblage were found with coverage 
percentages below those for their respective assem- 
blages. Potential co-dominant species were noted in 
several polygons, such as in polygons 8 and 17, 
where seven species were found in percentages that 


[Vol. 49 


MADRONO 


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112 MADRONO 


warranted co-dominant species designation, al- 
though none of these seven species was listed by 
GAP in any assemblage within the polygon. 

Overall, as seen in Table 3, of the 76 species 
listed by GAP for primary, secondary and tertiary 
assemblages for which data were collected (those 
species above the survey height), 33 were found to 
be correctly listed within their respective assem- 
blages, 13 were found to be incorrectly listed, and 
30 were found to be below percentages of all co- 
dominants. Of these 30, 15 were not observed in 
the field at all. In the 18 polygons, six species not 
listed by GAP in any assemblage (within respective 
polygons) were found in field surveys to be poten- 
tial primary species, 22 were found to be second- 
ary, and 23 tertiary, for a total of 51 additional 
species not listed by GAP but found to be present 
in cover sufficient to be considered as potential co- 
dominants. The Great Valley polygons (nos. 1—4) 
did not seem to differ from the Sierra Nevada poly- 
gons (nos. 5—18) in accuracy of listings. Unlike the 
results of Chung and Winer (1999), we found the 
accuracy of listings of primary, secondary, and ter- 
tiary species was about the same (~40%) with, re- 
spectively, 14 of 32, 15 of 33, and 4 of 11 plants 
listed correctly. 

Considering specifically data for oak species in 
Table 3, for the 18 polygons and for the 25 oak 
species listed, 20 were found to be listed correctly 
in their respective assemblages, three were listed 
incorrectly, and two were found to be below per- 
centages for any co-dominant. Three additional oak 
species not listed by GAP were found to be poten- 
tial primary co-dominants, 10 as secondary co- 
dominants, and nine as tertiary co-dominants. Thus, 
GAP listings for oaks were in good agreement with 
field data, but an additional 22 examples were 
found of oak species present at percentages large 
enough for inclusion as GAP co-dominants. Of 
these latter, the presence of two oak species in poly- 
gon 18 in percentages high enough to consider 
them primary suggests a significant discrepancy in 
cover type for that polygon between the listed spe- 
cies and field data. These results also indicate oaks 
may be found with greater spatial extent than in- 
ferred from the GAP database. 

We also examined listings in Table 3 of genera 
and species, other than oaks, considered to be me- 
dium or high emitters of isoprene or monoterpenes. 
For the 18 polygons, and for the 14 listings for 
plants such as Salix spp. and Populus spp., three 
were found to be correct, four were incorrect, and 
seven were found to be below percentages of co- 
dominants. Field survey data indicated one species 
was found which could be considered a primary co- 
dominant, six could be secondary, and four tertiary. 
Therefore, 11 species of the medium or high emit- 
ters, were found at low percentages compared to 
their respective GAP listings, but 11 species not 
listed were found to be potential co-dominants, 


[Vol. 49 


which seem to be canceling errors in the database 
from the perspective of BVOC emissions. 

As discussed earlier, the polygons surrounding 
polygon 15 were surveyed to test the GAP database 
for a possible registration error. The vegetation cov- 
er in polygons 16, 17, and 18 found in the field 
survey did not appear to match the GAP listings 
for polygon 15 any more closely than did the veg- 
etation cover within the boundaries of polygon 15. 
Hence, in this case we found no evidence of mis- 
registration. 


Implications of GAP Assessment Results for 
BVOC Emission Inventories in California 


The primary purpose for GAP is to identify the 
distribution and management status of plant com- 
munities, rather than to identify individual plant 
species. The quantitative nature of GAP represents 
an advance in landcover classification and the val- 
ues for plant cover and species percentages give an 
indication of the composition of plant communities. 
However, the GAP database is fundamentally about 
plant assemblages rather than species, and these as- 
semblages may vary in precise composition de- 
pending on geographic and environmental factors. 
In addition, a component of leaf mass, which the 
GAP database does not provide, must be overlaid 
on the species distribution data for BVOC emission 
calculation. Thus, the applicability of GAP for 
BVOC modeling requires ongoing discussion. 

We calculated isoprene emission indices (as de- 
scribed earlier) based on GAP data and found they 
differed less than +50% from the corresponding 
indices calculated from our field data for half the 
polygons surveyed. Isoprene emission indices 
summed over all 18 polygons based on GAP data 
were in good agreement with the sum of corre- 
sponding emission indices generated with data from 
field surveys, with a difference of only —6% using 
field survey data vs GAP data. Results were not as 
consistent for monoterpenes. Ten of 18 polygons 
had monoterpene emission indices based on field 
survey data differing by less than 100% from those 
generated with GAP data, and the sum of mono- 
terpene emission indices based on field surveys was 
78% less than the sum calculated from GAP data. 
When field data were used in place of GAP data 
the sum of total isoprene and monoterpene emis- 
sion indices dropped by 20%, and the change for 
individual polygons ranged from —100% to more 
than + 100%. These percentage changes for the sum 
of emission indices were greater than those found 
in polygons assessed in San Diego (Chung and Wi- 
ner 1999), 

Compared to previous databases estimating per- 
cent cover of vegetation in natural areas, the GAP 
database is species-specific and has a higher spatial 
resolution. Results of this study, and that of Chung 
and Winer (1999), indicate GAP may be useful for 
assigning species identities to plant cover in the 


2002] 


natural areas of California airsheds for BVOC in- 
ventory development. However, GAP should be 
used for this purpose with caution, as there can be 
discrepancies in species listings for individual poly- 
gons resulting in considerable differences in esti- 
mated BVOC emissions. 


Limitations of the Present Study 


GAP assessment in the southern Central Valley 
and surrounding mountains posed special problems 
in terms of sampling representative areas within 
privately owned parts of a polygon. In the Utah 
GAP validation project, 42% of the state was under 
the control of the US Bureau of Land Management, 
with private interests owning 21% (Edwards et al. 
1995). In the study of Chung and Winer (1999), the 
San Diego County Association of Government 
1990 ownership database indicated private interests 
owned 41% of San Diego County land. In the pre- 
sent GAP assessment project, suitable public lands 
within the vicinity of roads were limited, resulting 
in extended hikes from established roads to reach 
them. Even with such effort, our ability to conduct 
surveys in representative areas of a polygon’s major 
vegetation types as listed in the GAP database was 
limited. 

Given the effort needed to gather the field data, 
it was necessary to limit the number of polygons 
assessed and the area sampled. Moreover, the sam- 
ple area required for estimating the true sample 
cover of individual species in a polygon is not pre- 
cisely known. One reference (Bormann 1953) sug- 
gested that surveying 7% of a forested area using 
parallel belt transects provided a 65% chance the 
sample mean of the basal area of the trees would 
be within 10% of the true mean for more common 
species. The effort needed to obtain an accurate 
measure of relative cover may be similar. In the 
present study, each sample element for belt tran- 
sects occupied 0.6 ha, so for a polygon of 500 ha, 
two sample elements encompassing 1.2 ha were 
surveyed, or about 0.24% of the polygon area. For 
line transects, two sample elements in a 500 ha 
polygon would occupy about 1200 m/’, or about 
0.024% of the polygon. 

On the other hand, the effective size of the sam- 
ples may be larger. The vegetation cover compo- 
sition within the transects may approximate the 
cover composition of a square which immediately 
bounds the ends of the perpendicular transects. In 
that case the percentage of the polygon area sam- 
pled would be 10% and 3.6% of a 500 ha polygon 
for belt and line transects, respectively. 


CONCLUSIONS 


A ground-based assessment of the GAP database 
for the Great Valley and Sierra Nevada ecoregions 
of California was conducted in the southern San 
Joaquin Valley and adjacent mountains to evaluate 
use of GAP in developing a BVOC emission in- 


KARLIK ET AL.: GAP ASSESSMENT IN CENTRAL CALIFORNIA 113 


ventory for Central California. The species listed 
by GAP accounted for a range of 0 to 88% of the 
relative cover in the polygons, with a mean of 43%. 
Of the 76 species listed by GAP for primary, sec- 
ondary and tertiary assemblages for which data 
were collected (those species above the survey 
height), 33 were found to be correctly listed within 
their respective assemblages, 13 were found to be 
listed for the wrong assemblage, and 30 were below 
percentages of co-dominants of any assemblage. In 
the 18 polygons, a total of 51 additional species not 
listed by GAP were found to be present in amounts 
sufficient to consider them as potential co-domi- 
nants. However, the listings of oak species and oth- 
ers considered to be important due to their magni- 
tudes of biogenic emissions were generally in good 
agreement with field data. Summed over all 18 
polygons, BVOC emission indices based on field 
data were 20% less than those based on GAP, but 
for individual polygons the differences ranged from 
— 100% to more than + 100%. Registration error did 
not seem to be the cause of discrepancies between 
listed and field data. Thus, this database should be 
used with caution for developing BVOC invento- 
ries. Other databases more limited in geographic 
coverage may also be useful, and should be vali- 
dated for accuracy against field data, particularly 
for representativeness of species of interest. 


ACKNOWLEDGMENTS 


We thank the field technicians assigned to this project 
who traversed many canyons and mountain slopes to mea- 
sure plant cover, including Matthew Bates, Joseph Loeh- 
ner, Jason Robbins, and Jason Welch. We appreciate the 
cooperation of the U.S. National Park Service in providing 
no-cost access to Sequoia and Kings Canyon national 
parks for vegetation sampling, and thank the Kern Na- 
tional Wildlife Refuge for similar consideration. We ac- 
knowledge the many individual private property owners 
who permitted access to their land. Officials of the USDA 
Forest Service and the Bureau of Land Management fa- 
cilitated plant surveys within lands under their jurisdic- 
tion. 

We gratefully acknowledge the support of the Califor- 
nia Air Resources Board (Contract No. 97-320) for this 
research, and the assistance of Dr. Ash Lashgari and Dr. 
Michael Benjamin of that agency. The statements and 
conclusions in this article are those of the authors and not 
necessarily those of the California Air Resources Board. 


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MADRONO 


[Vol. 49 


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MApRONO, Vol. 49, No. 2, pp. 115-121, 2002 


SOME FACTORS INFLUENCING SEEDLING DENSITY OF CALIFORNIA 
BLACK OAK (QUERCUS KELLOGGID IN THE CENTRAL 
SIERRA NEVADA, CALIFORNIA 


BARRETT A. GARRISON 
California Department of Fish and Game, 
Sacramento Valley—Central Sierra Region, 1701 Nimbus Road, 
Rancho Cordova, CA 95670 
bagarris @dfg.ca.gov 


RoBIN L. WAcHS, JAMES S. JONES,! AND MATTHEW L. TRIGGS 
U.S. Forest Service, Tahoe National Forest, Foresthill Ranger District, 
22380 Foresthill Road, Foresthill, CA 95631 


ABSTRACT 


Seedling densities of California black oak (Quercus kelloggii) and overstory and understory vegetation 
attributes were quantified on four 21.1-ha study stands in Placer County, California. California black oak 
seedlings differed among stands (P < 0.001), and seedling densities (number/ha) at one stand (mean = 
58,733) were 4.0—5.3 times more abundant than the other stands (mean = 11,133—14,400). Seedling 
density increased with increasing average diameters (cm) of surrounding California black oaks (P < 
0.008). Larger diameter oaks are older and produce more acorns that can germinate into seedlings than 
smaller diameter oaks. For the stand with the greatest seedling density, seedling density increased with 
increasing basal area (m7/ha) of California black oak (P = 0.028). Variation in site conditions and the 
effects of California black oak diameters and basal area should be acknowledged when evaluating seedling 
densities and seedling-based recruitment for California black oak and prior to initiating management to 


promote seedling recruitment. 


Key Words: 


California black oak (Quercus kelloggii) is wide- 
ly distributed throughout California’s montane en- 
vironments, and its distribution extends into west- 
central Oregon and northern Baja California (Grif- 
fin and Critchfield 1972; McDonald 1990). Occur- 
ring in mostly pure stands or mixed with conifers, 
California black oak is shade intolerant and a vig- 
orous sprouter (McDonald 1969, 1990). Harvesting 
for firewood and saw logs is the land use most af- 
fecting this species, but California black oak re- 
mains mostly underutilized commercially despite 
its wide distribution and large timber volume (Bol- 
singer 1988; McDonald and Huber 1995). Califor- 
nia black oak appears to be declining in some plac- 
es as dying trees are not replaced because conifers 
are shading out oak seedlings and saplings in 
mixed-conifer stands, but not other places where 
oak sapling to tree ratios were adequate to support 
regeneration (Muick and Bartolome 1987; Mc- 
Donald and Tappeiner 1996). 

California black oak regenerates through stump 
sprouting or acorn germination. Sprouting is 
thought to be the primary regeneration method as 
stump sprouts grow fast to capture growing space, 
while acorn germination and growth into seedlings 
is thought to be an infrequent regeneration method 


' Present address: East Bay Municipal Utility District, 
1 Winemasters Way, Lodi, CA 95240. 


Quercus kelloggii, recruitment, seedlings, California. 


(McDonald 1969, 1990). Because of its sprouting 
ability, California black oak is thought to primarily 
occur in even-aged stands as fire and other distur- 
bances are the primary way stands are replaced 
(McDonald 1969, 1990; McDonald and Tappeiner 
1996), and sprouting maintains oak densities in ex- 
isting stands if enough sprouts grow into mature 
trees. 

Seedling occurrence is sporadic and has been as- 
sociated positively with oak canopy cover and re- 
duced solar radiation in woodlands (Standiford et 
al. 1997). Relationships between vegetation attri- 
butes and seedlings, however, have not been elu- 
cidated in forest habitats where timber management 
and fire suppression are dominant land uses. Ger- 
minated acorns change oak densities in existing 
stands as new trees are established, and acorns 
moved by animals away from parent trees can 
change oak distributions (Fuchs et al. 1997). 

Understanding seedling density and occurrence 
patterns would help land managers design and im- 
plement management actions intended to maintain 
or enhance California black oak where sexual re- 
production is favored. We observed four forest 
stands in the central Sierra Nevada with uneven- 
aged California black oak trees (Garrison et al. 
2002) with variable amounts of seedlings. Given 
the lack of information on seedling densities from 
forest habitats, we undertook an investigation to 


116 


document what stand attributes influenced Califor- 
nia black oak seedling density in these four stands. 
Our objectives were to: (1) quantify seedling den- 
sities in stands dominated by mature, but various 
aged California black oaks; and (2) assess how 
seedling density is affected by vegetation attributes 
in these stands. The seedling investigation is part 
of a more comprehensive study of the dynamics of 
California black oak and wildlife population and 
community responses to habitat attributes in these 
same stands. 


STUDY AREA 


We conducted this study on four 21.2-ha study 
stands in Placer County, California. Elevations 
ranged from 1220-1320 m, and the stands were 
located on plateaus and upper portions of steep riv- 
er canyons which characterized the study area. 
Study stands were located in larger size, homoge- 
neous forest stands with a tree layer dominated by 
large diameter (>50 cm diameter breast height 
[dbh]) California black oak and ponderosa pine (Pi- 
nus ponderosa). Other less dominant tree species 
included interior live oak (Q. wislizeni), Douglas- 
fir (Pseudotsuga menziesii), white fir (Abies con- 
color), sugar pine (P. lambertiana), and incense ce- 
dar (Calocedrus decurrens). Seedling and sapling 
California black oak and ponderosa pine dominated 
the subcanopy at one stand, while the other stands 
had little subcanopy. The shrub layer was generally 
sparse, and deerbrush (Ceanothus integerrimus) 
and manzanita (Arctostaphylos spp.) were the most 
common shrubs. The herbaceous layer was domi- 
nated by a sparse to dense cover of mountain mis- 
ery (Chamaebatia foliolosa) and seedlings of Cal- 
ifornia black oak. California black oak was rela- 
tively widespread throughout the study area and oc- 
curred in stands with trees of all ages with varying 
amounts of conifers. The study stands were repre- 
sentative of those with mature, large diameter Cal- 
ifornia black oak in the central Sierra Nevada, al- 
though these stands are rare in this area (Garrison 
et al. 1998). 


METHODS 


Seedling densities and associated vegetative 
characteristics were sampled from 30 0.04-ha plots 
in each stand (known as Stands 1—4). Each stand 
had two adjacent 10.6-ha subplots, and 15 0.04-ha 
circular plots were randomly selected and measured 
in each subplot at intersection points of a 25-m by 
25-m grid. To ensure that the entire subplot was 
sampled, one circular plot was randomly located on 
each of 11 transects in the grid and sampled in 
July—August 1994. Four additional circular plots 
were selected from the same transects and sampled 
in July-August 1995 for a total of 15 circular plots 
per subplot and 30 circular plots per stand. Seedling 
and understory vegetation data were measured at 
all circular plots in 1995. Data were taken on di- 


MADRONO 


[Vol. 49 


ameter (cm) and heights (m) of trees, cover (%) of 
the overstory canopy and understory, and stem den- 
sities (number/ha) and basal areas (m7/ha) of all live 
stems 212.7 cm dbh and =2 m tall. Seedlings (oaks 
<2.5 cm basal diameter) were counted from five 1- 
m? circular plots located at the 0.04-ha circular plot 
center and 5 m away from the plot center along 
axes pointing North (0°), East (90°), South (180°), 
and West (270°). Data were also collected on sap- 
lings (oaks 2.5—15 cm basal diameter and <2 m 
tall), but saplings occurred on 8% (10 of 120) of 
the vegetation plots so they were too rare to be 
analyzed. 

Because most data did not meet assumptions and 
distribution requirements for parametric statistics, 
the data were transformed (Zar 1996). Log trans- 
formations (log,) + 1.0) (Zar 1996) were used for 
averages for seedling counts, basal area, stem den- 
sity, heights and stem diameters for all trees, Cali- 
fornia black oak, and conifers. Cover for understo- 
ry and overstory canopy were given arcsine trans- 
formations [arcsine radian degrees (square root 
(proportion canopy cover + 1))] (Zar 1996). 

Analysis of variance with Bonferroni pairwise 
comparisons were used to determine differences 
among stands for the vegetation attributes. Pearson 
correlation coefficients for the transformed data 
was used to determine preliminary relationships be- 
tween vegetation attributes and seedling densities 
with data from all stands combined and separately 
for Stand 1. While combining data from all stands 
for the correlations is considered pseudoreplication 
(Hurlbert 1984), the correlations were for explor- 
atory purposes to select a smaller subset of habitat 
variables for more detailed analyses and no statis- 
tical inferences were made using these results. 

Vegetation attributes with statistically significant 
(P < 0.05) correlations were used in a backward 
stepwise general linear model to determine stand 
differences in seedling densities and which vege- 
tation attributes had the greatest effect on seedling 
density. A multiple linear regression analysis was 
conducted for Stand 1 because it had the greatest 
seedling densities and frequency of plots with seed- 
lings so it provided an opportunity to assess how 
habitat attributes influence seedling densities at the 
stand level. Understory cover measurements were 
not included in the general linear model or regres- 
sion because two of the five understory attributes 
were highly correlated (P < 0.02) with seedling 
density; this correlation indicated data interdepen- 
dence and redundancy that biased relationships be- 
tween the tree layer and seedlings. Statistical anal- 
yses were conducted using SYSTAT (SPSS Incor- 
porated 1999). Summary statistics and scatterplots 
for each stand were used to assist interpretation of 
the general linear model results. Throughout this 
paper, the term “‘oak”’ is synonymous with Califor- 
nia black oak as almost 100% of the tree oaks and 
100% of the oak seedlings and saplings measured 
were California black oak. 


2002] 


RESULTS 
Stand Attributes 


The four stands consisted of relatively large di- 
ameter California black oaks and smaller diameter 
conifers with closed canopies, large amounts of 
basal area and moderate stem densities (Table 1). 
Sixteen of the 21 (76%) vegetation attributes mea- 
sured differed (P < 0.024) among the four stands. 
Stand 1 had greater oak seedling densities (#/ha), 
greater conifer tree densities (#/ha), greater oak un- 
derstory cover (%), lesser oak tree densities (#/ha), 
smaller diameters (cm) of conifer trees, and shorter 
conifer trees (m) than the other stands (P < 0.001). 
Stem densities, understory and overstory cover, tree 
diameters, and tree heights were the attributes that 
mostly differed among the stands (P < 0.024) (Ta- 
ble 1). Basal area (m7/ha) of all trees and oak were 
similar (P > 0.197) among stands. At all stands, 
63-90% of the sample plots had at least one oak 
seedling indicating that some seedlings occurred 
over much of the area within and among stands 
(Fig. 1). Stands were at similar elevations (1220— 
1320 m); slopes varied from 5—30%, and aspects 
included E, SE, S, and W (Table 1). 


Habitat Attributes Affecting Seedling Densities 


Of the 20 vegetation attributes analyzed for their 
possible effects on seedling densities at the four 
stands, seven (35%) attributes were correlated (n = 
95-120; P < 0.038) with seedling density. Five of 
these seven attributes were from the tree layer (co- 
nifer tree density [#/ha], oak basal area [m?/ha], av- 
erage diameter [cm] of oaks and conifers, and av- 
erage height [m] of oaks) and were used in the gen- 
eral linear model. The other two attributes (% total 
understory cover and % oak understory cover) ere 
not included in the general linear model because 
they are highly correlated (n = 120, P < 0.02) and 
redundant variables for seedling density. At Stand 
1, four (20%) of the 20 vegetation attributes (% oak 
and conifer overstory cover, oak basal area [m2?/ha], 
and % oak understory cover) were correlated (n = 
30, P < 0.049) with seedling density. 

Average diameter (cm) of California black oak 
and stand were only attributes retained in the gen- 
eral linear model (Table 2). Seedling densities dif- 
fered (P < 0.008) among stands and increased (P 
< 0.001) as average oak tree diameter increased 
(Table 2, Fig. 2). This relationship was due some- 
what to the greater number of seedlings and larger 
diameter trees at Stand | (Table 1, Figs. 1 and 2). 
With Stand 1, seedling density increased with in- 
creasing oak basal area (P < 0.028). 


DISCUSSION 


California black oak seedlings were prevalent at 
one of four stands in the central Sierra Nevada, and 
the stands differed statistically in many vegetation 
attributes including densities, diameters, and 


GARRISON ET AL.: CALIFORNIA BLACK OAK SEEDLINGS 117 


heights of trees. Oak seedling densities differed 
among stands and increased with increasing aver- 
age diameters of the surrounding oak trees. More 
seedlings under larger diameter trees are expected 
because larger diameter California black oaks are 
older (Garrison et al. 2002) and acorn production 
increases with increasing age and diameter of Cal- 
ifornia black oak (McDonald 1969; Garrison et al. 
in press). Older trees also have longer periods of 
time to deposit acorns on the ground that germinate 
into seedlings so greater numbers of seedlings ac- 
cumulate under larger trees. 

All stands were dominated by California black 
oak and had equivalent total overstory canopies, 
but slopes, aspects, and oak overstory canopy cover 
varied somewhat among stands. Presence of Cali- 
fornia black oak seedlings is negatively associated 
with solar radiation and positively associated with 
canopy cover of California black oak (Standiford 
et al. 1997), and probabilities of seedling occur- 
rence at the four study stands were between 80— 
90% using the graphs from Standiford et al. (1997). 
Stand 1 had the greatest number of seedlings and 
the least amount of oak overstory cover (36%) as 
well as the least amount of solar radiation due to a 
relatively flat slope. Furthermore, seedling densities 
increased with increasing oak basal area in Stand 
1. Although seedling densities increased with in- 
creasing oak diameter in plots from all four stands, 
oak basal area was also correlated (r = 0.194, n = 
120, P < 0.034) indicating that both vegetation at- 
tributes have affect seedlings. 

Seedling presence and abundance varies across 
small and large geographic areas as seedlings oc- 
curred on 63—90% of our survey plots in the central 
Sierra Nevada yet densities varied four to fivefold 
among stands. Other studies reported California 
black oak seedlings present on 83% of survey plots 
in the southern Sierra Nevada (Standiford et al. 
1997) and 62% of sample plots in the range of the 
California black oak habitat in California (Bolsin- 
ger 1988). 

Seedlings are germinated acorns, so acorn pro- 
duction by parent trees ultimately affects seedling 
densities. California black oak acorn production has 
considerable spatial and temporal variation (Koenig 
et al. 1994) so acorns are limited at certain sites 
and particular times. In our stands, plots with larger 
oaks had lower stem densities with varying 
amounts of canopy cover so germinating acorns 
and seedlings received varying amounts of sunlight. 
Acorns deposited in areas with lesser canopy cover 
germinate and grow at greater levels that acorns in 
areas with greater canopy cover (Savage 1994; 
Standiford et al. 1997), and this pattern occurred at 
Stand 1 where seedling densities were greatest at 
moderate (15-55%) canopy coverages. Seedling re- 
generation of California black oak is clumpy as 
seedlings concentrate around drip lines of parent 
trees (Savage 1994; McDonald and Tappeiner 
1996). Furthermore, acorns are distributed and 


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Stand 3 


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ho 
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012345 6 7 8 9 10111213 1415161718 
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Fic. 1. 


GARRISON ET AL.: CALIFORNIA BLACK OAK SEEDLINGS 119 


Stand 2 


29 
20 
19 es 


10 


Number of plots 


i 12345 6 7 8 9 1011 12 131415 16 1718 
Seedlings per sq. m 


Stand 4 


Number of plots 


Oo 1 2 3 4567 8 9 101112131415 1617 18 
Seedlings per sq.m 


Frequencies of plots with varying densities of California black oak (Quercus kelloggii) seedlings (number/m7?) 


measured in 1995 at four 21.1-ha study stands in Placer County, California. The number of plots without seedlings in 
the 0—1 seedlings/m? density class were Stand 1 = 3, Stand 2 = 8, Stand 3 = 4, and Stand 4 = 11. 


cached by western gray squirrels (Sciurus griseus) 
(McDonald 1969) and Steller’s jay (Cyanocitta stel- 
leri) (personal observation, Fuchs et al. 1997). 
Many cached acorns are not consumed by wildlife 
and then germinate into seedlings. 

In undisturbed settings, regeneration of Califor- 
nia black oak from germinated acorns occurs as a 
steady accumulation of seedlings rather than large 
pulses (McDonald and Tappeiner 1996), although 
seedlings must be aged to confirm this. Because 


they grow slowly under the canopy of overstory 
trees, oak seedlings generally have similar heights 
although ages can be different (McDonald and Tap- 
peiner 1996). Seedling accumulations were obvious 
at Stand 1, but seedling ages were not determined 
to confirm whether they had accumulated over 
many years or resulted from a pulse of seedlings 
coincident with a bumper acorn crop. One good 
acorn crop, however, coincident with proper site 
conditions may result in a single event of great 


120 


MADRONO 


[Vol. 49 


TABLE 2. RESULTS OF BACKWARD STEPWISE GENERAL LINEAR MODEL ANALYSIS ON THE DENSITY OF CALIFORNIA BLACK 
OAK SEEDLINGS (NUMBER/m?2) AND VEGETATION ATTRIBUTES AT FOUR 21.1-HA STANDS IN PLACER COUNTY, CALIFORNIA. 
2 Attributes log, transformed. © Attributes dropped from model if P-value > 0.05. Oak diameter was the covariate. 


ance df F 


Std. 
coeffi- 
Attributes Coefficient SE cient 
Retained in model? 
Stand = — — 
Oak dbh> 0.46 0.17 O25 
Partial 
Correla- 
Dropped from model? tions 
Oak basal area? 0.09 = — 
Conifer dbh> —0.07 — — 
Conifer trees> —0.02 — — 
Conifer height’ —0.01 —- — 
Analysis of Covariance SS 
Stand 3.37 
Oak diameter® 0.70 
Error 7.50 


acorn germination and increase seedling densities. 
Acorn germination, however, is considered less sig- 
nificant as a recruitment source than sprouting by 
McDonald (1969, 1990) and McDonald and Tap- 
peiner (1996), yet Savage (1994) expected seed- 
lings to be a major source of recruitment of Cali- 
fornia black oak in the San Jacinto Mountains. 
Seedlings, rather than sprouts, appear to be the ma- 
jor recruitment source in the four study stands be- 
cause approximately 70% of the mature oak trees 
sampled were single stems (Garrison et al. 2002). 

Seedling densities increased with increasing oak 


ES 


1.0 


0.5 


CA black oak seedlings (#/sq. m) (log10) 


0.0 
Ve ere odd ee Ake) U8) 20 21 


CA black oak diameter (cm) (log10) 


Fic. 2. Scatterplot of California black oak seedlings/m? 
and California black oak diameter (cm) from four 21.1-ha 
study stands measured in 1994 and 1995 in Placer County, 
California. 


Toler- 
P-value R2 

0.59 3 11.48 0.000 0.40 

0.89 1 Tee) 0.008 

0.63 1 0.67 0.415 

0.74 1 0.36 0.549 

0.78 1 0.04 0.848 

0.85 1 0.00 0.960 
MS df F P-value R?2 
1.13 3 15.46 0.000 0.41 
0.70 1 9.55 0.003 
0.07 103 


diameter (P < 0.001) at all stands and increasing 
oak basal area at Stand | so diverse stand condi- 
tions with lesser and greater tree sizes and ages and 
basal areas should result in variable seedling den- 
sities. Larger diameter oaks have more wildlife 
habitat attributes such as dead branches, mistletoe 
(Phoradendron villosum), and acorns (Garrison et 
al. in press) so retaining large diameter trees is rec- 
ommended where land management activities 
might affect these trees. Acorn production varies 
across California black oak diameters from 35—115 
cm (Garrison et al. in press) but larger trees pro- 
duce more acorns so retaining larger trees will 
maintain acorn crops and provide additional wild- 
life habitat benefits. 

Stand | received a prescribed fire on 21 October 
1981, while the other stands had not burned within 
the last two decades. This prescribed fire and the 
larger diameter oaks, flater slope, and moderate oak 
overstory canopy cover were the most significant 
factors distinguishing Stand 1 from the other three 
stands and remains the most plausible reason why 
Stand | had more seedlings than the other stands. 
Prescribed fire appears to be a viable management 
tool in stands dominated by California black oak to 
promote seedlings and eliminate leaf litter (Ander- 
son 1993) as well as to reduce threats from more 
severe, stand-replacing fires. Prescribed fires con- 
ducted in late fall or early spring with low to mod- 
erate fuel loads caused relatively low levels of mor- 
tality to California black oak seedlings and sprouts 
(Kauffman and Martin 1990), so prescribed fires 
can be timed to promote seedling establishment 
from acorns, maintain existing seedling and sapling 
numbers, and damage or kill competing vegetation. 
Fall burns conducted prior to the drop of acorns 
and deciduous leaves might be the best time to burn 
based on our limited data and lack of assessment 


2002] 


of the effects of prescribed fire on seedlings in our 
study area. 

California black oak grows throughout its range 
in many different environmental conditions, and 
stands occur in even and uneven-aged conditions 
with varying amounts of conifers and other hard- 
woods (McDonald 1969, 1990; Garrison et al. 
2002) so management objectives and actions will 
differ accordingly. Retaining acorn-producing trees 
while opening the canopy to moderate levels 
through thinning could produce conditions suitable 
for acorn germination and seedling growth. Thin- 
ning oak stands also improves acorn production 
(Healy 1997; Standiford et al. 2000). Finally, this 
study was done on a limited number of sites (four) 
in the central Sierra Nevada with mature California 
black oaks so these results may have limited appli- 
cation to stands with different vegetative attributes. 


ACKNOWLEDGMENTS 


Ray Brumitt, Karen Durand, Catherine Fowler, Linda 
Louie, Jeanette Mar, Christopher Otahal, David Pratt, and 
Patty Sterling deserve special appreciation for their hard 
work sampling vegetation. Bob Heald, Doug McCreary, 
Phil McDonald, and Richard Standiford provided advice, 
information, and insights into the ecology and manage- 
ment of California black oak. Doug McCreary, Phil 
McDonald, Kristina Schierenbeck, and an anonymous re- 
viewer commented on earlier versions of this paper and 
gave helpful suggestions. This study was funded by the 
Deer Herd Plan Implementation Program of the California 
Department of Fish and Game and U.S. Forest Service. 
We thank Ken Mayer, Terry Mansfield, Eric Loft, Sonke 
Mastrup, Russ Mohr, Joelle Buffa, Susan Sharpley-Evans, 
Linda Tatum, Sheryl] Ducummon, Richard Johnson, Ray- 
mond LaBoa, Lisa Kreuger, Jeff Finn, and Ron Bertram 
for their interest, support, and assistance with this study. 


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Fucus, M. A., P. G. KRANNITZ, A. S. HARESTAD, AND FE L. 
BUNNELL. 1997. Seeds that fly on feathered wings: 
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GARRISON, B. A., R. L. WacuHs, T. A. GILES, AND M. L. 
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land Fisheries Division, Administrative Report 
1998-1. 

, C. D. OTAHAL, AND M. L. TricGs. 2002. Age 

structure and growth of California black oak (Quer- 


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cus kelloggii) in the central Sierra Nevada. Pp. 665— 

679 in Proceedings of the Fifth Symposium on Oak 

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scape. USDA Forest Service General Technical Re- 

port PSW-GTR-184. 

, R. L. WaAcus, T. A. GILES, AND M. L. Triaccs. In 
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KAUFFMAN, J. B. AND R. E. MARTIN. 1990. Sprouting shrub 
response to different seasons and fuel consumption 
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KoeEniIc, W. D., R. L. MumMmMeE, W. J. CARMEN, AND M. T. 
STANBACK. 1994. Acorn production by oaks in central 
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McDOoNna_Lp, P. M. 1969. Silvical characteristics of Cali- 
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. 1990. Quercus kelloggii Newb. California black 

oak. Pp. 661-671 in Silvics of North America, Vol. 

2, hardwoods. USDA Forest Service Agricultural 

Handbook 654. 

AND D. W. Huser. 1995. California’s hardwood 

resource: managing for wildlife, water, pleasing scen- 

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AND J. C. TAPPEINER. 1996. Silviculture-ecology 
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PHILLIPS. 1997. Factors influencing the probability of 
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, R. PHILLIPS, AND N. K. MCDOUGALD. 2000. The 
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Maprono, Vol. 49, No. 2, pp. 122-129, 2002 


CANOPY MACROLICHENS FROM FOUR FOREST STANDS IN THE 
SOUTHERN SIERRA MIXED CONIFER FORESTS OF 
SEQUOIA/KINGS CANYON NATIONAL PARK 


Davip C. SHAW 
Wind River Canopy Crane Research Facility, University of Washington, 
1262 Hemlock Road, Carson, WA 98610 
dshaw @u.washington.edu 


STEVEN A. ACKER 
Olympic National Park, 600 East Park Avenue, Port Angeles, WA 98362 


ABSTRACT 


Canopy macrolichens were sampled using the “‘litter pickup” technique in four forest stands in the 
mixed conifer forests of Sequoia/Kings Canyon National Park. The purpose was to provide a basis for 
assessing lichen abundance trends in permanent forest plots, and to compare differences in lichen com- 
munities between four forest types typical of the southern Sierra Nevada. Each stand was characterized 
by a different conifer: sugar pine (Pinus lambertiana Dougl.), white fir (Abies concolor Gord. & Glend.), 
giant Sequoia (Sequoiadendron giganteum (Lindl.) Buchh.) and Jeffrey pine (Pinus jeffreyi Grev. & Balf.). 
The standing crop of lichen litterfall was estimated at 33.6 kg/ha, 14.8 kg/ha, 6.9 kg/ha, and 7.6 kg/ha 
respectively. Seven macrolichens were present in the litterfall, in decreasing order of overall abundance: 
Letharia vulpina (L.) Hue, Hypogymnia imshaugii Krog, L. columbiana (Nutt.) J. W. Thomson, Bryoria 
fremontii (Tuck.) Brodo & D. Hawksw. and Melanelia exasperatula (Nyl.) Essl., M. subolivacea (Ny1.) 
Essl., and Lobaria (Schreber) Hoffm. sp. A single factor ANOVA indicated that L. vulpina was equally 
distributed throughout the four stands, while H. imshaugii and L. columbiana were not. H. imshaugii was 
the most abundant lichen in the White Fir stand, although L. vulpina closely approximated it there. L. 
vulpina was most abundant in the Sugar Pine, Giant Sequoia and Jeffrey Pine stands, and all other lichens 
were much less abundant. A complex of factors explains the differences in lichen abundance; stand 
density, stand structure, and tree species composition appear most important, although site environmental 
conditions cannot be ruled out due to the lack of replication and small sample size in this study. The 
White Fir and Sugar Pine stands had 2—3 times the tree density as the Giant Sequoia and Jeffrey Pine 
stands. Giant sequoia and incense cedar (Calocedrus decurrens (Torr.) Florin) shed bark and therefore do 
not have abundant epiphytes on branches and tree boles. White fir appears to have a generally positive 
effect on lichen abundance, except in extremely dense stands. The abundance of H. imshaugii and L. 
columbiana were highly correlated with abundance of sugar pine. Although species diversity is low, 


standing crop of lichen litterfall is high, and may exceed many other forests in North America. 


Key Words: 


Macrolichens of forest canopies can be used to 
make inferences about a variety of ecosystem char- 
acteristics, including air quality, stand structure and 
history, stand age, and overall forest health (Segal 
and Nash 1983; Wetmore 1986; Boucher and Stone 
1992; Bates and Farmer 1992; McCune 1993; 
Rhoades 1995). The distribution of these arboreal, 
non-crustose lichens across the landscape reflects 
the dynamic mosaic of environmental conditions 
(Hale 1974). Within the mixed-conifer forests of 
the southern Sierra Nevada Mountains in Califor- 
nia, canopy macrolichens have received limited 
study. 

The National Park Service and other government 
agencies are interested in determining whether the 
lichens are increasing or decreasing in abundance, 
because lichens may have value as indicators of 
environmental problems (McCune 2000). Smith 
(1980) did a taxonomic survey of the macrolichens 
in Sequoia/Kings Canyon National Park and found 
40 species in 13 mostly forested study sites. An air 


Sierra Nevada, lichens, biomass, litterfall, canopy. 


pollution impact survey of all the lichens of Se- 
quoia/Kings Canyon National Park has identified 
204 species (Wetmore 1986). Wetmore concluded 
that considering the dry climate, the lichen flora 
was diverse and healthy. 

The purpose of this study was twofold: 1. To 
document the relative abundance of canopy ma- 
crolichens in four forest stands that are part of a 
permanent forest plot system in Sequoia/Kings 
Canyon National Park (Harmon et al. 1987; Riegel 
et al. 1988). These data provide a baseline for fu- 
ture sampling to determine temporal trends in can- 
opy macrolichen abundance. 2. To compare the rel- 
ative abundance of canopy macrolichens in four 
forest stands dominated by different species of co- 
nifers and representing different environmental 
conditions in the lower montane, mixed conifer for- 
ests of the southern Sierra Nevada Mountains. 

The lower montane, mixed-conifer forests of the 
southern Sierra Nevada Mountains of California 
(1,600 m to 2,300 m) are characterized by giant 


2002] 


sequoia (Sequoiadendron giganteum (Lindl.) 
Buchh.) (Cupressaceae), white fir (Abies concolor 
(Gord. & Glend.) Lindl. ex Hildebr.) (Pinaceae), 
California red fir (Abies magnifica A. Murr.) (Pin- 
aceae), sugar pine (Pinus lambertiana Dougl.) (Pin- 
aceae), Jeffrey pine (P. jeffreyi Grev. & Balf.) (Pin- 
aceae), and incense cedar (Calocedrus decurrens 
(Torr.) Florin) (Cupressaceae). Along a moisture 
gradient, giant sequoia occurs in mesic locations 
that do not dry out in the summer, white fir-mixed 
conifer (sugar pine and incense cedar) occurs in 
generally drier habitats, and Jeffrey pine occurs in 
the most xeric sites (Rundel et al. 1977; Vankat 
1982). Fire and fire suppression play an extremely 
important role in stand composition and structure. 
In general, fire suppression results in an increase in 
the abundance of white fir (Rundel et al. 1977). 

Appropriate sampling for canopy macrolichen 
studies can be challenging, particularly for studies 
of trends in abundance over time. Canopy access 
using tree climbing is the most direct means of 
sampling canopy macrolichens, but sampling tree 
crowns to determine stand level abundance (i.e. 
biomass) requires very large amounts of time in tall 
forests (Clement and Shaw 1999). As an alterna- 
tive, McCune (1994) has developed a method to 
quantify the relative abundance of lichens in a for- 
est stand by sampling litterfall. This “‘litter pick- 
up’’ technique allows one to estimate the mass of 
canopy macrolichens at the stand level, which can 
then be used to compare relative abundance to other 
stand types and to determine stand-level trends in 
abundance over time. 


METHODS 
Study Site/Reference Stands 


The study site is located in the northwest portion 
of Sequoia National Park (Latitude 36°N and Lon- 
gitude 118°W) (Fig. 1). We chose four of the six 
reference stands described by Riegel et al. (1988), 
each dominated by a different species of conifer; 
Jeffrey pine, white fir, sugar pine (mixed conifer), 
and giant Sequoia (in a riparian setting). The stands 
are between 2,012 and 2,219 m in elevation and 
representative of three vegetation types: Sierran 
mixed-conifer (sugar pine and white fir), giant Se- 
quoia-mixed conifer (riparian), and Jeffrey pine 
(Riegel et al. 1988). The reference stands were es- 
tablished for long-term monitoring of vegetation 
and are cooperatively managed by the Sequoia/ 
Kings Canyon National Park, Oregon State Uni- 
versity, and the US Forest Service, Pacific North- 
west Research Station (Acker et al. 1998). The ref- 
erence stands were established in 1984 and re-mea- 
sured in June of 1994. All trees >5 cm have been 
tagged and mapped, and each tree has the diameter 
at breast height measured. Information is collected 
on crown ratio, crown vigor, tree mortality and 
damage. All data presented on stand structure 
comes from the 1994 measurement. 


SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON 128 


Sequoia & Kings 


Canyon National GaN 


* FRESNO 


a 
A) 


A 


WOLVERTON 


Mixed 
Conifer 
SUWANEE 
CREEK 


HALSTEAD 


GENERAL SHERMAN 
TREE 


124 
> 
z 
+ 


Riparian Giant 
a) 
yo Giant ) 


Sequoia 
CRESCENT 
MOW 


FOREST 


4 MORO ROCK 


Fic. 1. Location of study site in Sequoia National Park, 
California (reproduced from Riegel et al. 1998). Four of 
these six reference stands were sampled, including White 
Fir, Jeffrey Pine, Riparian Giant Sequoia (Giant Sequoia) 
and Mixed Conifer (Sugar Pine). 


The study site has a mediterranean climate, with 
cool, moist winters and hot dry summers. Precipi- 
tation averages 1172 mm/year (1932—1983 mean at 
Giant Forest/Lodgepole, Sequoia National Park) 
and falls mostly as snow between November and 
April. The hot dry summers have a strong influence 
on arboreal lichen communities, which are charac- 
terized by low species numbers and dominance by 
a few drought-tolerant species. 

The Jeffrey Pine reference stand (1.0 ha) is on a 
moderately steep SE facing slope, (Table 1), with a 
glaciated granodiorite rock substrate. Exposed rock 
is common at the site. The canopy is open, domi- 
nated by Jeffrey pine (124 trees/ha), with California 
black oak (Quercus kellogii Newb.) (Fagaceae) (60 
trees/ha) (Table 2). Dense clumps of shrubs, espe- 
cially green manzanita (Arctostaphylos patula 
Greene) (Ericaceae), are present. White fir (18 
trees/ha), sugar pine (2 trees/ha) and incense cedar 
(5 trees/ha) occur in the lower plot where the slope 
flattens. This is a xeric, low productivity site. Jef- 
frey pine is adapted to the coarse textured soil 
found in the fissures of the glaciated granite (Riegel 
et al. 1988). 

The White Fir reference stand (0.9 ha) is located 


124 MADRONO [Vol. 49 
TABLE |. SITE CHARACTERISTICS OF THE FOUR REFERENCE STANDS IN SEQUOIA NATIONAL PARK. 

Reference Stand size Elevation Topographic Average 

stand (ha) (m) Aspect position slope (%) 

Sugar Pine 1.1 2091 southeast midslope-bench 11 
White Fir 0.9 2012 southwest bench 20 
Giant Sequoia 2.0 DANG) southwest/southeast lower slope 10 
Jeffrey Pine 1.0 2109 southeast upper slope 23 


on a flat area above the east-side of Suwanee Creek 
(Table 1). There are scattered outcrops of bedrock 
in the stand. The stand has a dense canopy of white 
fir (420 trees/ha) and California red fir (33 trees/ 
ha) near the stream which grades into a mixed-co- 
nifer forest with scattered sugar pine (56 trees/ha) 
and incense cedar (177 trees/ha) on the east side of 
the reference stand (Table 2). White fir is most 
abundant in all size- and canopy-classes,\while in- 
cense cedar, sugar pine and California red fir are 
more abundant in the intermediate and suppressed 
canopy classes and smaller diameter-classes (Riegel 
et al. 1988). 

The Sugar Pine reference stand (1.1 ha) is locat- 
ed to the west of Suwanee Creek approximately 
200 m from the White Fir reference stand on a mid- 
slope bench (Table 1). The forest is a mosaic of 
large old sugar pine (110 trees/ha) and white fir 
(473 trees/ha) trees forming a relatively open can- 
opy in the dominant (sugar pine 20 trees/ha, white 
fir 19 trees/ha) and codominant (sugar pine 11 
trees/ha, white fir 42 trees/ha) canopy classes (Table 
2, Riegel et al. 1988). There are clumps of sup- 
pressed white fir and incense cedar (78 trees/ha) 
interspersed throughout the stand where white fir 
dominates the smaller size and canopy classes. Cal- 


ifornia black oak (16 trees/ha) and California red 
fir (7 trees/ha) are present in low numbers. The 
abundance of white fir in small size classes is 
thought to be a result of fire suppression (Riegel et 
al. 1988). 

The Giant Sequoia reference stand (2.0 ha) is on 
a lower slope, and straddles both sides of Crescent 
Creek (Table 1). There is a narrow corridor of her- 
baceous vegetation along the creek. The stand has 
a typical mixed conifer over-story dominated by gi- 
ant sequoia (24 trees/ha), which tower above the 
surrounding white fir (222 trees/ha) and California 
red fir (64 trees/ha) (Table 2). The true firs have a 
reverse J-shaped size distribution with a predomi- 
nance of small stems, as is typical of shade tolerant 
species (Riegel et al. 1988). 


Macrolichen Sampling 


Canopy macrolichens were sampled on June 20— 
24, 1994 using 2-m radius (12.57 m7) litter pickup 
plots (McCune 1994). Litter refers to material Gin 
this case lichens) fallen from the canopy. At fifteen 
randomly chosen grid points in each reference 
stand, a stake was placed in the center of the plot 
and a 2-m string was used to measure the radius of 


TABLE 2. SPECIES COMPOSITION, NUMBER OF TREES PER HECTARE (TPH), TOTAL NUMBER OF SPECIES, DIAMETER (IN cm) 
CHARACTERISTICS, TREE SPECIES EVENNESS, TREE SPECIES RICHNESS BASED ON NUMBERS (KREBS 1989), TREE SPECIES 
RICHNESS BASED ON AREA (KREBS 1989) OF THE FOUR REFERENCE STANDS. 


Stand 

TPH by species Sugar Pine White Fir Giant Sequoia Jeffrey Pine 
White Fir 472.6 420.5 Dies 18.0 
Red Fir Toll 33.0 63.5 0) 
Incense Cedar THD 177.3 0) 5.0 
Jeffery Pine 0 0) 1.0 124.0 
Sugar Pine 109.7 Se 6.0 2.0 
Ponderosa Pine 0 0) 0) 1.0 
Cal. Black Oak 15.9 0) 0 60.0 
Giant Sequoia 0 0) D335) 0) 
TOTAL TPH 683.2 686.4 3525 210.0 
# Tree Species 5) 4 5 6 
median dbh Or 15.4 12.8 8.6 
quad mean dbh 33.5 B72 81.3 31.8 
max dbh 154.1 148.7 600.0 1333)5 I 
Basal area (m7?/ha) 60 74 164 17, 
Evenness 0.61 0.71 0.63 0.71 
Rich No. 4.9 (0.27) 4.0 (0.00) 4.6 (0.51) 6.0 (0.00) 
Rich Area 5.0 (0.00) 4.0 (0.00) 4.7 (0.47) 5.9 (0.35) 


2002] 


TABLE 3. TOTAL STANDING CROP OF CANOPY MACROLICHENS (kg/ha) ON THE FOREST FLOOR AND FREQUENCY OF OCc- 


CURRENCE IN 2 M RADIUS PLOTS. STANDARD DEVIATION IN PARENTHESES. 


SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON 


Stand 
Sugar Pine White Fir Giant Sequoia Jeffrey Pine 
Total Lichens 
Kg/ha 55:6 (27.7) 14.8 (18.5) 6:95:29) 7.6 (16.5) 
Frequency (%) 100 100 100 100 
Letharia vulpina 
Kg/ha 15.04 (13.3) G5. (f233) 4.8 (1.5) T.33"(iG2) 
Frequency (%) 100 73 93 100 
Letharia columbiana 
Kg/ha 4.4 (5.4) 095153) 0.2 (0.4) 0.15 (0.18) 
Frequency (%) 93 100 73 80 
Hypogymnia imshaugii 
Kg/ha 14.1 (15.4) TS. (120) 8: (2:5) 0.08 (0.18) 
Frequency (%) 93 93 80 33 
Bryoria fremontii 
Kg/ha 0.01 (0.03) 0) 0.08 (0.3) 0.01 (0.030 
Frequency (%) 20 0 20 13 
Melanelia spp. 
Kg/ha 0 0 0.06 (0.1) 0) 
Frequency (%) 0 0) 33 0 


the plot. Flagging was located in four directions to 
denote the boundaries of the litter pickup plot. All 
fresh macrolichens (i.e., had not decayed beyond 
an identifiable state) were collected and placed in 
paper bags. Litter attached to wood was also col- 
lected, as was litter caught in shrubs up to 1 m off 
the ground that was not attached to the shrubs. 

Macrolichens were transported to the lab, 
cleaned, and sorted to species. The lichens were 
then dried at 60°C for 24 hours and weighed. Li- 
chen identifications were made using Hale and Cole 
(1988), and names follow Brodo et al. (2001). Spe- 
cies identifications were verified and unknown sam- 
ples were identified by Bruce McCune, Oregon 
State University. Reference specimens are deposit- 
ed in the University of Washington Herbarium. 


Analysis 


Biomass on each 2-m radius plot was trans- 
formed to g/ha for data analysis. The mean for the 
15 plots in each reference stand was used to rep- 
resent stand level abundance and reported as kg/ha 
with standard deviation. Total lichen biomass and 
biomass of Letharia vulpina (L.) Hue (Parmeli- 
aceae), L. columbiana (Nutt.) J. W. Thomson (Par- 
meliaceae), and Hypogymnia imshaugii Krog (Par- 
meliaceae) were compared between reference 
stands using a single factor Analysis of Variance 
(Zar 1999), (d.f. = 3 between groups, and 56 d.f. 
within groups, a = 0.05). 

Although the study included only four stands, we 
explored various descriptors of forest structure and 
composition as potential predictors of total lichen 


litterfall biomass and biomass of each lichen spe- 
cies. Variation in lichen biomass was compared to 
variation in stand-level tree density and basal area, 
stem density of individual tree species, and tree 
species evenness and richness (from the rarefaction 
method (Krebs 1989)). 


RESULTS 
Species 


Seven species were found in the macrolichen lit- 
terfall of these four reference stands: Letharia col- 
umbiana, L. vulpina, Hypogymnia imshaugii, 
Bryoria fremontii (Tuck.) Brodo & Hawksw. (Par- 
meliaceae), Melanelia exasperatula (Nyl.) Essl. 
(Parmeliaceae), and M. subolivacea (Nyl.) Essl. 
(Parmeliaceae) and a Lobaria (Schreber) Hoffm. 
(Lobariaceae) sp. fragment. The Lobaria fragment 
was unidentifiable to species, and is not discussed 
further. The Letharia species and H. imshaugii were 
present in all four stands, while Bryoria fremontii 
was absent from the White Fir stand. The two Mel- 
anelia species were present only in the Giant Se- 
quoia stand which at six species, had the highest 
macrolichen litterfall species diversity. The other 
stands had four species, including the Lobaria sp. 
fragment at the White fir stand. 


Abundance 


The Sugar Pine stand had the highest standing 
crop of lichen litterfall (33.6 kg/ha) (Table 3). The 
White Fir stand had about % as much (14.8 kg/ha) 
and the Giant Sequoia (6.9 kg/ha) and Jeffrey Pine 


Biomass lichens (kg/ha) 


Number of Sugar Pine per hectare 


Fic. 2. Density of Sugar Pine per hectare on the four 
reference stands versus the standing crop of litterfall l- 
chen biomass per hectare for total lichens, Letharia vul- 
pina (LEVU), L. columbiana (LECO), and H. imshaugii 
(HYIM). Sugar Pine density corresponds to forest stands: 

= Jeffrey Pine, 6 = Giant Sequoia, 56 = White Fir, 110 
= Sugar Pine. 


(7.6 kg/ha) stands had about one fourth that much 
lichen litterfall biomass as the Sugar Pine stand. In 
three of the four reference stands, lichen litterfall 
biomass was dominated by a combination of L. vul- 
pina and H. imshaugii. In the Giant Sequoia, White 
Fir, and Sugar Pine stands, L. vulpina accounted for 
44% to 70% of lichen litterfall biomass and H. im- 
shaugii accounted for 25% to 49%. The Jeffrey 
Pine stand was unusual in that nearly all the lichen 
litterfall biomass (97%) was contributed by a single 
species, L. vulpina. The only other species to ac- 
count for 10% or more of the lichen litterfall bio- 
mass of any stand was L. columbiana, which was 
13% of the biomass for the Sugar Pine stand. 

The ANOVA indicated significant differences 
between reference stands in biomass of total lichens 
(P < 0.01), L. columbiana (P < 0.01), and H. im- 
shaugii (P < 0.01). No significant difference was 
found for L. vulpina (P = 0.16). Biomass of L. 
columbiana and H. imshaugii generally increase 
with increasing density of sugar pine (Fig. 2). Tree 
density, basal area, tree species evenness, tree spe- 
cies richness based on numbers, or tree species 
richness based on area (Table 2, 3) shows little re- 
lationship to the variation in total biomass of li- 
chens. 


DISCUSSION 
Species Richness and Distribution 


The lichen litter pick-up technique documented 
only seven species of lichens in these four forest 
stands. This is low species diversity, even for dry 
habitats. Smith (1980) found 40 species of macroli- 
chens in the Ash Mountain to Grant Grove (High- 
way 198) region of Sequoia-Kings Canyon Nation- 
al Park and also included Cedar Grove. He sampled 


MADRONO 


[Vol. 49 


13 sites using a floristic survey method that includ- 
ed all substrates, not just canopy lichens. The litter 
pick-up technique is not a ‘stand-alone’ method for 
surveys of species diversity, because species cap- 
ture is low. The technique is best used in conjunc- 
tion with other survey techniques that specifically 
search for different species of lichens (McCune and 
Lesica 1992; McCune 1994). However, it is a good 
technique for determining the relative abundance of 
the predominant forest canopy species. This is im- 
portant for monitoring change in lichen communi- 
ties. 

In a study of lichens on conifers and their rela- 
tion to air pollution in the Southern California 
mountains outside of Los Angeles, Sigal and Nash 
(1983) reported 16 species, the same number re- 
ported by Hasse for the same area in 1913 (Gin Sigal 
and Nash 1983). The lichen flora showed a strong 
relationship to air pollution: only eight of the orig- 
inal 16 species reported by Hasse were present in 
the most heavily polluted forests of the San Ber- 
nardino and San Gabriel Mountains. Sigal and Nash 
(1983) also rated the sensitivity of lichen species to 
air pollution, including several species we ob- 
served. They ranked L. vulpina as tolerant, M. su- 
bolivacea as moderately tolerant, and B. fremontii 
as very sensitive. 

Smith (1980) has given species accounts of all 
40 species he observed in Sequoia/Kings Canyon 
National Park, including the six species observed 
in this study. According to Smith, Bryoria fremontii 
is uncommon, and was only collected once from 
the bark and branches of Pinus murrayana (Sierra 
lodgepole pine, P. contorta subsp. murrayana (Bal- 
four) Engelmann (Pinaceae)) in the Stony Creek 
area. We found B. fremontii in the Jeffrey Pine, 
Giant Sequoia, and Sugar Pine reference stands. 
Hypogymnia imshaugii was uncommon and was 
collected on A. concolor in the Crystal Cave Junc- 
tion area. Smith found H. enteromorpha (Ach.) 
Nyl. (Parmeliaceae) to be common and sometimes 
very abundant in all areas above 450 m. This in 
contrast to our finding of AH. imshaugii in all four 
sites, and a lack of collections for H. enteromorpha. 
Hale and Cole (1988) note that in the past, virtually 
all fertile Hypogymnias in California were called 
H.. enteromorpha, but that this name is now limited 
to populations along the coast that are characterized 
by grossly inflated branches, and that this species 
does not occur in the Sierra Nevada. Hale and Cole 
(1988) also indicate that H. imshaugii is very com- 
mon in Sequoia National Park. 

Smith considers Letharia columbiana and L. vul- 
pina to be two of the most common and abundant 
lichens in the park between 1200 m to 2700 m el- 
evation. He found them growing on numerous tree 
species all through the study region. We also found 
these two lichens to be abundant. Interestingly, L. 
vulpina was the more abundant of the two species 
with 3 to 10 times the biomass of L. columbiana in 
the reference stands. Smith found Melanelia subo- 


2002] 


livacea (called Parmelia subolivacea Nyl.) abun- 
dant in all 13 study sites and in some trees the 
upper branches were completely covered by the li- 
chen. It was present on a wide variety of conifers 
and hardwoods. Parmelia exasperata De Not. was 
described as commonly found on Quercus, wide- 
spread in the Ash Mountain area, Potwisha, Buckey 
Flats and Deer Ridge. This may be what we iden- 
tified as M. exasperatula. We found these two spe- 
cies were present only in litterfall of the Giant Se- 
quoia reference stand. 


Abundance 


Letharia vulpina was the dominant lichen in 
three of these forest stands, and was generally 
equally distributed throughout the four forest 
stands. L. columbiana and H. imshaugii were not 
equally distributed and showed strong patterns of 
increase with increasing sugar pine and white fir. 
The extreme xeric conditions of the Jeffery Pine 
stand may have a negative influence on L. colum- 
biana and H. imshaugii. 

McCune (1994) has investigated canopy litter- 
fall relationships in the Pacific Northwest of North 
America. He found that the ratio 1:100 (litter : can- 
opy lichens) was fairly consistent in Douglas-fir 
forests for late summer standing crop of lichen lit- 
ter. Thus about 100 times the amount of lichen 
found on the forest floor in late summer will be in 
the canopy. This relationship has not been tested 
for forests of the Sierra Nevada. However if it is 
valid for the Sierra Nevada, the canopy biomass 
of macrolichens in the four reference stands would 
range from 0.7 Mg/ha in the Giant Sequoia stand 
to 3.4 Mg/ha in the Sugar Pine stand, with inter- 
mediate values for the Jeffrey Pine and White Fir 
stands (0.8 Mg/ha and 1.5 Mg/ha, respectively). 
These numbers are surprisingly large, perhaps in 
part because litter was collected in June rather 
than late summer. Typically a large pulse of lichen 
litter from winter storms will gradually disappear 
over the next 6-12 months depending on the spe- 
cies (McCune and Daley 1994). Another possibil- 
ity is that the mildly toxic Letharia spp are resis- 
tant to herbivory, resulting in greater persistence 
on the forest floor. 

Some of the most productive old-growth Doug- 
las-fir stands in the Pacific Northwest have 1.3 to 
1.9 Mg/ha of macrolichens in the canopy (McCune 
1993; McCune et al. 1997). Boucher and Nash 
(1990) estimated 0.75 Mg/ha macrolichens for can- 
opies of Blue Oak in California (36°N Latitude) 
while Turner and Singer (1976) estimated 1.9 Mg/ 
ha for a Pacific Silver Fir stand in the western Cas- 
cades of Washington. For further information on 
the biomass of epiphytes see Boucher and Stone 
(1992) and Rhoades (1995). The relationship of li- 
chen litter biomass to lichen biomass in the cano- 
pies of Sierra Nevadan forests is a key area for 
future research. 


SHAW AND ACKER: CANOPY MACROLICHENS IN SEQUOIA/KINGS CANYON IDF 


Factors Influencing Lichen Abundance 


Lichen species composition and abundance in 
forest canopies is influenced by a multitude of fac- 
tors. Among other things, this includes tree spe- 
cies, bark texture/chemistry, stand age, ecological 
continuity of the forest (Bates and Farmer 1992), 
tree density, forest structure, disturbance history, 
air pollution, climatic conditions, and forest man- 
agement practices (Hale 1974; McCune 1993; 
Rhoades 1995). Within the southern Sierran 
mixed-conifer forests that we sampled, the most 
obvious influences on lichen species composition 
and abundance include tree species composition, 
stand density, and forest stand structure. It should 
be stressed that the generality of our interpretation 
is limited by the small sample size and no repli- 
cation of stand types. 

Differences in tree density did not directly cor- 
respond to differences in lichen biomass as stands 
with similar tree density differed in lichen biomass 
by a factor of 2 (Sugar Pine and White Fir) and 
stands with similar lichen biomass differed in tree 
density by 50% (Giant Sequoia and Jeffrey Pine). 
Tree species composition may explain some of 
these differences. Though they were similar in den- 
sity, the Sugar Pine and White Fir stands were very 
different in stand structure and species composition. 
The White Fir stand was uniform in stand structure 
creating more evenly shaded tree boles, and had 
Over twice aS many incense cedar (78/ha in the 
Sugar Pine to 177/ha in the White Fir stand). In- 
cense Cedar has exfoliating bark that sheds lichens. 
The Sugar Pine stand was more open with twice 
the number of sugar pine trees (110/ha in sugar 
pine to 56/ha in the fir stand) and a complement of 
dominant and codominant trees in the overstory, 
which provides for more sunlight on tree boles and 
branches. Thus, the Sugar Pine stand may have had 
an optimal combination of tree species composition 
and stand structure to provide for abundant lichen 
biomass. 

Giant Sequoia represented 74% of the basal area 
and 80% of the stand wood volume in the Giant 
Sequoia stand although accounting for only 7% of 
the stems. Giant sequoia also has exfoliating bark, 
hence the lichens are rare on the tree bole, and only 
abundant on dead wood and cones (Steve Sillett 
and Joel Clement, personal communication). This 
might explain why the Giant Sequoia and Jeffrey 
Pine stands were similar in lichen litterfall biomass 
even though the sequoia stand had 50% more trees. 
The Jeffrey Pine stand approached a woodland set- 
ting, with widely scattered trees, among outcrops 
of rock. Letharia vulpina was the dominant lichen 
in this forest, perhaps showing a tolerance for xeric 
conditions and compatibility for Jeffery pine bark 
texture and chemistry. 

The physical settings of the forest plots, such as 
aspect and proximity to streams, may also play a 
role in lichen abundance. The Giant Sequoia stand 


128 MADRONO 


had a stream running through it, the White Fir stand 
and Sugar Pine stands were adjacent a stream, and 
the Jeffrey Pine stand was not influenced by a 
stream. A xeric to mesic environmental gradient 
was not measured in a systematic and replicated 
way in this study, and therefore conclusions re- 
garding the overall effect of tree species composi- 
tion as the major influencing factor associated with 
lichen abundance should be taken as a hypothesis 
needing further study. 


CONCLUSIONS 


We observed low species diversity of canopy h- 
chens in the mixed-conifer forests of the southern 
Sierra Nevada Mountains, yet an unusually high 
stand biomass of lichen litterfall. The Sugar Pine 
stand would be one of the highest biomass esti- 
mates for lichens in North America if the 1:100 
ratio of litterfall to canopy lichen biomass for 
Northwestern forests (McCune 1994) holds true in 
Sierran forests. The early summer sample period 
and possibility of longer persistence on the forest 
floor by Letharia spp. may explain these higher 
numbers. Tree species composition (especially 
abundance of sugar pine), and canopy openness/ 
vertical structure appear to play a role in the abun- 
dance of canopy macrolichens, although the lack of 
replication within stand types and along the envi- 
ronmental moisture gradient preclude a definitive 
analysis. Characteristics of forest stands are con- 
trolled by a complex of factors, but in the future, 
anthropogenic influences such as fire suppression 
and controlled burning, air pollution, and climate 
change may become very important in determining 
lichen abundance. Long-term monitoring of lichens 
is important for understanding their role in the dy- 
namics of ecosystems and how they will respond 
to anthropogenic influences. 


ACKNOWLEDGMENTS 


This research was supported by the Wind River Canopy 
Crane Research Facility, a cooperative scientific endeavor 
of the University of Washington, Gifford Pinchot National 
Forest, and Pacific Northwest Research Station, US Forest 
Service, and the Sequoia/Kings Canyon National Park. 
Susan C. Shaw assisted with field sampling. Bruce 
McCune, Eric B. Peterson, Jeanne M. Ponzetti, and Tom 
Rambo provided editorial comment and review of the 
manuscript, which is especially appreciated. 


LITERATURE CITED 


ACKER, S. A., W. A. McCKEE, M. E. HARMON, AND J. FE 
FRANKLIN. 1998. Long-term research on forest dy- 
namics in the Pacific Northwest: a network of per- 
manent plots. Pp. 93-106 in F Dallmeier and J. A. 
Comiskey (eds.), Forest biodiversity in North, Central 
and South American, and the Caribbean: research and 
monitoring. UNESCO, Paris, France. 

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Bropo, I. M., S. D. SHARNOFF, AND S. SHARNOFF. 2001. 
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CLEMENT, J. P. AND D. C. SHAW. 1999. Crown structure 
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HARMON, M. E., K. CROMACK, JR., AND B. G. SMITH. 1987. 
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2002] 


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ZAR, J. H. 1999. Biostatistical analysis. Prentice-Hall, En- 
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MApRONO, Vol. 49, No. 2, pp. 130-131, 2002 


CALYSTEGIA SILVATICA (CONVOLVULACEAE) IN WESTERN 
NORTH AMERICA 


R. K. BRUMMITT 
The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, U.K. 


At the kind request of the Jepson Manual editor, 
I contributed the account of Calystegia to the recent 
edition of the Jepson Manual (Brummitt in Hick- 
man 1993: 517-521) despite my never having set 
foot in California at the time. My knowledge of the 
taxa in California, where half the genus is endemic, 
was based entirely on extensive study of herbarium 
specimens sent on loan from the major herbaria in 
the late 1960s. I was unaware of any introduced 
species established in the state, though I was fa- 
miliar with several species of the genus which had 
become very well naturalized as weeds in various 
other parts of the world. 

In 1997 I attended a meeting at the California 
Academy of Sciences, and, to my great delight, was 
able to see in the field for the first time many of 
the Calystegia taxa native to the state. But, to my 
surprise, I was able to add one introduced species 
to the Californian list before ever stepping into the 
field, when I reidentified a specimen in the CAS 
collections as C. silvatica (Kit.) Griseb., native of 
southern Europe. The specimen was collected at 
Stinson Beach in Marin Co. in the late 1950s by 
J.T. Howell and then identified and laid away as C. 
sepium (L.) R.Br. My host at CAS, Tom Daniel, 
immediately agreed to take me to Stinson Beach to 
see if the plant was still there, and, sure enough, it 
was there in great abundance on waste ground over 
a distance of perhaps 100 m, swamping other veg- 
etation and climbing to a height of 3 m, very con- 
spicuous even to a passing motorist. It was surpris- 
ing that such a conspicuous plant has been so over- 
looked, and we collected additional material: Marin 
Co., Stinson Beach, roadside in town, alt. 20 m, 
122°39’W, 37°54'N, abundant, rampant over vege- 
tation to 3 m, 31 May 1997, R.K. Brummitt & T.F. 
Daniel 19614 (CAS, K, MO, RSA, UC). 

Alerted to the presence of this species in the area, 
I noticed it again a few days later some 11 km east 
of Stinson Beach on waste ground in the town of 
Mill Valley, and collected it again: Marin Co., Mill 
Valley, near foot of Reed Street, roadside in sub- 
urban area, 122°33'W, 37°54'N, climbing over 
bushes and roadside vegetation to 3 m, 10 June 
1997, R.K. Brummitt 19672 (CAS, K). Again, the 
strongly climbing stems and the large white flowers 
were very conspicuous. 

C. silvatica is readily distinguished from any 
Calystegia in northern or mid California by its very 
vigorous habit, large glossy leaves, and particulaly 
its large flowers (corolla 50-75 mm) with two large 


braceoles at the base which are inflated and over- 
lapping and more or less obscuring the calyx. In 
southern California it is approached in its flowers 
and bracteoles by C. macrostegia (House) Brum- 
mitt subsp. macrostegia and subsp. amplissima 
Brummitt from the Channel Islands, but that spe- 
cies is woody at the base whereas C. silvatica is 
rhizomatous, and the two species are almost cer- 
tainly not closely related despite their superficial 
similarity. 

The pan-temperate complex of C. sepium and re- 
lated species, including C. silvatica, is difficult to 
resolve taxonomically, consisting of a series of geo- 
graphical taxa with minor distinguishing characters. 
My concept of C. silvatica includes plants native 
of eastern N. America, southern Europe and as far 
east as Iran, and China, which occur sympatrically 
with C. sepium except in China and are distin- 
guished from it by their large inflated and overlap- 
ping bracteoles. Those from eastern N. America 
and China are characterised by sometimes having 
twin peduncles in leaf axils and a rather square leaf 
sinus, and I refer them to subsp. fraterniflora 
(Mack. & Bush) Brummitt. In the Mediterranean 
region those from central and southern Italy east- 
wards have a longer range of flower size and big- 
ger, more inflated bracteoles with a rounded to 
emarginate apex, and these are referred to subsp. 
silvatica. Plants from northern Italy and southern 
Switzerland westward to Spain tend to have a 
smaller range of flower size and an obtuse apex to 
the bracteoles. Despite more than thirty years of 
hesitation over whether to give this variation formal 
taxonomic recognition—which necessitated a 
lengthy note on the matter in Flora Europaea 
(Brummitt 1972) instead—I have recently formally 
separated these southwestern European plants as 
subsp. disjuncta Brummitt (see below). 

Both of the Mediterranean subspecies were in- 
troduced into the British Isles about a century ago, 
probably as garden ornamentals, and both are now 
serious weeds there (but, curiously, not so in other 
north European countries). Subsp. silvatica is also 
known as a weed in Australia, where subsp. dis- 
juncta appears not to have been introduced. In 
North America, however, it seems to have been 
only subsp. disjuncta that has become established, 
this being known from specimens collected in 
Washington State from 1927 onwards and also from 
British Columbia and Oregon. Its occurrence in 
California is thus not very surprising. Subsp. fra- 


2002] 


terniflora has been recorded as a rare alien in the 
British Isles, almost certainly introduced from 
North America, but has not become established. 

In the standard text on the plants of the Pacific 
Northwest, Hitchcock (1959) included all these taxa 
in Convolvulus sepium L. The plant figured there 
under this name on p. 88 is very probably C. sil- 
vatica subsp. disjuncta. The plant referred to in the 
text as var. fraterniflorus is probably also this taxon 
and this epithet is misapplied here. The statement 
that it is native of the eastern United States is in- 
correct. It was already described by Hitchcock here 
as a difficult weed. 

All taxa in the C. sepium complex spread vig- 
orously by rhizomes and tend to become aggressive 
weeds, often swamping the vegetation the stems 
climb over. The rhizomes are, however, not quite 
so deep-rooted as those of Convolvulus arvensis L., 
a major weed in California, and they are not quite 
such a persistent pest. Seed dispersal in C. sepium 
and allies is less significant than vegetative spread, 
since there is no obvious dispersal mechanism and 
the seeds tend to merely fall to earth beneath the 
parent plant. Furthermore, all plants in this complex 
are self-sterile and single populations normally do 
not set seed at all. C. silvatica tends to be more 
vigorous in its vegetative spread than the variants 
of C. sepium are, and its potential as a serious weed 
is considerable. It seems that in Britain new popu- 
lations may be established by accidental transport 
of pieces of rhizome (see, for example, notes in 
Brummitt & Chater 2000). 


Calystegia silvatica (Kit.) Griseb., Spic. Fl. Rum. 
2: 74 (1844) subsp. disjuncta Brummitt in La- 
gascalia 18: 339 (1996). 


Additional specimens documenting known range 
in North America. 
BRITISH COLUMBIA. West Vancouver, 16 Ma- 


BRUMMITT: Calystegia silvatica in Western North America 131 


rine Drive, roadside, 6 July 1955, W. Bird 1239 
(BM); Vancouver, Bush Crown & 25 Ave., 5 May, 
1956, W. Bird 2008 (BM): Vancouver, Canoe Pass 
Delta, bank between road and river, 1 Sept. 1957, 
W. Bird 3533 (BM). 

WASHINGTON. Marysville, fields, July 1927, 
J.M. Grant (US); Bank of R. Spokane, July 1931, 
Sister M. Milburge (WTU); Seattle, end of Green 
Lake, dumping ground, 28 July 1933, J.W. Thomp- 
son 9619 (NY); Port Blakely, in thicket, 17 June 
1934, W.J. Eyerdam (L, MO); Whatcom Co., Birch 
Bay, 19 July 1937, W.C. Muenscher 8343 (DAO); 
King Co., Fort Lawton, roadside hedges, 25 July 
1937, W.J. Eyerdam (EK MO); % mile NE of Seattle, 
near Sand Point, in thicket, 10 June 1949, W.J. 
Eyerdam (BM); Shelton—Woodsport road, moist 
sandy soil, 30 July 1950, P.E. Freer 320 (WTU). 

OREGON. Just east of Corvallis, across Van Bu- 
ren St. bridge, 15 June 1960, L.R.J. Dennis 2227 & 
G. van Vechten (NY). 

It is a pleasure to thank many botanists we have 
assisted me in the field in California on my three 
visits to look at Calystegia in 1997, 1998 and 1999, 
especially Barbara Ertter (Berkeley) and Tom Dan- 
iel (California Academy of Sciences). 


LITERATURE CITED 


BRUMMITT, R. K. 1972. Calystegia. Pp. 78-79 in T. G. 
Tutin et al., (eds.), Flora Europaea, Vol. 3. Cambridge 
University Press, Cambridge, U.K. 

BRUMMITT, R. K. AND A. O. CHATER. 2000. Calystegia 
(Convolvulaceae) hybrids in West Wales. Watsonia 
23:161—165. 

HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher 
plants of California. University of California Press, 
Berkeley, CA. 

Hitcucock, C. L. 1959. Pp. 85-89 in C. L. Hitchcock, A. 
Cronquist, and M. Ownbey (eds.), Vascular Plants of 
the Pacific Northwest, Vol 4. University of Washing- 
ton Publications in Botany, 17. University of Wash- 
ington Press, Seattle, WA. 


MADRONO, Vol. 49, No. 2, pp. 132—133, 2002 


NOTEWORTHY COLLECTIONS 


ARIZONA 


MANCOA PUBENS (A. Gray) Rollins (BRASSICA- 
CEAE).—Cochise County, San Pedro Riparian National 
Conservation Area, Kolbe site ca. 2 km SW of San Pedro 
Inn Bed and Breakfast, ca. 3 km S of Hereford Bridge, 
ca. 100 m E of San Pedro River. N31°24.961' 
W110°6.227’, elevation 1274 m, June 6, 2001. Found in 
sacaton grassland habitat with young mesquite; upper 
flood plain in sandy-loamy soil; level, open aspect. As- 
sociated species include Sporobolus wrightii, Prosopis ve- 
lutina, Erigeron concinnus, Xanthocephalum gymnosper- 
moides, Helenium thurberi, Conyza coulteri, Conyza can- 
adensis, Plantago sp., Salix goodingii, Elizabeth Makings 
365 (ASU). 

Previous knowledge. West Texas to Coahuila, Mexico 
(Rollins, The Cruciferae of Continental North America, 
Stanford University Press, 1993). Other regional collec- 
tions: Hinckley s.n. (ARIZ) Jefferson Davis County, Texas 
in 1937; Waterfall s.n. (ARIZ) Jefferson Davis County, 
Texas in 1943; Andres Rodriguez 861 (TEX) Coahuila, 
Mexico in 1983; four collections from Jefferson Davis 
County, Texas (TEX); two collections from Presidio 
County, Texas (TEX); and four collections from Brewster 
County, Texas (TEX). 

Significance. First record for the species in Arizona. 
This collection is approximately 500 km from nearest 
known collections in Jefferson Davis County, Texas. This 
was an isolated annual/biennial, inconspicuous in its in 
sacaton grassland community, which may explain why it 
has been under collected. This habitat is widely recog- 
nized as a major corridor for migratory birds. Seed size 
make wind dispersal unlikely, therefore, bird dispersal 
may explain the considerable range extension for this spe- 
cies. 


—ELIZABETH MAKINGS, Department of Plant Biology, 
Arizona State University, PO. Box 871601, Tempe, AZ 
85287-1601. 


W ASHINGTON 

BACCHARIS PILULARIS DC (ASTERACEAE).—Pacific 
Co: Fort Canby State Park, Beard’s Hollow. R11W, TION, 
S23, SW % of NE %4, USGS 7.5 minute “Cape Disap- 
pointment”’ quad, 30 m SW of Beard’s Hollow overlook 
on State Route 100, 1.5 km W of Ilwaco. Growing on SW 
facing cliffs in low coastal seacliff meadows above dune 
forest, with Poa unilateralis, Festuca rubra, Sedum ore- 
gonensis, Vulpia bromoides, and below Calamagrostis 
nutkaensis meadows. Colony of 30-50 upright plants, 
0.5—2 m tall, at 10-30 m elevation. One plant grows on 
sand 100 m to the north, at an elevation of 4 m, with 
Alnus rubra, Ammophila arenaria, Leymus mollis. First 
seen in 1981, revisited and collected while in flower on 
9-15-2001. K. Sayce (WS, OSC, WTU). 

Previous knowledge. Baccharis pilularis occurs from 
Oregon to northern Mexico on coastal bluffs to oak wood- 
lands, occasionally on serpentine, 0-750 (1500) m. The 
nearest known occurences are Gearhart and Cannon 
Beach, Clatsop County, Oregon (information supplied by 


Richard Halse, OSU Herbarium, Corvallis). Baccharis pi- 
lularis is increasingly common to the south, and is a mem- 
ber of shrub communities, including chaparral and coastal 
sage, in California. 

Significance. This is the northernmost known location 
and only Washington site for Baccharis pilularis, and is 
30 km north of the next known site in Oregon. This small 
population may represent remnants of a species that was 
more widespread in warmer climates. 


—KATHLEEN SAYCE, P.O. Box 91, Nahcotta WA 98637. 
kas @sbpac.com 


Moorea, SOCIETY ISLANDS, FRENCH POLYNESIA 


ARUNDO DONAX L. (POACEAE).—Poa Poa, University 
of California, Gump Biological Research Station, hillside 
grounds of main house and beyond, 17°30’S, 149°49’'w, 
elev. 100 m, 25 October 2000, Mitchel P. McClaran and 
James W. Bartolome 00-04 (ARIZ, UC). 

Previous knowledge. Native to Mediterranean (Allred 
1993, In: Hickman (ed.), The Jepson Manual, University 
of California Press, Berkeley, CA, p. 1235) cultivated 
worldwide, and regularly naturalized. Known from nearby 
Huahine and Raiatea islands (Welsh 1998, Flora Societen- 
sis, Electronic PrePrint Services Inc., Orem, UT). 

Significance. First record of this non-native species for 
Moorea. Several large plants spreading from landscape 
grounds to wild hillside. Based on evidence from Southern 
California, this species spreads via vegetative reproduc- 
tion, can dominate riparian vegetation through competitive 
exclusion, and can facilitate an increase in fire occurrence 
(Bell 1997, In: Brock et al. (eds.). Plant Invasions: Studies 
from North America and Europe. Backhuys Publishers, 
Leidens, Netherlands, p. 103-113). 


CHLORIS BARBATA Sw. (POACEAE).—Poa Poa, in waste 
area surrounding Supermarche Aré market, 17°29’S, 
149°49’'W, elev. 2 m, 23 October 2000, Mitchel P. Mc- 
Claran and James W. Bartolome OO-O1 (ARIZ, BISH, 
UC). 

Previous knowledge. Native to New World tropics and 
naturalized worldwide, and known from nearby Tahiti and 
Raiatea islands (Welsh 1998). 

Significance. First record of this non-native species for 
Moorea. Naturalized and common along roadsides and 
other waste areas near human settlements. This distribu- 
tion 1S consistent with observations from other South Pa- 
cific archipelagoes, including Hawaii, Samoa, Tonga, Fiji, 
Guam, and Belau (Whistler 1995, Wayside Plants of the 
Islands, Isle Botanica, Honolulu, HI). 


HYPARRHENIA RUFA (Nees) Stapf (POACEAE).—ca. 2 
km S of Afariatu, near and in abandoned livestock pasture, 
assoc. with Setaria sphacelata (Schumach.) Stapf & C.E. 
Hubb. ex. M.B. Moss, 17°34’S, 149°47'W, elev. 4 m, 24 
October 2000, Mitchel P. McClaran and James W. Bar- 
tolome 00-02 (ARIZ, BISH, UC). 

Previous knowledge. Native to tropical Africa and 
America (Renvoize 1984, The Grasses of Bahia, Kew Bo- 
tanical Garden, England). Known from nearby Tahiti and 
Raiatea islands (Welsh 1998). 


2002] 


Significance. First record of this non-native species for 
Moorea. Not common, but plants present beyond bound- 
aries of area seeded for livestock pasture. Naturalized, and 
likely to continue to spread based on experience in Aus- 
tralia, Hawaii, and Venezuela. Considered a weed in Aus- 
tralia because it spreads from seeded areas into native veg- 
etation (Lonsdale 1994, Australian Journal of Ecology 19: 
345-354). Spread increases the occurrence of fire, which 
hastens its increase and the decrease of native species in 
Hawaii and Venezuela (Smith and Tunison 1992, In: Stone 
et al. (eds.), Alien Plant Invasions in Native Ecosystems 
of Hawaii: Management and Research, University of Ha- 
wail Cooperative National Park Resources Study Unit, 
Honolulu, HI, p. 394—408, and Baruch 1996, In: Solbrig 
et al. (eds.), Biodiversity and Savanna Ecosystem Pro- 
cesses: A Global Perspective, Springer-Verlag, New York, 
NY, p. 79-93), although it will spread and dominate native 
vegetation in the absence of fire and grazing (San Jose 
and Farinas 1991, Acta Oecologia 12:237—247). 


SETARIA SPHACELATA (Schumach.) Stapf & C.E. Hubb. 
ex. M.B. Moss (POACEAE).—ca. 2 km §S of Afariatu, 
near and in abandoned livestock pasture, assoc. with Hy- 
parrhenia rufa (Nees) Stapf, 17°34'S, 149°47'W, elev. 4 
m, 24 October 2000, Mitchel P. McClaran and James W. 
Bartolome 00-03 (ARIZ, BISH, UC). 


NOTEWORTHY COLLECTIONS 133 


Previous knowledge. Native to tropical and subtropical 
Africa, and Yemen (Hacker 1992, In: ‘t Mannetje & Jones 
(eds.), Plant Resources of South-east Asia, No. 4, Pudoc 
Sci. Publ., Wageningen, Netherlands, p. 201—203). Known 
from seeded pasture, Taiarapu Plateau on nearby Tahiti 
island (Welsh 1998). 

Significance. First record of this non-native species for 
Moorea. Not common, but plants present beyond bound- 
aries of area seeded for livestock pasture. Possibly less 
likely to spread from seeded area than Hyparrhenia rufa 
based on its absence from Australian weed lists (Lonsdale 
1994). However, that prediction may be incorrect because 
this collection is apparently the more fecund cultivar 
‘Splenda’, based on intermediate values for some char- 
acters (number of culm nodes, culm diam., lf width, and 
panicle length) between vars. sericea and _ splendida 
(Hacker 1992). This cultivar, developed in Australia in the 
1980s, had high yields in Southeast Asian and Southern 
Pacific trials, and unlike the two varieties, will produce 
fertile seeds in abundance (Hacker 1992). 


—MITCHEL P. MCCLARAN, School of Renewable Natural 
Resources, 325 Bioscience East, University of Arizona, 
Tucson, AZ 85721 and James W. Bartolome, Department 
of Environmental Science, Policy, and Management, 145 
Mulford Hall, University of California, Berkeley, CA 
94720. 


MADRONO, Vol. 49, No. 2, p. 134, 2002 


REVIEW 


Seeing things whole: the essential John Wesley 
Powell. Edited by WILLIAM DEBuys. 2001. Island 
Press, Covelo, CA. 388 pp. $27.95. ISBN 1-55963- 
872-9. 


Guessing right about the direction history will 
take is always a doubtful proposition, but as Wil- 
liam deBuys makes clear in Seeing things whole: 
the essential John Wesley Powell, the famous one- 
armed explorer and scientist did guess right about 
enough important issues of his time to still be worth 
reading today. 

Actually, as deBuys makes clear, to say that 
Powell “‘guessed right’’ is not to do him the justice 
he deserves. A true polymath, Powell spent decades 
studying the landscape and peoples of the region 
he cared about—the American West. First as ex- 
plorer of the Grant Canyon, and then, over time, as 
geologist, ethnologist, bureau administrator, social 
planner, public figure, and philosopher, Powell 
worked over the same territory again and again 
from endlessly evolving perspectives, seeing pat- 
terns where his contemporaries clearly did not. 

Powell has not been ignored by the historians of 
recent times, and it is a tribute to the Major’s sig- 
nificance that he has been the subject of biographies 
by both Wallace Stegner and Donald Worster. What 
deBuys adds to these two excellent histories is 
Powell in his own words. This selection of Powell’s 


writings varies from excerpts from his well-known 
Colorado River writings, to things much less 
known including such forgotten statements as the 
Major’s address to Montana’s 1889 constitutional 
convention. Particularly welcome are several chap- 
ters from Powell’s often quoted (but not so often 
read) Report on the arid lands. 

What deBuys shows us is not a Powell who es- 
caped all the constraints of his times, but rather a 
powerful and original thinker who worked hard to 
see what was going on in his time. Certainly, by 
contemporary standards, Powell could not escape 
19th century expectations that the native people of 
North America were destined to disappear, and as 
an ecologist he had trouble (as many still do) com- 
ing to grips with the role of the fire in the West, 
but what deBuys shows clearly is that in a number 
of critical areas Powell did break through into new 
ways of thinking about a number of still critical 
issues. Certainly, his recognition that the nation’s 
land laws would victimize both many of those who 
tried to settle the West and the land itself, was un- 
precedented and still highly valid today. William 
deBuys is to be thanked for bringing a useful se- 
lection of Powell’s original writings back into easy 
reach. 


—WILLIAM TWEED. Sequoia-Kings Canyon National 
Parks, Three Rivers, CA 93271. 


Maprono, Vol. 49, No. 2, p. 135, 2002 


REVIEW 


Inventory of rare and endangered plants of Cali- 
fornia, sixth edition. California Native Plant Soci- 
ety, Rare Plant Scientific Advisory Committee, Da- 
vid P. Tibor, convening editor. 2001. California Na- 
tive Plant Society, Sacramento, CA. 388 pp. Soft- 
cover $29.95. ISBN 0-943460-40-9. 


The California Native Plant Society’s recent pub- 
lication of the sixth edition of the Inventory of rare 
and endangered plants of California offers a wel- 
come update on the constantly changing condition 
of endangered plants and habitats. Plants species’ 
rare, threatened or endangered status has expanded 
by a hefty 19% since the fifth edition, issued in 
1994, so it is an essential addition to any library. 

This version basically adheres to the same suc- 
cessful formula of its predecessors. However, some 
new features that appear in this /nventory are the 
inclusion of rare, non-vascular bryophytes (mosses, 
liverworts and hornworts), each species’ typical el- 
evation ranges, county indexing for all 2073 plants, 
and a new common name appendix. Inside covers 
have helpful lists of often-used, and too-often for- 
gotten acronyms. 

The Jnventory is a massive collaborative effort 
utilizing the talents of hundreds of scientific con- 
tributors who have spent untold hours in the field 
identifying, mapping, and gathering botanical data 
to be added to or updated in the Department of Fish 
and Game database. The importance of this work 
has its basis in that critical and balanced scientific 
input. But with a readership as diverse as devel- 
opers, biological consultants, educators and private 


landowners, the /nventory also must and does 
adeptly make this information understandable and 
available to all intellectual levels, thereby providing 
the greatest opportunity for early detection as well 
as avoidance of potential environmental conflicts. 

Introductory chapters espouse both CNPS poli- 
cies as well as those of various state & federal 
agencies—simplifying and clarifying the some- 
times overwhelming mass of plant protection 
guidelines and legislations. Peggy Fiedler again 
provides the scientific rationale for the plant endan- 
germent rating system. 

The actual “Threatened and Endangered Plant In- 
ventory’ is alphabetically organized by scientific 
name for easy access. In most cases, specific entries 
contain the following information: scientific names 
as well as family and common names, associated 
authorities, CNPS’ rarity rating (and those given 
state and Federal status as well), occurrences by 
county with map coordinates, key identifying char- 
acteristics, typical habitat and pertinent historical 
notations with literary referencing. 

The Inventory of rare and endangered plants is 
one of the most tangible and fruitful of CNPS’ 
many pursuits. Convening Editor Tibor and his le- 
gions of collaborators have not only achieved their 
intent for the /Jnventory, which encourages “ 
conservation planning and enforcement of environ- 
mental laws that protect rare species,’ they have 
ably furthered the goals of the California Native 
Plant Society. 


—MELANIE BAER-KEELEY. Division of Resources Man- 
agement, Branch of Vegetation Management, Sequoia 
Kings Canyon National Park, Three Rivers, CA 93271. 


MADRONO, Vol. 49, No. 2, p. 136, 2002 


CALIFORNIA BOTANICAL SOCIETY 
2002—2003 SCHEDULE OF SPEAKERS 


All Meetings are held at 7:30 p.m. on the 3rd Thursday of the month 


Sepia 


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Room 2063, Valley Life Sciences Building 
University of California, Berkeley 


Martin Bidartondo, Dept. of Plant Biology, UC Berkeley 
Obligate cheaters of mycorrhizal networks. 


Truman Young, Dept. of Environmental Horticulture, UC Davis 
Ecological restoration: an emerging conservation strategy. 


Randy Jackson, ESPM, UC Berkeley 
Spring-fed wetland structure and function in California oak savannas. 


David Ackerly, Dept. of Biological Sciences, Stanford University 
Fire, drought, and the evolution of chaparral shrubs. 


Annual banquet (University of San Diego) 
Jon Rebman, San Diego Natural History Museum 
Discoveries on a floristic frontier: Baja California 


Kim Steiner, California Academy of Sciences 
The evolution of a specialized pollinator system in southern Africa. 


Todd Dawson, Dept. of Integrative Biology, UC Berkeley 
Giants in the mist: coastal redwoods and the land-sea interface. 


Marcel Rejmanek, Div. of Biological Sciences, UC Davis 
Seed dispersal and coexistence of tree species in tropical forests 


Open to non-members. Refreshments served after the presentation. 


Volume 49, Number 2, pages 61—136, published 17 December 2002. 


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VOLUME 49, NUMBER 3 JULY-SEPTEMBER 2002 


RECEIVED BY: §§ F / 
INDEXING (-/- IND 


RIAN PENINSULA AND THE BALEARIC 

ISLANDS 

Roberto Gamarra.and Cindy Talbott ROCHE s.:....0020002000cs0eecsneceacessaseneenoens 7 
HyYMONOCLEAS ARE AMBROSIAS (COMPOSITAE) 

Jonni Strother and Bruce G. BOVAWUD so.diccescsssesscastasaincewaeedcaconaseesvaraeenes 143 
THE FLORA OF ToRA ISLET AND NOTES ON GUADALUPE ISLAND, BAJA CALIFORNIA, 

MEeExIco 

Jon P. Rebman, Thomas A. Oberbauer, and José Luis Leén de la Luz.... 145 
GENETIC STRUCTURE OF SENECIO LAYNEAE (COMPOSITAE): A RARE PLANT OF THE 

CHAPARRAL 

Glenda D. Marsh and Debra R. AYTES. «.polgg¥2)...2<02000:20pPh- aisle nonacsesesesestes 150 
Woop ANATOMY AND SUCCESSIVE CAMBIA IN SIMMONDSIA (SIMMONDSIACEAE): 

EVIDENCE FOR INCLUSION IN CARYOPHYLLALES S. L. 

SHOR WOR COI GUISE cook se mcncecc Ath Bc acacin aac ee NOR << pea cc. Notes svesencnetacescnes 158 
A NEw SUBSPECIES OF NAVARRETIA LEUCOCEPHALA (POLEMONIACEAE) FROM VERNAL 

POoLs IN EASTERN WASHINGTON 


Curtis R. BjOrRE Serge AOR A ee ON at oo RI). nn ceg cess ceeees 165 
RESURRECTION OF A CENTURY-OLD SPECIES DISTINCTION IN CALAMAGROSTIS 
Barbara YWAIsOn GAG SGHIMGFOVR Sr Nase IER wating nnn ee ToT OAR cones 169 


PLANT ZONATION IN A SHASTA COUNTY SALT SPRING SUPPORTING THE ONLY KNOWN 
POPULATION OF PUCCINELLIA HOWELLII (POACEAE) 
Lary evng Mary Bat ca GAD KAO¥ ul CRA re srs ikon eee cecceceeeseee 178 


SPHAGNUM BALTICUM IN A SOUTHERN RocKY MOUNTAIN IRON FEN 


David J. Cooper, Richard E. Andrus and Christopher D. Arp ..............+++- 186 
EVIDENCE OF A NOVEL LINEAGE WITHIN THE PONDEROSAE 

AnniM. Patten and Steven I. Bi HgfOl aed oo..c.ccscoincscSMede ah on odeevecasecosseseesse 189 
JTURSTOR pool acca ey Ueno Se | 22 eee 193 
COULTER 22 28 Re SAO oe 2 a See ae ena oe eee 193 
Ones re nee a ne See oe he eae «lj cusp vu Vana encvarsiuwassnendetcnveadecs 194 
oy a STEPTGHIGIN), 2s. Sees 8 00h 09 of Spa) AO Er atene On fee) Se ee ee ee ee 195 


A Cactus OpyssEy: JOURNEYS IN THE WILDS OF BOLIVIA, ARGENTINA, AND PERU, 
BY JAMES D. MAuUSETH, ROBERTO KIESLING, AND CARLOS OSTOLAZA 
Mae cateeBtatea Cork FAG NN TE ee ea note ens a er Setice tobi o css onaces Suansdinesedsibdieibineueseecres 198 


BIENNIAL GRADUATE STUDENT MEETING AND ANNUAL BANQUET ..........cesceecceecceeceeees 199 
Eon A eae ys re hoe oan Raney sive cddeesevaserassaswcevesene 200 


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Maprono, Vol. 49, No. 3, pp. 137—142, 2002 


DISTRIBUTION OF THE GENUS CRUPINA IN THE IBERIAN PENINSULA 
AND THE BALEARIC ISLANDS 


ROBERTO GAMARRA 
Dpto. de Biologia (Botanica), Facultad de Ciencias, 
Universidad Autonoma de Madrid, Cantoblanco, E-28049 Madrid, Spain 


CINDY TALBOTT ROCHE* 
109 Meadow View Drive, Medford, OR 97504 USA 
crupinaqueen @charter.net 


ABSTRACT 


Crupina vulgaris, with populations in Sonoma and Modoc counties, is a relatively recent Mediterranean 
invader among the 1045 alien plant species naturalized in California, apparently introduced from the 
Iberian Peninsula. Although the genus Crupina comprises two species in Spain, only one is known in the 
Western Hemisphere. This study compiled distribution maps for both C. vulgaris and C. crupinastrum in 
the Iberian Peninsula using 939 collections from 17 herbaria in Spain and 154 locations recorded in the 
literature (floras, botanical explorations, and phytosociological studies). Crupina vulgaris is more com- 
mon, with a distribution occupying most of the eastern half of the Peninsula, reaching north to the southern 
slopes of the Pyrenees Mountains which form the boundary between Spain and France. It is rare in the 
western third of the Peninsula, including Portugal, and is absent on the Balearic Islands. Found throughout 
the southeastern quadrant of the Peninsula and Mallorca in the Balearic Islands, Crupina crupinastrum 
is most abundant in the Andalusian provinces. In southeastern Spain where their distributions overlap, C. 
crupinastrum is more prevalent than C. vulgaris. Detailed distribution maps in the native range of invasive 
species are useful for understanding biological invasions, comparisons of native and introduced habitats, 
and searches for potential biological control agents. 


RESUMEN 


Crupina vulgaris es una planta de origen mediterraneo, de entre las 1045 especies naturalizadas en 
California, que se comporta como invasora y cuenta con poblaciones, relativamente recientes, en los 
condados de Sonoma y Modoc. Los estudios moleculares han demostrado que las cinco poblaciones 
norteamericanas derivan de 3 0 mas introducciones procedentes de la Peninsula Ibérica. Aunque el género 
Crupina comprende dos especies en Espana, solamente una es conocida en el hermisferio occidental. En 
este estudio se presentan los mapas de distribuci6n de C. vulgaris y C. crupinastrum en la Peninsula 
Ibérica, obtenidas tras la revision de 939 recolecciones encontradas en 17 herbarios de Espana y 154 
referencias bibliograficas (floras, catalogos floristicos, y estudios fitosociolé6gicos). Crupina vulgaris es la 
especie mas comun, con una distribuci6n que ocupa la mitad oriental de la Peninsula, alcanzando hacia 
el Norte las vertientes meridionales de los Pirineos. Es rara en el tercio oeste, incluyendo Portugal, y esta 
ausente en las islas Baleares. Presente en todo el cuadrante suroriental de la Peninsula y en Mallorca 
(islas Baleares), Crupina crupinastrum es mas abundante en las provincias andaluzas. En el Sureste de 
Espana, donde ambas distribuciones se solapan, C. crupinastrum prevalece sobre C. vulgaris. Los mapas 
de distribuci6n detallados de las especies invasoras en su zona de origen, son utiles para comprender las 
invasiones, la comparaci6n entre habitat autdctonos y aléctonos, y las investigaciones para buscar poten- 
ciales agentes de control bioldégico. 


Key words: Crupina crupinastrum, Crupina vulgaris, invasive species, weed distribution 


Among the 1045 alien plant species naturalized 
(17.5% of the flora) in California (Randall et al. 
1998), Crupina vulgaris Cass. (Asteraceae: Cyna- 
reae) is a relatively recent addition. Rejmanek and 
Randall (1994) indicated that it was very likely in- 
troduced to California within the past 25 years. This 
contrasts sharply with numerous other Mediterra- 
nean species introduced in the Spanish and Mexi- 
can periods (1769-1848) (Bossard et al. 2000). 
When Crupina vulgaris was discovered in Idaho in 
1968, it was reported in Madrofio as a species new 


* Corresponding author. 


to North America (Stickney 1972). Under the Fed- 
eral Noxious Weed Act of 1974 (Public Law 93 
629), Crupina vulgaris was included on the Federal 
Noxious Weed List as a new invader. Both Crupina 
vulgaris and its congener Crupina crupinastrum 
(Moris) Vis. were among 1200 species designated 
as economically important foreign weeds posing 
potential problems in the United States (Reed 
1977). A search of the interception records for Fed- 
eral Noxious Weeds by USDA APHIS PPQ from 
1983 through 1998 showed that Crupina vulgaris 
has not been intercepted crossing the U.S. borders 
since it was listed and records kept on weed inter- 


138 


ventions (Polly Lehtonen, USDA APHIS, personal 
communication). 

Although the first flora in California to include 
Crupina vulgaris was The Jepson Manual (Hick- 
man 1993), it was first discovered in California in 
1976 at Santa Rosa, Sonoma County (unpublished 
CDFA reports, Miller and Thill 1983). It was sub- 
sequently discovered in 1984 at Lake Chelan, Che- 
lan County, Washington (Alverson and Arnett 
1986), and in 1987 at Dry Creek, Umatilla County, 
Oregon (Couderc-LeVaillant and Roché 1993). The 
Sonoma County population was declared eradicated 
by 1982 (Miller and Thill 1983), but it was redis- 
covered in 1989 about 1 km distant from the orig- 
inal infestation (unpublished CDFA records, Davis 
and Sherman 1991). In 1990 another new popula- 
tion was reported near Adin, Modoc County, Cali- 
fornia (unpublished CDFA records, Couderc-Le- 
Vaillant and Roché 1993). Thus, five infestations in 
four western States were detected between 1968 
and 1990. Crupina vulgaris is inconspicuous due to 
its small, delicate stature and extremely difficult to 
detect when populations are sparse. All six original 
discoveries of C. vulgaris in the western United 
States were made by professional botanists or weed 
scientists; none were reported by casual observers 
or landowners. 

Multiple introductions have been suspected for 
the invasion (Couderc-Le Vaillant and Roché 1993; 
Patterson and Mortensen 1985), although a large 
propagule hinders long distance dispersal (Roché 
and Thill 2001). Since the previous publications in 
Madrono, earlier collections of C. vulgaris in North 
America have been reported from Massachusetts 
(Sorrie and Somers 1999). Specimens at the Har- 
vard University Herbaria collected by C.E. Perkins 
in 1877 and 1879 from Boston and South Boston 
Flats, Suffolk County, Massachusetts (NEBC, W. 
Kittridge personal communication), indicate that C. 
vulgaris was among the numerous species intro- 
duced in ship’s ballast from seaports in the Medi- 
terranean region. Based on its absence in current 
floras, C. vulgaris failed to establish in the north- 
eastern United States, and apparently arrived inde- 
pendently in western North America. Recent mo- 
lecular studies (RAPD) revealed that the five cur- 
rent populations derived from three or more intro- 
ductions from the Iberian Peninsula (Garnatje et al. 
2002). Although the genus Crupina is represented 
by two species in the Iberian Peninsula and Bale- 
aric Islands, C. crupinastrum and C. vulgaris, only 
one has been reported in North America. 

The objective of the study was to compile an 
accurate distribution map for Crupina in the part of 
its native range where its North American popula- 
tion founders originated. Such a map would serve 
as a foundation for further investigations, such as 
ecological studies comparing native and introduced 
habitats, including behavior of the invader, and fac- 
tors that contribute to differences in species re- 
sponse in the two hemispheres. A distribution map 


MADRONO 


[Vol. 49 


is also useful for searches for potential biological 
control agents. Floras provide general distribution 
information that is inadequate for these purposes. 
For example, Flora Europaea indicates that C. vul- 
garis grows on “dry grassland and stony slopes”’ 
in 17 countries (Amaral Franco 1976), which leads 
one to believe that it can be easily encountered any- 
where in these habitats in southern Europe, as far 
north as west central France and the southern 
Ukraine. In fact, much of this distribution is based 
on centuries-old collection records, some of which 
represent localized populations that failed to persist 
under changing land use patterns in the past 100 
years. In our study we included the more ruderal 
(within the Mediterranean region) congener, C. cru- 
pinastrum, in order to provide supporting infor- 
mation for inferences about plant migration and the 
invasion process. 


METHODS 


A complete listing of all recorded locations for 
both species of Crupina in the Iberian Peninsula 
was compiled from two types of sources: 1) liter- 
ature citations of locations from floras, botanical 
explorations, and phytosociological studies and 2) 
herbarium specimens from 17 herbaria in Spain 
(listed in the acknowledgments, with institutional 
abbreviations from Holmgren et al. 1990). Because 
the two species closely resemble each other, each 
herbarium sheet was examined and annotated. On 
mature specimens, verification was based on cyp- 
sela characters, while immature specimens and oth- 
ers without fruits were verified using trichome char- 
acters (Couderc-LeVaillant 1984). Maps were pre- 
pared using software (CYANUS) based on 10 km 
UTM grid square. 


RESULTS 


The distributions of C. vulgaris and C. crupinas- 
trum are shown on maps in Figs. 1 and 2, respec- 
tively. Of the 939 herbarium specimens examined, 
572 were assigned to C. vulgaris and 367 to C. 
crupinastrum. The maps include 134 citations from 
the literature for which there was no doubt about 
the identification. An additional 120 literature ci- 
tations for C. vulgaris were not included on the 
maps because they lacked corroborating herbarium 
vouchers and were from regions where the distri- 
bution of C. crupinastrum overlapped with that of 
C. vulgaris. In some provinces there were locality 
descriptions for which UTM coordinates could not 
be determined. A complete record of herbarium la- 
bel and literature citations will be published in 
Spain (Gamarra and Roché 2002). 


Crupina vulgaris 


In the Iberian Peninsula, C. vulgaris is the more 
common species, with a distribution occupying 
most of the eastern half of the Peninsula (Fig. 1). 
All of the phytosociology literature and plant dis- 


2002] 


‘Dizerress 


= ee 


GAMARRA AND ROCHE: CRUPINA IN THE IBERIAN PENINSULA 139 


e 

EDS 
Ne 1 

e s ses 4 al 


‘ oN 
codes Sa Na 


36° 


Fic. 1. 


Distribution of Crupina vulgaris in the Iberian Peninsula, each symbol indicates documented presence within 


a 10 km UTM grid square. Dotted lines are provincial boundaries; refer to Fig. 2 for names. 


tribution records indicate that C. vulgaris is not 
found in forested areas and in subalpine grasslands, 
as well as areas of siliceous soils, which tend to be 
coarse and acidic. It does not occur on the Balearic 
Islands. The most northerly populations of C. vul- 
garis reach the southern slopes of the Pyrenees and 
the Pre-Pyrenean Mountains. In the western third 
of the peninsula its presence is sparser and almost 
always tied to calcareous substrates, which are in- 
frequent in this region. In Portugal it is very rare, 
limited to certain populations near the locality of 
Elvas, very close to the border with Spain, and far- 
ther away, in Sezimbra, but always on calcareous 
soils. Using locations from herbarium labels dating 
from the 1970’s, one of the authors (Roché) and a 
colleague searched all suitable habitat in the Elvas 
locality in 1999 and failed to find a single plant. If 
C. vulgaris is still present there, it is extremely 
scarce. 

According to Rivas Martinez et al. (1990), this 
species appears in all the chorologic provinces, al- 
though in the north and northeastern provinces of 
Cantabro-Atlantica and Gaditano-Onubo-Algar- 
viense, it occurs only in isolated locations. As stat- 


ed earlier, in the history of Spanish botanical sci- 
ence, it has never been found in the Balearic prov- 
ince. 

The elevation within its distribution ranges be- 
tween 100 and 1200 (rarely 1500) m, reaching the 
major territory of the Sierra de Segura and the Pyr- 
enees, but it is never found in the high mountains. 
It prefers basic soils, principally calcareous soils 
derived from limestone or clay soils rich in bases 
originating from evaporites (formed by the evapo- 
ration of brackish water), including substrates rich 
in gypsum; and is only rarely found over schist or 
slate. It is reported from rangeland, dry grazed ar- 
eas, low matorral (e.g., thyme fields), open ever- 
green oak woodlands, and clearings in deciduous 
oak forests. It is also found along roadsides and on 
the margins of perennial crops such as vineyards or 
olive groves where they border appropriate native 
habitat. 


Crupina crupinastrum 


This species primarily inhabits the southeastern 
quadrant of the peninsula and the Balearic Islands, 


140 MADRONO 


[Vol. 49 


Pizerress 


Fic. 2. 


42° | 


| | 


Distribution of Crupina crupinastrum in the Iberian Peninsula, each symbol indicates documented presence 


within a 10 km UTM grid square. Abbreviations for names of provinces: Alicante (A), Albacete (Ab), Almeria (Al), 
Asturias (O), Avila (Av), Badajoz (Ba), Barcelona (B), Burgos (Bu), Caceres (Cc), Cadiz (Ca), Cantabria (S), Castellon 
(Cs), Cordoba (Co), Ciudad Real (CR), Cuenca (Cu), Gerona (Ge), Granada (Gr), Guadalajara (Gu), Huelva (H), Huesca 
(Hu), Jaen (J), La Corufia (C), La Rioja (Lo), Ledn (Le), Lerida (L), Lugo (Lu), Madrid (M), Malaga (Ma), Murcia 
(Mu), Navarre (Na), Orense (Or), Palencia (P), Pontevedra (Po), Salamanca (Sa), Sevilla (Se), Segovia (Sg), Soria (So), 
Tarragona (T), Teruel (Te), Toledo (To), Valencia (V), Valladolid (Va), Vizcaya (Vi), Guipuzcoa (SS), Zamora (Za), 


Zaragoza (Z). 


and is most abundant in the Andalusian provinces 
(Almeria, Cadiz, Cordoba, Granada, Huelva, Jaén, 
Malaga, Sevilla) (Fig. 2). In contrast to the northern 
presence of C. vulgaris, C. crupinastrum only ex- 
tends as far north as the provinces of Segovia and 
Burgos, where it is rarely encountered and could be 
interpreted as isolated individuals occurring as 
ephemeral introductions. In the Balearic Islands, it 
is found only on Mallorca. It does not appear on 
the Pitiusas Islands (the southwest islands within 
the Balearic group), which would be an extension 
of the levantan populations. Toward western Iberia, 
it is very rare and we found only isolated occur- 
rences in the provinces of Sevilla (Castilleja de 
Guzman) and Caceres (Guadalupe), which did not 
extend as far as Portugal. 

According to Rivas Martinez et al. (1990), this 
species appears principally in the provinces Bética 
and Castellano-Maestrazgo-Manchega, with some 


presence in the provinces Murciano-Almeriense 
and Balear, and is very rare in the Luso-Extrema- 
durense (Portugal, Caceres and Badajoz). 

The presence of Crupina crupinastrum is linked 
to basic soils, principally substrates of limestone 
and evaporites, although some populations in the 
province of Cérdoba grow over schists. The ele- 
vation ranges between 100 and 1500 m, very rarely 
exceeding this limit, but occasionally doing so in 
the more southern mountains as in Gador and in 
the Sierra Nevada. It appears on the sides of roads, 
grazed lands, pastures, low matorral, and open ev- 
ergreen oak woodlands. 


DISCUSSION 


Both species of Crupina share the southeastern 
quadrant of the Iberian Peninsula, in some cases 
growing in mixed communities. In this region C. 


2002] 


crupinastrum is much more abundant than C. vul- 
garis. However, nowhere in the Iberian Peninsula 
would one describe either species as abundant in 
the plant communities where they occur. It is note- 
worthy that the more ruderal C. crupinastrum, 
which appears more frequently in disturbed sites 
(e.g., roadsides) than C. vulgaris, was not the spe- 
cies introduced in North America. Because cypselas 
of the two species are the same size with the same 
pappus characteristics, differing only in shape at the 
point of attachment (see illustrations in Reed 1977), 
this anomaly suggests that the invasion founders 
originated from locations where C. crupinastrum 
does not grow along with C. vulgaris. 

On sites supporting Crupina in the Iberian Pen- 
insula, grazing of sheep and goats is the primary 
land use, especially historically. In the Mediterra- 
nean region, Crupina has been identified among 
low chaparral species which maintain populations 
by epizooic transport associated with herds of do- 
mestic sheep and goats (Schmida and Ellner 1983). 
Transhumanant herds of sheep were likely the mi- 
gratory vectors responsible for dispersing and 
maintaining isolated ephemeral populations in the 
Iberian Peninsula. Well established trails (cafiadas) 
connect summer and winter pastures (Montserrat 
and Fillat 1990), which may be as close as moun- 
tain grasslands with adjacent valleys, or extend 
nearly the entire north-south distance of Spain 
(Mangas 1992). 

The distribution of C. vulgaris far to the north 
of C. crupinastrum in the Iberian Peninsula is es- 
pecially significant because the Pyrenean and Pre- 
Pyrenean Mountain region was the source of major 
immigration to the United States of laborers for the 
sheep industry through the early 1970’s. Basque 
sheepherders were legendary in the western U.S. 
For example, in 1970 about 90% of the 1700 men 
under contract to the Western Range Association 
were Basques (Lane and Douglass 1985). Although 
the Spanish government required equal immigra- 
tion opportunities for all Spanish nationals, it was 
primarily due to improving economic conditions in 
the Basque Country in the 1960’s and 1970’s that 
the numbers of Asturians, Leonese, Castillians and 
Andalusians swelled the herder ranks (Lane and 
Douglass 1985). After that time, this link with rural 
Spain ended when the range association shifted its 
herder recruitment efforts to Latin America and 
Mongolia. 

Despite the recent (1968—1990) discoveries of C. 
vulgaris in western North America, it is probable 
that it arrived decades earlier. Arriving in small 
numbers to remote areas, with slow growing colo- 
nies of inconspicuous individuals, it is not surpris- 
ing that it could escape detection for long periods 
of time. By showing where the distributions of C. 
vulgaris and C. crupinastrum overlap and diverge, 
and where the distribution of C. vulgaris overlaps 
a region that was home to numerous immigrants to 
rangelands of the western United States, these dis- 


GAMARRA AND ROCHE: CRUPINA IN THE IBERIAN PENINSULA 141 


tribution maps provide a resource for elucidating 
the case history of how C. vulgaris became an in- 
vader in the Western Hemisphere, a chronicle that 
has not yet been fully revealed. They also serve as 
a reference for future studies concerning ecology 
and potential control. 


ACKNOWLEDGMENTS 


The following herbaria in Spain graciously loaned or 
allowed us to review their materials: Sociedad de Ciencias 
Aranzadi, San Sebastian (ARAN): Institut Botanic de Bar- 
celona, Barcelona (BC); Universitat de Barcelona (Depar- 
tament de Biologia Vegetal), Barcelona (BCC); Univer- 
sitat de Barcelona (Facultat de Farmacia), Barcelona 
(BCF); Universidad de Cérdoba, Cérdoba (COFC); Uni- 
versidad de Granada, Granada (GDA): Universidad de 
Granada, Granada (GDAC); Universitat de Girona, Girona 
(HGI); Instituto Pirenaico de Ecologia, Jaca (JACA): Co- 
legio Universitario Santo Reino, Jaén (JAEN); Real Jardin 
Botanico, Madrid (MA); Universidad Complutense (Cien- 
cias Biolégicas), Madrid (MACB); Universidad Complu- 
tense (Farmacia), Madrid (MAF); Universidad de Malaga, 
Malaga (MGC); Universidad de Sevilla, Sevilla (SEV): 
Universitat de Valéncia, Valencia (VAL); and Museo de 
Ciencias Naturales de Alava, Vitoria (VIT). We thank the 
following individuals for their generous sharing of their 
time and knowledge: Daniel Gémez, Federico Fillat, Mi- 
kel Lorda, Pedro Uribe, Mauricio Velayos, José Pizarro, 
Josep Vicens and Antonio Sanchez-Cuxart. The manu- 
script was reviewed by K. L. Chambers. C. Roché extends 
special thanks to Josep Montserrat, for his encouragement 
and his suggestions of contacts for the project. 


LITERATURE CITED 


ALVERSON, E. AND J. ARNETT. 1986. Plant life of the North 
Cascades: Lake Chelan-Sawtooth Ridge, Stehekin 
Valley and Glacier Peak. Douglasia Occasional Paper, 
Vol. 2. Washington Native Plant Society, Seattle, WA. 

AMARAL FRANCO, J. 1976. Crupina. P. 301 in T. G. Tutin 
et al. (eds.), Flora Europaea, Vol. 4. Cambridge Univ. 
Press, London, U.K. 

BOSSARD, C. C., J. M. RANDALL, AND M. C. HOSHOVSKY 
(eds.). 2000. Invasive plants of California’s wildlands. 
University of California Press, Berkeley, CA. 

COUDERC-LEVAILLANT, M. 1984. L-amphiploidie dans le 
genre Crupina DC. Essai de systématique synthéti- 
que. Thése Docteur Es-Sciences Naturelles, Univ. 
Paris-Sud, Orsay, France. 

AND C. T. ROCHE. 1993. Evidence of multiple in- 
troduction of Crupina vulgaris in infestations in the 
western United States. Madrono 40:63-—65. 

Davis, L. H. AND R. J. SHERMAN. 1991. Crupina vulgaris 
Cass. (Asteraceae: Cynareae), established in Sonoma 
County, California, at Annadel State Park. Madrono 
38:296. 

GAMARRA, R. AND C. T. ROCHE. 2002. Cartografia Coro- 
l6gica Ibérica. Aportaciones. Botanica Complutensis. 
Universidad Complutense, Madrid, Spain. 

GARNATIE, T., R. VILATERSANA, C. T. ROCHE, N. GARCIA- 
JACAS, A. SUSANNA, AND D. C. THILL. 2002. Multiple 
introductions from the Iberian Peninsula responsible 
for invasion of Crupina vulgaris Cass. in western 
North America. New Phytologist 154:419—428. 

HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher 
plants of California. University of California Press, 
Berkeley, CA. 


142 MADRONO 


HOLMGREN, P. K., N. H. HOLMGREN, AND L. C. BARNETT. 
1990. Index herbariorum, 8th ed. New York Botanical 
Garden, Bronx, NY. 

LANE, R. H. AND W. A. DouGLass. 1985. Basque sheep- 
herders of the American West. University of Nevada 
Press, Reno, NV. 

MANGAS, J. M. 1992. Vias pecuarias. Cuadernos de la tras- 
humancia, No. 0, Instituto Nacional para la Conser- 
vacion de la Naturaleza, Madrid, Spain. 

MILLER T. AND D. THILL. 1983. Today’s weed: common 
crupina. Weeds Today 14:10—11. 

MONTSERRAT, P. AND FE FILLAT. 1990. The systems of 
grassland management in Spain. Pp. 37-70 in A. 
Breymeyer (ed.), Managed grasslands, ecosystems of 
the world. Elsevier, Amsterdam, Netherlands. 

PATTERSON, D. E. AND D. A. MorTENSEN. 1985. Effects of 
temperature and photoperiod on common crupina 
(Crupina vulgaris). Weed Science 33:333-339. 

RANDALL, J. M., M. REJMANEK, AND J. C. HUNTER. 1998. 
Characteristics of the exotic flora of California. Fre- 
montia 26(4):3—-12. 

REED, C. E 1977. Economically important foreign weeds. 
Ag. Handbook No. 498. U.S. Govt. Printing Office, 
Washington, DC. 


[Vol. 49 


REJMANEK, M. AND J. M. RANDALL. 1994. Invasive alien 
plants in California: 1993 summary and comparison 
with other areas in North America. Madrono 41:161— 
Wa Tle 

RivAs MARTINEZ, S., P. CANTO, E FERNANDEZ GONZALEZ, 
C. NAVARRO, J. M. PIZARRO, AND D. SANCHEZ MATA. 
1990. Biogeografia de la Peninsula Ibérica, Islas Ba- 
leares y Canarias. X Jornadas de Fitosociologia, Gra- 
nada, Spain. 

ROCHE, C. T. AND D. C. THILL. 2001. Biology of common 
crupina and yellow starthistle, two Mediterranean 
winter annual invaders in western North America. 
Weed Science 49:439—447. 

SCHMIDA, A. AND S. ELLNER. 1983. Seed dispersal on pas- 
toral grazers in open Mediterranean chaparral, Israel. 
Israel Journal of Botany 32:147—159. 

SORRIE, B. A. AND P. Somers. 1999. The vascular plants 
of Massachusetts: a county checklist. Massachusetts 
Division of Fisheries and Wildlife, Natural Heritage 
and Endangered Species Program, Westborough, MA. 

STICKNEY, P. E 1972. Crupina vulgaris (Compositae: 
Cynareae), new to Idaho and North America. Madro- 
no 21:402. 


MaApbroNo, Vol. 49, No. 3, pp. 143-144, 2002 


HYMENOCLEAS ARE AMBROSIAS (COMPOSITAE) 


JOHN L. STROTHER 
University Herbarium, 
University of California, Berkeley, CA 94720-2465 
strother @uclink4.berkeley.edu 


BRUCE G. BALDWIN 
Jepson Herbarium and Department of Integrative Biology, 
University of California, Berkeley, CA 94720-2465 


ABSTRACT 


Inclusion of Hymenoclea within the taxonomic circumscription of Ambrosia necessitates new combi- 
nations: Ambrosia monogyra, A. xplatyspina, A. salsola, A. salsola var. fasciculata, and A. salsola 


var. pentalepis. 


Key words: Ambrosia, Compositae, Hymenoclea 


After review of similarities and differences be- 
tween and among species of Hymenoclea Torrey & 
A. Gray ex A. Gray and Ambrosia Linnaeus, es- 
pecially with regard to restriction sites in chloro- 
plast DNAs, Miao et al. (1995) concluded that the 
two species of Hymenoclea do not constitute a 
clade, are separately allied to franserioid members 
of Ambrosia, and are better included in Ambrosia 
than maintained as a distinct genus. They listed Hy- 
menoclea as a synonym of Ambrosia Linnaeus 
subg. Franseria (Cav.) Miao et al., Pl. Syst. Evol. 
194:252, 1995. 

Baldwin et al. (1996) documented natural hy- 
bridization between Hymenoclea salsola Torrey & 
A. Gray ex A. Gray and the franserioid species Am- 
brosia dumosa (A. Gray) W. W. Payne and between 
H. salsola and A. ambrosioides (Cavanilles) W. W. 
Payne, another franserioid species. They noted “.. . 
normal pairing of chromosomes in interspecific hy- 
brids”’ (1.e., between species of Ambrosia and Hy- 
menoclea) as indicating ““Close genetic similarity 

” of parental species and went on to say, ““Hy- 
bridization between species of Ambrosia and Hy- 
menoclea may reflect inadequacy of the long-stand- 
ing generic classification of Ambrosiinae ... .” 

We have considered the findings of Miao et al. 
(1995) and Baldwin et al. (1996) and we are con- 
vinced that hymenocleas should be treated within 
the taxonomic circumscription of Ambrosia: 


Ambrosia monogyra (Torrey & A. Gray ex A. 
Gray) Strother & B. G. Baldwin, comb. nov. 
Basionym: Hymenoclea monogyra Torrey & A. 
Gray ex A. Gray, Mem. Amer. Acad. Arts ns. 4: 
79. 1849. Syntypes: “‘Along the valley of the 
Gila, Lieut. Emory. Also at ‘Ojito,) New Mexi- 
co? Dr. Gregg.’ Peterson and Payne (1973, p. 
253-254) cited the Emory collection at NY as 
type and thereby effected lectotypification. 

Ambrosia salsola (Torrey & A. Gray ex A. Gray) 


Strother & B. G. Baldwin, comb. nov. Basionym: 
Hymenoclea salsola Torrey & A. Gray ex A. 
Gray, Mem. Amer. Acad. Arts n.s. 4:79. 1849. 
Type: California, ““Sandy, saline uplands near the 
Mojave River, ..., Fremont.’’ Lectotype (Peter- 
son and Payne 1973, p. 254): 1844, Frémont 400 
(NY). 

Ambrosia salsola (Torrey & A. Gray ex A. Gray) 
Strother & B. G. Baldwin var. fasciculata (A. 
Nelson) Strother & B. G. Baldwin, comb. nov. 
Basionym: Hymenoclea fasciculata A. Nelson 
var. fasciculata. [cf. Hymenoclea fasciculata A. 
Nelson, Bot. Gaz. 37:270. 1904. Type: Nevada, 
““Kernan,”’ 29 Apr 1902, L. N. Goodding 662 
(NY). The variety fasciculata dates from publi- 
cation of Hymenoclea fasciculata A. Nelson var. 
patula A. Nelson, Bot. Gaz. 47:431. 1909. Type: 
Nevada, “‘“Moapa,”’ 8 Apr 1905, L. N. Goodding 
2178 (RM). Peterson and Payne (1973, 1974) 
treated the two types, Goodding 662 and 2178, 
as convarietal. Initially, Peterson and Payne 
named that variety Hymenoclea salsola Torrey & 
A. Gray ex A. Gray var. fasciculata (A. Nelson) 
K. M. Peterson & W. W. Payne (Brittonia 25:255. 
1973). Under the applicable Code (Stafleu et al. 
1972) in 1973, Peterson and Payne should have 
used the varietal epithet “‘patula’’ because auto- 
nyms were “not to be taken into consideration 
for purposes of priority.”’ Peterson and Payne 
subsequently renamed that same variety Hymen- 
oclea salsola Torrey & A. Gray ex A. Gray var. 
patula (A. Nelson) K. M. Peterson & W. W. 
Payne (Brittonia 26:397. 1974), which is now an 
illegitimate name because under the current Code 
(Greuter et al. 2000), the autonymic varietal 
name has priority at varietal rank. ] 

Ambrosia salsola (Torrey & A. Gray ex A. Gray) 
Strother & B. G. Baldwin var. pentalepis (Ryd- 
berg) Strother & B. G. Baldwin, comb. nov. Bas- 
ionym: Hymenoclea pentalepis Rydberg in N. L. 


144 MADRONO 


Britton et al., N. Amer. Fl. 33:14. 1922. Type: 
Arizona, “‘Pima Canon,”’ 10 Apr 1901, D. Grif- 
fiths 2630 (NY) = Hymenoclea salsola Torrey & 
A. Gray ex A. Gray var. pentalepis (Rydberg) L. 
D. Benson, Amer. J. Bot. 30:631. 1943. 

Hymenoclea hemidioica A. Nelson, Amer. J. Bot. 
25:117. 1938. Syntypes: Arizona, Mohawk 
Mountains, 29 Mar 1935, A. Nelson 1340 and 
1341 (RM?, not seen). 


We use the name Ambrosia xplatyspina (Sea- 
man) Strother & B. G. Baldwin, comb. nov. [Bas- 
ionym: Hymenoclea xplatyspina Seaman, Madrono 
23:111. 1975, pro sp.], for hybrids between A. du- 
mosa and A. salsola. 

We believe the name Ambrosia sandersonii S. L. 
Welsh, Rhodora 95:396. 1993[1994] [=Hymeno- 
clea sandersonii (S. L. Welsh) N. H. Holmgren in 
A. Cronquist et al., Intermount. Fl. 5:473. 1994], 
also refers to hybrids, perhaps to hybrids between 
A. eriocentra (A. Gray) W. W. Payne and A. salsola 
(Baldwin et al. 1996). We treat Welsh’s name as 
Ambrosia xsandersonii S. L. Welsh, pro sp. 


[Vol. 49 


ACKNOWLEDGMENTS 


We thank D. J. Keil, T. K. Lowrey, R. L. Moe, and A. 
R. Smith for helpful comments on early drafts of this pa- 
per. 


LITERATURE CITED 


BALDwIn, B. G., D. W. KyuHos, S. N. MARTENS, FE C. Va- 
SEK, AND B. L. WEsSA. 1996. Natural hybridization 
between species of Ambrosia and Hymenoclea sal- 
sola (Compositae). Madrofo 43:15—27. 

GREUTER, W., ET AL. 2000. International code of botanical 
nomenclature. Regnum Vegetabile 138:v—xvili, 1— 
474. 

Miao, B., B. TURNER, B. SIMPSON, AND T. MABRY. 1995. 
Chloroplast DNA study of the genera Ambrosia s.1. 
and Hymenoclea (Asteraceae): systematic implica- 
tions. Plant Systematics and Evolution 194:141—255. 

PETERSON, K. M. AND W. W. PAYNE. 1973. The genus Hy- 
menoclea (Compositae: Ambrosieae). Brittonia 25: 
243-256. 

AND . 1974. Erratum: the correct name for 
the appressed-winged variety of Hymenoclea salsola 
(Compositae: Ambrosieae). Brittonia 26:397. 

STAFLEU, FE A., ET AL. 1972. International code of botan- 
ical nomenclature. Regnum Vegetabile 82:7—426. 


Mapbrono, Vol. 49, No. 3, pp. 145-149, 2002 


THE FLORA OF TORO ISLET AND NOTES ON GUADALUPE ISLAND, 
BAJA CALIFORNIA, MEXICO 


JON P. REBMAN 
San Diego Natural History Museum, P.O. Box 121390, 
San Diego, CA 92112-1390, USA 
jrebman @sdnhm.org 


THOMAS A. OBERBAUER 
County of San Diego, Department of Planning and Land Use, 
5201 Ruffin Rd. Suite B5, San Diego, CA 92123, USA 


Jose Luts LEON DE LA Luz 
Centro de Investigaciones Bioldégicas del Noroeste, Apdo. Postal 128, 
La Paz, 23000, Baja California Sur, México 


ABSTRACT 


During a natural history expedition to Guadalupe Island and its adjacent islets in June of 2000, the 
previously unexplored islet, Toro, was botanically surveyed. The flora of this islet was found to have 32 
species and one putative interspecific hybrid. This diversity represents 30 dicots and 2 monocots, in 22 
plant families. Eighteen of the plant taxa are endemic to the Guadalupe Island group, resulting in a 56.3% 
endemism in the islet’s flora. A few native plant communities were observed on the islet, within which 
only three plant taxa are obviously exotic, rare in occurrence, and presumably introduced by seabirds. 
The botanical data obtained from this undisturbed islet helps us to fill in the missing pieces about the 
overall flora of Guadalupe Island and its adjacent islets especially in relation to the ecological devastation 
caused by feral goats on the main island. Brief notes on other floristic components of Guadalupe Island 
and its islets, and new plant records collected during the expedition are also presented. 


RESUMEN 


Durante una expedicion para estudiar la historia natural de la Isla Guadalupe e islotes adyacentes, en 
junio de 2000, inventariamos la flora del islote Toro, que nunca habia sido explorado. En este islote 
encontramos 32 especies de plantas y un aparente hibrido interespecifico, incluyendo 30 dicotiledonas y 
2 monocotiledonas, de 22 familias. Dieciocho de los taxa son endémicos a la Isla Guadalupe e islotes 
adyacentes, por lo tanto Toro tiene una tasa de endemismo de 56.3%. Se encontraron algunas comunidades 
de plantas nativas, en las que solamente 3 taxa eran ex6ticas, poco abundantes, probablemente introducidas 
por aves marinas. La informacion botanica de este islote sin disturbio puede ayudarnos a llenar las piezas 
faltantes de la flora de Guadalupe, especialmente en relaci6n con la devastaci6n ecolégica causada por 
cabras en la isla principal. Ademdas, en este articulo presentamos notas breves sobre otros componentes 


floristicos de la Isla Guadalupe y sus islotes, y nuevos registros de plantas colectadas durante la expe- 


dicion. 


Key words: Guadalupe Island, Baja California, Mexican flora, expedition, endemism 


The Guadalupe Island Expedition in June of 
2000 involved 16 scientists from the U.S.A. and 
Mexico representing arachnology, botany, conser- 
vation biology, entomology, marine ecology, orni- 
thology, and phycology. This expedition was or- 
ganized by the San Diego Natural History Museum 
and funded by the Biotic Surveys and Inventories 
section of the National Science Foundation (Grant 
No. 0074462). The primary emphases of this bi- 
national, multidisciplinary endeavor were to ob- 
serve, record, and collect natural history informa- 
tion on the biodiversity of Guadalupe Island and 
the previously unexplored Toro Islet. The botanists 
of the expedition were Dr. José Luis Leon de la 
Luz of the Centro de Investigaciones Bioldgicas del 
Noroeste, Tom Oberbauer of the Department of 
Planning and Land Use for the County of San Di- 


ego, and Dr. Exequiel Ezcurra, and the senior au- 
thor from the San Diego Natural History Museum. 

The biological data obtained on this trip provides 
us with additional biodiversity information on all 
of the terrestrial areas of Guadalupe Island and its 
islets and will be used for making conservation de- 
cisions for the island and evaluating the impacts of 
introduced species such as feral goats and cats. The 
floristic information from the expedition can be 
used to supplement our current knowledge on the 
entire Guadalupe Island flora as presented by Mo- 
ran (1996). 


THE ISLAND ENVIRONMENT 


Guadalupe is an oceanic island located approxi- 
mately 260 km off the Pacific coast of the Baja 


146 


California peninsula, in northwestern Mexico. The 
island’s general geographic position is 29°03'N lat- 
itude and 118°17’W longitude. Guadalupe Island is 
about 36 km long on its N-S axis and 12 km wide 
on the E-W axis, with an approximate surface area 
of 250 km/?. The island is the peak of a seamount, 
which may have originated from several eruptive 
episodes, with the oldest exposed rocks being dated 
around 7 million years old (Moran 1996). There are 
three islets off of the southern end of the main is- 
land: Negro (30 m in elevation), Toro (220 m), and 
Zapato (190 m). Each islet has approximately 1 km? 
of surface area. 

The climate of the island is maritime and heavily 
influenced by the cold California Current, with its 
characteristic features of wind, fog, and winter rain- 
fall. A meteorological station on the southern end 
of Guadalupe (the driest area of the island) indi- 
cates almost 120 mm of rainfall annually and a 
mean monthly temperature of 17—19°C that is rel- 
atively stable throughout the year. It should be not- 
ed that the annual precipitation value given above 
does not include the contribution of moisture from 
fog condensation, which is a common event on 
Guadalupe Island. 

Floristically, the island is considered an “‘outli- 
er’ of the California Floristic Province (Moran 
1996) since it is composed of many native plant 
species either disjunct from this vegetation type on 
the mainland or on other islands, or endemic spe- 
cies with northern affinities. Although the island is 
home to Mexico’s only populations of many plant 
species from the California Floristic Province, its 
most striking attribute is its endemism, which oc- 
curs in many biological groups. According to Mo- 
ran (1996) in respect to plants, almost 22% of the 
native species are endemic, including two mono- 
specific genera. Although there have been 220 dif- 
ferent plant taxa documented over time on Guada- 
lupe Island and its islets, the activities of feral goats 
released in the early 19th century by whalers have 
devastated most of the main island’s flora. This im- 
pact on the flora is not only from the direct browse 
of goats, but also from the damage to the insular 
substrates as a result of soil erosion. At present, the 
main island’s original flora and natural plant com- 
munities have practically vanished. It is estimated 
(Moran 1996) that at least 26 native plant species 
could now be extinct, including one endemic, 
monospecific genus and many other plant species 
seem to be on the threshold. To add to the problem, 
the vegetation is now dominated mainly by weedy 
species; 62 exotic plant taxa have been recorded on 
the island, of which many are quite aggressive and 
capable of competing for niches and displacing na- 
tive species. 

Toro (also known as Islote de Adentro or Inner 
Islet) is a small islet that lies directly off of the 
southern end of Guadalupe Island, situated between 
the main island and another southern islet, Zapato. 
Toro Islet is a dome-shaped rock that reaches ap- 


MADRONO 


[Vol. 49 


proximately 500—700 feet (165-220 m) in eleva- 
tion. The islet has very steep, vertical walls that 
arise out of the water at an angle of almost 90 de- 
grees and that have, heretofore, prohibited its ex- 
ploration. However, during this expedition the sci- 
entists used a helicopter to facilitate landing and 
exploration on the islet. On its top, Toro has a small 
basin in its center with a high western ridge. A 
narrow ridge extends to the north and drops into a 
steep slope facing the main island. There are two 
types of rocky substrates that exist on this volcanic 
islet. One type of substrate is a rocky, fractured 
basaltic material with darker coloration and the oth- 
er is a tan, hard solid rock with characteristics of 
andesite. Plant species occur on both substrates but 
most seem to prefer the broken basaltic materials. 


THE FLORA OF TORO 


The known flora (see Table 1) of Toro Islet that 
was documented during the expedition consists of 
32 species and | putative hybrid. This diversity rep- 
resents 30 dicots and 2 monocots, in 22 plant fam- 
ilies. Eighteen of the taxa are endemic to Guada- 
lupe Island, resulting in a 56.3% rate of endemism 
in the islet’s flora. The Asteraceae are the best rep- 
resented on the islet with 5 genera, 6 species and 
one putative interspecific hybrid in the genus He- 
mizonia. Other families such as Malvaceae, Scrop- 
hulariaceae, and Fabaceae are represented with two 
genera. Only Cryptantha, Hemizonia, and Mesem- 
bryanthemum are represented with two species in 
the same genus. 

Only three plant taxa (Hordeum murinum ssp. 
glaucum, Mesembryanthemum crystallinum, and M. 
nodiflorum) are obviously exotic. As of yet, their 
populations on the islet are small and their occur- 
rences quite rare. It is likely that they are rather 
recent introductions, which presumably arrived by 
seabirds. In fact, Hordeum seems to be currently 
confined to the steep northern slope of the islet in 
an area near Western Gull (Larus occidentalis) 
nests. 

Of the 32 plant species found on the islet, eleven 
are succulent in nature with fleshy leaves, stems, or 
both, and are represented by species in eight dif- 
ferent plant families (Rebman 2001). Six of these 
succulent taxa are endemic to the Guadalupe Island 
and its adjacent islets. One of the leaf succulent 
species, Baeriopsis guadalupensis belongs to an en- 
demic, monotypic genus in the Sunflower family 
(Asteraceae). 

The vegetation of Toro Islet is best described as 
a maritime, succulent scrub and is similar to that 
found on the other southern islets, Zapato and Ne- 
gro. This type of vegetation probably dominated 
most of the southern portions of the main island as 
well, but is mostly extirpated now by the impacts 
of feral goats. This vegetation is dominated by Azri- 
plex barclayana, Cistanthe guadalupensis, Euphor- 
bia misera, Spergularia macrotheca var. talinum, 


2002] 


TABLE 1. 
of plant specimens made by Rebman et al. 


Plant taxon (family) 


Atriplex barclayana (Benth.) D. Dietr. (Chenopodiaceae) 
Baeriopsis guadalupensis J. T. Howell (Asteraceae) 
Castilleja fruticosa Moran (Scrophulariaceae) 


Cistanthe guadalupensis (Dudley) Carolin in Hershkovitz (Portulacaceae) 


Coreopsis gigantea (Kellogg) H. M. Hall (Asteraceae) 
Cryptantha foliosa (Greene) Greene (Boraginaceae) 


Cryptantha maritima (Greene) Greene var. maritima (Boraginaceae) 
Dichelostemma capitatum Alph. Wood ssp. capitatum (Themidaceae) 


Dudleya guadalupensis Moran (Crassulaceae) 
Erysimum moranii Rollins (Brassicaceae) 

Eschscholzia palmeri Rose (Papaveraceae) 

Euphorbia misera Benth. (Euphorbiaceae) 

Galvezia speciosa (Nutt.) A. Gray (Scrophulariaceae) 
Hemizonia greeneana Rose ssp. greeneana (Asteraceae) 
Hemizonia hybrid (Asteraceae) 

Hemizonia palmeri Rose (Asteraceae) 


Hordeum murinum L. ssp. glaucum (Steud.) Tzvelev. (Poaceae) 


Lavatera lindsayi Moran (Malvaceae) 
Lomatium insulare (Eastw.) Munz (Apiaceae) 


Lotus argophyllus (A. Gray) Greene ssp. ornithopus (Greene) Raven (Fabaceae) 


Lupinus niveus S. Watson (Fabaceae) 
Lycium californicum Nutt. (Solanaceae) 


Mammillaria blossfeldiana Boed. var. shurliana (Gates) Wiggins (Cactaceae) 


Mesembryanthemum crystallinum L. (Aizoaceae) 
Mesembryanthemum nodiflorum L. (Aizoaceae) 


Mirabilis laevis (Benth.) Curran var. crassifolia (Choisy) Spellenb. (Nyctaginaceae) 


Perityle incana A. Gray (Asteraceae) 
Phacelia floribunda Greene (Hydrophyllaceae) 
Plantago ovata Forssk. (Plantaginaceae) 


Rhus integrifolia (Nutt.) Brewer & S. Watson (Anacardiaceae) 
Spergularia macrotheca (Hornem.) Heynh. var. talinum (Greene) Jepson 


Sphaeralcea palmeri Rose (Malvaceae) 
Stephanomeria guadalupensis Brandegee (Asteraceae) 


Sphaeralcea palmeri, and Stephanomeria guada- 
lupensis. The succulent, perennial Cistanthe gua- 
dalupensis appears to be the most common of all 
of the species on the islet. It was in full flower 
during our visit in the early part of June, but other 
conspicuously flowering species included Baeriop- 
sis guadalupensis, Eschscholzia palmeri, Perityle 
incana, and two species of Hemizonia. The plant 
community of the central basin on the islet is dom- 
inated by Atriplex barclayana, Cistanthe guadalu- 
pensis, and Sphaeralcea palmeri with a strong pres- 
ence of Dudleya guadalupensis, Euphorbia misera, 
Lavatera lindsayi, Lycium californicum, Mammilla- 
ria blossfeldiana var. shurliana, Spergularia ma- 
crotheca, and Stephanomeria guadalupensis. On 
the upper slopes of the basin Baeriopsis guadalu- 
pensis, Coreopsis gigantea, Eschscholzia palmeri, 
Hemizonia greeneana ssp. greeneana, and H. pal- 
meri are prominent. The endemic Stephanomeria 
guadalupensis is most common on the lower slopes 
and bottom of the basin. The steep northern slope 
was the only location where Erysimum moranii, 
Phacelia floribunda and Rhus integrifolia were 
found. One individual plant of a putative interspe- 
cific hybrid between Hemizonia greeneana ssp. 


REBMAN ET AL.: FLORA OF TORO ISLET 147 


FLoristic List OF TORO ISLET, BAJA CALIFORNIA, MExico. Note that Coll. # refers to the collection numbers 


Coll. # 


6753, 6763 
6758, 6882 
6774 

6765 

6759 
678la 
6781b 
6773 

6757 

6772 

6755 

6779 

6761 

6756, 6766 
6775 

6760 

6770 
6754, 6764 
6780 

6769 

6778 

6884 

6784 

6767 

n/a 

Onn 7. 

6776 

6782 

6783 

6771 

6762 

6768 

6883 


greeneana and H. palmeri was found on the north- 
ern ridge. 

The flora of Toro Islet is very similar to that of 
Zapato Islet (also known as Islote de Afuera or Out- 
er Islet) with a few exceptions. It is noteworthy that 
Eriogonum zapatoense was not found on Toro, 
though an effort was made to look for it. This en- 
demic buckwheat species apparently only occurs on 
Zapato, where it is found on the upper ridge and 
basin rim and is relatively common there. Similar 
habitats were examined on Toro Islet, but it was 
not seen. Zapato Islet has a large basin that drops 
down to near sea level where plants such as Ap- 
hanisma blitoides occur. In general, Zapato appears 
to have a greater diversity of habitat types than 
Toro. Other species that occur on Zapato and not 
on Toro are Atriplex californica, Calystegia ma- 
crostegia ssp. macrostegia, Crassula connata, 
Crossosoma californica (which may no longer oc- 
cur on Zapato), Descurainea pinnata ssp. menziesii, 
Hutchinsia procumbens, Lavatera occidentalis, Oli- 
gomeris linifolia, Parietaria hespera var. hespera, 
Perityle emoryi, and Pholistoma racemosum. Plant 
taxa found on Toro Islet and not on Zapato include: 
Cryptantha maritima, Hemizonia palmeri, a puta- 


148 


tive Hemizonia hybrid, Hordeum murinum, Lupinus 
niveus, Mesembryanthemum nodiflorum, and Plan- 
tago ovata. All of these plants are found on the 
adjacent main island, which lies in closer proximity 
to Toro. 

While Toro Islet was explored quite fully, there 
are still possibilities for plants that were missed, 
particularly annual species. A botanical survey of 
Toro in early spring of a rainy year would probably 
identify additional annuals in the islet’s flora in gen- 
era such as Descurainea, Hutchinsia, Oligomeris, 
Parietaria, and Pholistoma, all of which have been 
recorded on Zapato Islet. 


MISCELLANEOUS BOTANICAL NOTES AND 
OBSERVATIONS FROM THE EXPEDITION 


In total, 149 plant collections were made during 
the entire expedition of which, four specimens doc- 
ument new distributional records for both native 
and exotic species not previously known to occur 
on the island. The new records of naturalized, ex- 
otic species include: Atriplex rosea L. (Rebman et 
al. 6817) and A. semibaccata R. Br. (Rebman 
6750), both found near the airstrip in the center of 
the main island; and Schismus barbatus (L.) Thell. 
(Rebman 6752) found in the vicinity of the North- 
east Anchorage along the trail in Barracks Canyon. 

Of particular note was the discovery of a new 
native species for the island, Lonicera hispidula 
Douglas var. vacillans A. Gray. This pink-flowered 
honeysuckle is also found on Santa Cruz, Santa 
Catalina, and San Clemente islands of California, 
but is the first record (Rebman et al. 6800) of its 
occurrence in Mexico. Only one individual plant 
was found on the upper parts of a 5-meter-high 
rocky cliff cascade. 

Moran (1996) lists the weedy exotic Chamomilla 
suaveolens (Pursh) Rydb. (=Matricaria matrica- 
rioides (Less.) Porter) as part of the Guadalupe flo- 
ra, but it should be noted that the specimen (Wig- 
gins & Ernst 113) upon which this is based was 
misidentified and is actually the native species C. 
occidentalis (E. Greene) Rydb. Another plant spe- 
cies, Dichelostemma capitatum, was listed in Mo- 
ran (1996) as possibly introduced to Guadalupe Is- 
land. However, its presence on Toro and Zapato as 
well as on major portions of the main island does 
not support the concept that it was introduced. 

In preparation for the expedition, all of the data 
from specimens previously collected on Guadalupe 
Island and its adjacent islets that are housed in the 
SD Herbarium were compiled into a database. Dig- 
itized images, mostly scanned herbarium speci- 
mens, of the endemic plant taxa for the island ar- 
chipelago were also captured. 

Botanical specimens collected on the expedition 
were done so in replicate, whenever possible, so 
that specimens would be available for study in sev- 
eral herbaria. The first collections will be deposited 
at the SD Herbarium of the San Diego Natural His- 


MADRONO 


[Vol. 49 


tory Museum, and duplicates will be in Mexican 
herbaria at HCIB in La Paz and BCMEX in Ense- 
nada. Plant materials including seeds and/or speci- 
men samples for genetic analyses were collected 
and sent to several institutions. Samples of species 
in the Asteraceae including Baeriopsis, Perityle in- 
cana, Stephanomeria, and the putative Hemizonia 
hybrid material were sent to the Jepson Herbarium 
(UC/JEPS) at the University of California Berkeley. 
Leaf material of Castilleja fruticosa was sent to the 
University of Washington (WTU) and samples of 
Lavatera lindsayi, Lupinus niveus, and Phacelia 
floribunda were sent to the University of South Da- 
kota. Triteleia guadalupensis material was sent to 
the University of Wisconsin for research on the 
Themidaceae. 

While conducting botanical explorations on Za- 
pato Islet during the expedition, some observations 
were made on the reproductive biology of two plant 
species endemic to the Guadalupe Island group. 
These observations are based only on morphology 
and their functionality still needs to be investigated 
further. It appears that Mammillaria blossfeldiana 
var. shurliana is a gynodioecious species with two 
sexual conditions. Although many of the cacti were 
not in flower at the time of our visit, a few flow- 
ering individuals showed evidence for two flower 
types occurring on different plants. Most flowering 
individuals appeared to have only bisexual flowers, 
with both functional stamens and pistils. However, 
a few plants were obviously different and seemed 
to be functionally female. These pistillate individ- 
uals had flowers with smaller tepals, abortive an- 
thers not producing pollen, and pistils with a larger 
size and increased number of stigmatic lobes. 
Plants showing these different floral characteristics 
grew in the same immediate vicinity on the islet. 
Gynodioecy is not uncommon in the Mammillaria 
species of Baja California since it has been docu- 
mented in M. dioica and some of its relatives in the 
region (Lindsay and Dawson 1952). However, this 
is the first report of this sexual condition in this 
cactus species. 

The other notable flower forms occurred in Cis- 
tanthe guadalupensis. Some individuals had flow- 
ers with “‘normal-sized’”’ petals (corolla diameter 
approximately 3—4 cm.) and yellow anthers, while 
other plants had “‘normal-sized’’ flowers but had 
pink-purple anthers. Furthermore, a fewer number 
of individual plants had significantly smaller flow- 
ers (corolla diameter approximately 2 cm.) and 
much shorter inflorescences. These plants with 
smaller flowers and inflorescences exhibited only 
yellow anthers and no fruit formation was seen. All 
of these three different flower forms were observed 
on individuals growing in close proximity to one 
another on Zapato. The comparable stages of flow- 
ering and similar moisture conditions of the local 
habitat seem to rule out any environmental cause 
for these floral variations. Therefore, it is hypoth- 
esized that Cistanthe guadalupensis is an andro- 


2002] 


dioecious species. It appears that the individuals 
that have smaller flowers, shorter inflorescences, 
and no fruit formation are functionally staminate, 
while the plants with “‘normal-sized”’ flowers and 
viable pistils that develop into fruits are hermaph- 
rodites. The variability in anther color on different 
hermaphroditic plants is not yet understood and 
may not affect the reproductive system of this spe- 
cies. 


CONCLUSION 


The botanical data obtained from Toro Islet dur- 
ing this expedition not only helps to fill in the miss- 
ing pieces about the overall flora of Guadalupe Is- 
land and its adjacent islets, but it also yields im- 
portant information for conservation efforts. Due to 
the extinction of various plant species as a result of 
the ecological devastation caused by feral goats on 
the main island, the undisturbed Toro Islet is a very 
important resource for plant taxa that can be used 
for restoration of the main island if and when the 
goats are removed. The diversity of native and en- 
demic plants in this region is a rich heritage and 
serious conservation strategies need to be imple- 
mented in order to lessen the threats for their sur- 
vival into the future. 


REBMAN ET AL.: FLORA OF TORO ISLET 149 


ACKNOWLEDGMENTS 


We are indebted to Drs. Michael Hager and Exequiel 
Ezcurra of the San Diego Natural History Museum and 
William Everett of the Endangered Species Recovery 
Council, for arranging and organizing the expedition. We 
thank the appropriate Mexican authorities for granting 
permits allowing the visitation to the island and the col- 
lection of plant species. We are grateful to the captain and 
crew of the Shogun for transportation to the island; Thom- 
as Thrailkill and Robert Gannon for field logistics; and 
Mel Cain for his remarkable piloting of the helicopter. We 
would also like to recognize Judy Gibson and botany vol- 
unteers at the SD Herbarium for their help in compiling 
a database of Guadalupe Island collections and scanning 
herbarium specimens. A special thanks to Patricia Beller 
for her efforts in obtaining the papers and permits needed 
to conduct the trip and for her help with the Spanish re- 
sumen. This material is based upon work supported by the 
National Science Foundation under Grant No. 0074462. 


LITERATURE CITED 


LINDSAY, G. AND E. Y. DAwson. 1952. Mammillarias of 
the islands off northwestern Baja California, Mexico. 
Cactus and Succulent Journal (U.S.A.) 24:77-84. 

Moran, R. V. 1996. The flora of Guadalupe Island, Mex- 
ico. Memoirs of the California Academy of Sciences 
19. California Academy of Sciences, San Francisco, 
CA. 

REBMAN, J. 2001. The succulents of Islote Toro, Baja Cal- 
ifornia, Mexico. Cactaceas y Suculentas Mexicanas 
46:52—55. 


MADRONO, Vol. 49, No. 3, pp. 150—157, 2002 


GENETIC STRUCTURE OF SENECIO LAYNEAE (COMPOSITAL): 
A RARE PLANT OF THE CHAPARRAL 


GLENDA D. MArsH! 
California State University, Sacramento, 6000 J Street, 
Sacramento, CA, 95819 


DEBRA R. AYRES 
Evolution and Ecology, One Shields Avenue, 
University of California, Davis, CA, 95616 


ABSTRACT 


The genetic structure of Senecio layneae, a rare plant endemic to the fire adapted chaparral in the 
central foothills of the Sierra Nevada, California, USA, was investigated over the entire known range of 
the species. Genetic variation was assessed using 63 ISSR and 42 RAPD polymorphic DNA markers. 
Multivariate analysis differentiated populations from each of three counties and grouped two populations 
from El Dorado County together. ISSR markers more accurately portrayed population membership patterns 
than did RAPD markers. A species-wide F,; of 0.28 (by AMOVA) and a F.; of 0.22 within El Dorado 
County showed that significant genetic differentiation exists in the species and between local (within 5.5 
km) populations. These results argue for maintenance of disturbed openings in the chapparal which this 
federally listed, threatened species can recolonise after fire, and for the use of local seed sources (within 
5.5 km) for re-introduction efforts. Additionally, the preservation of multiple populations is indicated in 
order to maintain the existing pattern of genetic diversity across the landscape. 


Key words: ISSR, RAPD, genetic structure, S. layneae, chaparral 


Senecio layneae E. L. Greene (Composital), 
Layne’s Butterweed, is a perennial herb with an 
underground rootstock that forms small clones. It 
is restricted to open rocky areas within chaparral 
plant communities on gabbroic or serpentine-de- 
rived soils in the central Sierran foothill counties of 
Yuba, El Dorado, and Tuolomne in California (Cal- 
ifornia Native Plant Society 1994; USFWS 1996). 
Plants occur in sunny openings within the chaparral 
and die off as vegetation grows up around them 
(Baad and Hanna 1987). Flowering between April 
and June, the inflorescence is composed of disk 
flowers and a handful of unevenly distributed ray 
flowers. Primarily insect pollinated (G. Marsh per- 
sonal observation), its dandelion-like seeds are suit- 
able for dispersal by the wind. It is not known if a 
persistent, heat resistant seed bank of S. layneae 
exists in the soil beneath the chaparral. 

Senecio layneae was federally listed as threat- 
ened in 1996 (USFWS 1996). Chief threats include 
residential and commercial development, road 
maintenance, decreased fire frequency, off-road ve- 
hicle use, competition from invasive alien vegeta- 
tion, shading from native tree and shrub species, 
excessive horse grazing practices and gold mining 
(USFWS 1996). Populations of between 3 and per- 
haps 1000 plants (M. Baad personal communica- 
tion) are scattered within the chaparral of western 
El Dorado County including the Pine Hill gabbroic 


' Present address: 2130 51st Street, Box 113, Sacra- 
mento, CA, 95817. E-mail: gdmarsh@juno.com. 


intrusion and adjacent serpentine (USFWS 1996). 
In Tuolumne County, several populations of be- 
tween 2 and 500 plants (A. Franklin personal com- 
munication) were first documented in 1984, and oc- 
cur in the chaparral dominated BLM Red Hills 
Management Area (BioSystems Analysis, Inc. 
1984). In Yuba County two populations, one of ap- 
proximately 200 plants on public land, and the oth- 
er composed of scattered plants found on private 
lands, are found in chaparral and open Ponderosa 
Pine forest (Bureau of Land Management 1997). 
They were first reported in 1997, expanding the 
range of S. layneae northward 93 km. 

The protection of genetic diversity within species 
has become a priority for conservation efforts (Hol- 
singer and Gottleib 1991; Falk 1992). Such genetic 
diversity may be evident between individuals with- 
in a population or between the populations in a re- 
gion. The long-term objective is to maintain the 
evolutionary viability of the taxon, and maximize 
its chances for persistence in the face of changing 
environments (Huenneke 1991). Genetic diversity 
contributes to short-term ecological success in the 
face of stochastic environmental events such as lo- 
cal population destruction, pathogens, or herbivory 
(Holsinger and Gottlieb 1991). In rare plant pre- 
serve planning, an effort to preserve the species’ 
entire genetic diversity must be a goal (Frankel and 
Soulé 1981). 

The genetic variation maintained within a species 
is distributed among regions, populations, and in- 
dividuals within populations (Barrett and Kohn 


2002] 


1991). Variation in rates of gene flow, intensity of 
natural selection, and random genetic drift create a 
hierarchy which is referred to as genetic structure. 
Genetic structure arises when gene flow between 
populations is limited, favoring development of dis- 
tinct arrays of genetic characters within each pop- 
ulation (Wright 1951; Nei 1973). It is important to 
determine the spatial scale at which differentiation 
of populations occurs in order to understand the 
pattern of population divergence and microevolu- 
tion of a species. Preservation of this pattern should 
be a priority for species conservation (Driscoll 
1998). 

Several life history factors, primarily those re- 
lated to pollen and seed dispersal, have been as- 
sociated with the development of genetic structure 
(Hamrick and Godt 1989). Wind carried seed, and 
an outcrossed breeding system, such as occurs in 
Senecio layneae (G. Marsh personal observation), 
promote gene flow, hampering the development of 
genetic structure and reducing the affects of genetic 
drift. Alternatively, Senecio layneae appears to re- 
colonize sunny openings left in the chapparal after 
disturbance such as fire or brush clearing, resulting 
in a historically patchy distribution. Small, patchily 
distributed populations would lead to the develop- 
ment of genetic structure through drift. Our primary 
research goal was to examine the genetic structure 
of this species and to infer the spatial scales at 
which gene flow is common and rare. Additionally, 
we will infer how historical fire regime could have 
influenced the genetic structure of populations seen 
today. 

Previously, much information about the genetic 
structure of plants was based almost entirely on 
data from electrophoretic surveys of soluble en- 
zymes (allozymes). Recently, RAPD-PCR-based 
methods have been used to investigate plant genetic 
Structure G-e., Huff et al. 1993: N’Goran et al. 
1994; Baruffi et al. 1995; Ayres and Ryan 1997; 
Martin et al. 1997). These nuclear genetic markers 
are presumed to be selectively neutral and can re- 
veal patterns in variation due to neutral process 
such as random genetic drift and gene flow. This 
knowledge does not necessarily inform us regard- 
ing the arrangement of adaptively important traits 
across the landscape (Olfelt et al. 2001). However, 
even with these limitations, information from mo- 
lecular markers provides insight into population 
isolation due to restricted gene flow, and diver- 
gence due to drift which are potent factors in spe- 
cies evolution. 

A new type of marker, ISSR (inter-simple se- 
quence repeat), appears to be even more efficient 
than RAPD’s (random aplified polymorphic DNA’s) 
and has been used in recent investigations of nat- 
ural plant populations (Robinson et al. 1997; Wolfe 
et al. 1998a, b; Esselman et al. 1999). ISSR markers 
are repeat-anchored primers that amplify regions 
between SSR’s (simple sequence repeats or micro- 
satellites) (Tsumura et al. 1996). ISSR primers an- 


MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 15] 


1 Yuba County 


El Dorado County 
EN Tuolumne County 


uae 


300 0 


300 Kilometers 


Fic. 1. Locations of four sample sites for Senecio lay- 
neae in Yuba County, El Dorado County, and Tuolumne 
County, California, 1999. 


neal directly to SSR’s and no prior knowledge of 
the genome is required (Tsumura et al. 1996; Wolfe 
and Liston 1998). Additionally, ISSR’s produce a 
complexity of patterns as well as a level of poly- 
morphisms detected per single PCR experiment that 
largely exceeds that of RAPD’s (Zietkiewicz et al. 
1994; Esselman et al. 1999; Ajibade et al. 2000). 
They are interpreted as dominant markers similar 
to RAPD data and are scored as binary data with 
‘band present’ or ‘band absent’ (Wolfe et al. 
1998a). Our secondary research goal was to com- 
pare the accuracy of ISSR and RAPD markers in 
determining genetic structure in S. layneae. 


MATERIALS AND METHODS 
Sample Collection 


Plant material was collected in the early spring 
when the dormant rootstocks began producing new 
growth. Four sites were sampled: Yuba County near 
Brownsville, Pine Hill and Cameron Park in El Do- 
rado County, and the Red Hills in Tuolumne Coun- 
ty (Fig. 1). One apical meristem was removed from 
20 individuals from each of four populations. Plants 
were patchily distributed and sampled individuals 
were haphazardly chosen at least 2 meters apart to 
avoid sampling the same individual twice. Each 
sample was placed on ice until returned to the lab 
at which time they were stored at —70°C until DNA 
was extracted. 


DNA Extraction 


DNA was extracted according to Ayres and Ryan 
(1997) with the addition of a RNAase treatment 
between the two alcohol precipitations: 100 pl of 
TE (10 mM Tris-HCL and 1 mM EDTA, pH 7.8) 
and | unit of RNAase (Sigma) were added to the 


152 MADRONO 


first DNA pellet and the tubes were then incubated 
at 35°C for 20 min. Precipitation in ethanol fol- 
lowed. The DNA was quantified from the absor- 
bance at 260 nm in a spectrophotometer. 


PCR and Electrophoresis 


Prior to screening primers, the optimum PCR an- 
nealing temperature was determined by running a 
temperature gradient reaction using a single acces- 
sion. The optimum annealing temperature was 
found to be 39°C for RAPD’s, and 54°C for ISSR’s. 
Amplification was performed with the following 
thermocycler programs: for RAPD’s, 94°C for 15 
sec, 39°C for 30 sec, 72°C for 2.0 min, repeated 40 
times, and then a 4°C holding temperature; for 
ISSR’s the program was identical except that the 
annealing temperature was 54°C. 

RAPD and ISSR PCR were performed in an Ep- 
pendorf Mastercycler gradient (Westbury, NY). 
DNA amplification reactions were performed in a 
volume of 15 pl containing approximately 30 ng of 
plant DNA, 0.20 pM/liter primer (Operon Tech- 
nologies, Inc., Alameda, CA, primers A4, Cl, C8, 
C10, D7, G8, G13, G14, G18, G19, H9 for RAPD); 
University of British Columbia kit 800, primers 
807, 823, 836, 846, 848, 857, 859, 860 for ISSR), 
200 wM/liter each of dATP cCTP, dGTP, dTTP 
(Promega, Madison, WI), 0.6 units Taq polymerase 
(Promega, Madison, WI), 3 wmol/liter MgCl2, 10% 
by volume MgCl2-free 10 reaction buffer A (Pro- 
mega, Madison, WI), and 10.5 wl of double dis- 
tilled water. The reaction mixture was overlain with 
a drop of mineral oil. PCR products were loaded 
on to 1.5% agarose gels for electrophoresis in 
0.50 TBE buffer, followed by staining in ethidium 
bromide, and visualized and photographed under 
UV light. Molecular sizes of the visualized bands 
were estimated by referencing a 100 base-pair lad- 
der (Gibco Co.). Polymorphic, reproducible bands 
were scored as present or absent. 

Eighty-four decanucleotide RAPD primers from 
Operon Technologies, Inc. (Alameda, CA) and 84 
ISSR primers from the University of British Co- 
lumbia were screened against one DNA sample to 
find primers that amplified DNA of S. layneae. The 
primers that produced bands were then subjected to 
a second evaluation using one accession from three 
populations (Yuba, Cameron Park, Pine Hill). Prim- 
ers that amplified DNA from all three populations 
were subjected to a third screening using three ac- 
cessions from each of the three populations (9 total 
DNA samples) to identify primers that produced 
polymorphic bands. Each of the above screenings 
was conducted two times to ensure reproducibility 
of results. Any primer which did not produce the 
same results was not used. Ultimately, 11 RAPD 
and 8 ISSR primers were selected for this study. 


[Vol. 49 


Data Analysis 
ISSR and RAPD analyses 


Of 80 samples from which DNA was extracted, 
74 accessions were ultimately included in all RAPD 
and ISSR data analyses. Five accessions with di- 
vergent band patterns were determined to be from 
a different Senecio species as at the time of sample 
tissue collection it was not possible to differentiate 
this second species of Senecio. These were dropped 
from further analyses. One accession was lost dur- 
ing DNA extraction. 

Fifty-eight RAPD and seventy-five ISSR poly- 
morphic markers were produced by PCR. Analysis 
showed that 28 markers had band patterns identical 
to other markers; these duplicates were dropped 
from further analysis. The remaining 42 RAPD and 
63 ISSR markers were used in multivariate analyses 
of inter-individual and population genetic distance. 
For comparative data analyses of each marker type, 
a sub-sample of 35 markers of each type (for a total 
of 70) were randomly selected to provide an equal 
number of markers. 


Cluster analysis 


Matrices of RAPD and ISSR phenotypes, in 
which marker presence (1) or absence (0) was re- 
corded, were analyzed for inter-individual genetic 
distance separately and together using the Euclidean 
distance coeficient of the SIMINT subprogram of 
NTSYS-pc, version 2.01d (Rohlf 1993). Resulting 
genetic distance matrices were used to construct den- 
drograms using the SAHN subprogram and employ- 
ing unweighted pair group averaging (UPGMA) in 
NTSYS-pce. 


AMOVA 


Analysis of molecular variance (AMOVA) (Ex- 
coffier et al. 1992) was used to partition the genetic 
distance (assessed using Euclidean distance) among 
individuals within a population and among popula- 
tions. This hierarchical analysis of variance parti- 
tions the total variance into covariance components 
(Schneider et al. 2000). The covariance components 
are used to compute fixation indices in terms of in- 
breeding coefficients (Schneider et al. 2000). AMO- 
VA was originally developed for RFLP haplotypes, 
but has been used for RAPD phenotypes (Huff et al. 
1993). By extension, it is also appropriate for ISSR 
phenotypes. The F,; statistic was computed and test- 
ed for significance after 1000 permutations (Schnei- 
der et al. 2000). These analyses were undertaken us- 
ing Arlequin ver. 2.0 (Schneider et al. 2000) avail- 
able at http:/Agb.unige.ch/arlequin/software/. 


Mantel test 


A Mantel test (Mantel 1967) of the correlation 
between genetic distances and geographic distanc- 
es, and the correlation between the RAPD and ISSR 
genetic distance matrices were calculated for all 


2002] 


MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 153 


RAPD and ISSR Euclidean Distance 


PH 


CP 


PH 


CP 


PH — Pine Hill 

CP — Cameron Park 
T — Tuolumne 

Y - Yuba 


El Dorado 


Tuolumne 


Yuba 


eee OT ee Oo ee ee 


2.83 3.82 


4.81 5.80 


Genetic Distance 


Fic. 2. 


UPGMA clustering of genetic distance of Senecio layneae individuals collected from Pine Hill (PH) and 


Cameron Park (CP) in El Dorado County, Tuolumne County (T), and Yuba County (Y). 


pairs of individuals using the MXCOMP program 
in NTSYS-pce (Rohlf 1993). The significance of the 
matrix was evaluated by comparing the observed 
Mantel test statistic Z, with its random distribution 
obtained after 1000 permutations. Interindividual 
geographical distances were found by calculating 
the Euclidean distance based on the UTM coordi- 
nates of each population. 


RESULTS 


Eleven RAPD primers and eight ISSR primers 
produced 42 and 63 distinct polymorphic bands, re- 
spectively. ISSR’s produced almost 2-fold more 
polymorphic bands per primer than RAPD’s (P(t = 
4.125) > 0.001). Polymorphic bands ranged in size 
from 400 bp to 1800 bp. 


Genetic Structure of Senecio Laynea 


Multivariate analysis resulted in a dendrogram 
clearly showing three population clusters with all 
individuals grouped within their geographical pop- 
ulations at the county level (Fig. 2). While the El 
Dorado county populations of Pine Hill and Cam- 
eron Park were placed in a single large cluster, most 


individuals were grouped into generally discrete 
subclusters within the main county cluster (Fig. 2). 

Highly significant (P < 0.000001) genetic dif- 
ferences between populations were detected in the 
analysis of molecular variance (AMOVA) carried 
out on the genetic distance matrix. Of the total ge- 
netic diversity, 72% was attributable to individual 
differences within a population, and 28% to diver- 
gence among populations (Fy; = 0.28) (Table 1). 
Analysis of just the two El Dorado County popu- 
lations resulted in a F,, of 0.22. 


Analysis of Differences between RAPD and ISSR 


Equalizing the number of polymorphic bands for 
each marker type and analyzing each marker type 
separately resulted in somewhat similar aggrega- 
tions of individuals except for the Tuolumne Coun- 
ty population. This population was genetically dis- 
tinct when assayed by ISSR, but was intermixed 
with the Cameron Park and Pine Hill (El Dorado 
County) individuals when analyzed with RAPD 
markers (dendrogram not presented). The number 
of individuals placed in the wrong geographical 
population was significantly greater using RAPD’s 
than, ESSR’s. (0:01 <= P(y? = 9.407),.<.0.05)...An 


154 


TABLE 1. 


MADRONO 


[Vol. 49 


AMOVA OF GENETIC DISTANCE IN SENECIO LAYNEAE. The total data set contained 74 individuals from four 


populations; El Dorado County analysis includes two populations from that county. P is the probability of obtaining a 
larger than observed variance component or fixation index by chance alone. 


Percentage 

Source of variation of variation 
Among all populations Dp 
Within populations 220i 
Among El Dorado County pop’s Dies) 
Within El Dorado County pop’s 67.29 


AMOVA based on 35 ISSR markers resulted in a 
much stronger degree of between population dif- 
ferentiation than an AMOVA based on 35 RAPD 
markers (F,; = 0.38 and 0.26, respectively). 


Mantel Tests 


The correlation between genetic distance (as- 
sessed using all markers) and geographical distance 
separating individual plants was not significant (r 
= 0.277, P < 0.49). The correlation between ge- 
netic distance matrices based on 42 RAPD markers 
and 63 ISSR markers was significant, but low (r = 
0.345, P < 0.002). 


DISCUSSION 


Significant genetic structure exists in S. layneae 
(Fsy = 0.28) resulting in three distinct geographic 
populations in El Dorado, Tuolumne, and Yuba 
Counties. Further, significant genetic structure (Fy, 
= 0.22) exists between two populations, separated 
by 5.5 km, in El Dorado County. These statistical 
findings were in agreement with multivariate clus- 
ter analysis. In contrast, in Wyethia reticulata, a 
clonal sunflower restricted to chaparral in El Do- 
rado County, populations separated by as little as 
500 m had a distinct genetic identity (Ayres and 
Ryan 1997). The lack of significant correlation be- 
tween genetic distance and geographic distance in 
S. layneae could be interpreted in two ways: either 
gene flow is occurring among all populations and 
they are not isolated from each other, or populations 
have been isolated for a long enough time for in- 
dependent divergence to have occurred through 
random processes or natural selection. Given the 
preceding evidence, which supports substantial 
population divergence, we conclude that gene flow 
is extremely rare among populations of S. layneae, 
and populations have diverged through random ge- 
netic drift or selection. 

Several characteristics that may counter the de- 
velopment of genetic structure are known for S. 
layneae. Senecio layneae is primarily outcrossing 
with insect pollination (G. Marsh unpublished data 
and personal observation), disperses its seed on the 
wind, and has a perennial life history. Outcrossing 
generally reduces genetic structure and promotes 
higher genetic variation within populations. Wide 
seed dispersal prevents divergence among popula- 


Pp Fixation indices 
<0.001 FST = 0.279 
<0.001 
<0.001 ES =0216 
<(0.001 


tions via small amounts of long-distance migration 
and promotes higher genetic variation within pop- 
ulations. Perennial life history, especially long life, 
reduces the effects of drift and increases the chanc- 
es of migration, thus hindering divergence of pop- 
ulations and loss of genetic variation. (Loveless and 
Hamrick 1984; Hamrick and Godt 1989; Linhart 
and Grant 1996; Ayres and Ryan 1999). These fac- 
tors may be responsible for the intermingling of 
individuals from the Pine Hill and Cameron Park 
populations in El Dorado County within a single 
cluster: gene flow between the two sites, 5.5 km 
apart, does occasionally occur. Small populations of 
plants, scattered between these two populations, 
may act as a genetic bridge allowing this gene flow 
to occur. However, the species dispersal abilities do 
not extend to populations more than 90 km apart. 
The Red Hills Management Area sample site is ap- 
proximately 98 km south of the Cameron Park sam- 
ple site, and the Yuba County sample site is ap- 
proximately 93 km north of the Cameron Park site. 
A comprehensive search for additional populations 
acting as genetic bridges between these three areas 
would shed more light on the patterns of genetic 
diversity in S. layneae. 

Patchy spatial distribution can increase isolation 
and reduce gene flow, enhancing differentiation 
among populations if pollinator behavior reduces 
gene flow between patches. Patchy spatial distri- 
bution could also be at work in the apparent sub- 
structuring in the Pine Hill and Cameron Park pop- 
ulations. In addition, the ease and rapidity with 
which seeds of S. layneae germinate suggest that 
no seed bank exists, and so seed bank contributions 
probably do not play a role in the maintenance of 
within population diversity. 


Utility of ISSR Markers 


In this study, ISSR primers produced significant- 
ly more polymorphic bands per primer than did 
RAPD primers. More importantly, when equal 
numbers of polymorphic bands were used in cluster 
analysis of population membership patterns, ISSR- 
based genetic distance estimates more accurately 
portrayed population assignment of S. /ayneae in- 
dividuals than RAPD-based estimates. Wolfe et al. 
(1998a) found that UPGMA dendrograms derived 
from ISSR markers showed more highly resolved 


2002] 


phylogenetic reconstructions of Penstemon popu- 
lations in the section Spectibilies than trees based 
on rDNA, cpDNA, and allozyme identity coeffi- 
cients. ISSR’s better resolved genotypes for three 
of four populations studied than did RAPD’s in ex- 
amining clonal diversity in rare Calamagrostis por- 
teri ssp. inseperata (Esselman et al. 1999). These 
results suggest a difference in the utility of RAPD 
and ISSR markers. In addition, the correlation be- 
tween RAPD-based and ISSR-based genetic dis- 
tance was low. These differences could arise due to 
differences in areas of the genome that each type 
of marker amplifies, because of higher error in 
RAPD markers, or because of higher precision in 
ISSR markers. 


Conservation Implications 


One of the main goals of conservation programs 
for species that are rare or threatened is to maintain 
existing levels of genetic variation (Avise 1994). 
While most of the genetic variation in Senecio lay- 
neae is found within populations, almost % of var- 
iation is found among populations. This is enough 
to argue for conservation of S. /Jayneae populations 
throughout its range. We also determined that pop- 
ulations separated by 5.5 km, while occasionally 
exchanging genes, are still quite distinct genetically 
(Fs; = 0.22), which argues for the use of local seed 
(within 5.5 km) for restoration projects. The nature 
of the disjunct and patchy distrubution of S. lay- 
neae populations is historical having to do with cy- 
cles of fire creating appropriate habitat within the 
chapparal. However, human activities, such as fire 
suppression and urban development, certainly influ- 
ence habitat availability in El] Dorado County ex- 
tinguishing local populations and further isolating 
remaining populations. Information about the fate 
and persistence of S. /ayneae individuals and pop- 
ulations throughout a fire cycle will be necessary 
to formulate a conservation strategy for this species 
that goes beyond management of local populations 
in each of the counties where it is currently found. 

Plants employ several mechanisms to cope with 
periodic fire. The responses to fire of three other 
rare plants found on the Pine Hill gabbro complex 
in El Dorado County demonstrate some of these 
strategies (D. Ayres unpublished data). Fire kills 
plants of the herbaceous trailing vine Calystegia 
stebbinsii but promotes seed germination from the 
soil seed bank. Plants grow rapidly and flower pro- 
fusely 2—3 yr after fire. As the canopy closes during 
the interfire period C. stebbinsii almost completely 
dies out, but populations can once again establish 
after fire as long as the soil seed bank has been 
replenished. The fire response of the low growing 
woody shrub Ceanothus roderickii is quite similar, 
except that the juvenile plants do not begin to flow- 
er until 5—6 years after fire, and some plants survive 
and flower under mature chaparral. Populations re- 
quire a fire-free period of at least 6 years to replen- 


MARSH AND AYRES: GENETIC STRUCTURE OF SENECIO LAYNEAE 155 


ish the seed bank in order to exist in perpetuity. 
Fire leaves unharmed the underground rhizomes of 
W. reticulata and promotes vigorous flowering and 
seed set. This plant remains in the understory in the 
interfire period when the canopy closes over, but 
with much reduced flowering vigor. 

While we suspect that S. Jayneae does not have 
a fire resistant seed bank due to a relatively soft 
seed coat and an absence of any seed dormancy, 
this is but one of several questions regarding the 
fire response of S. layneae that needs to be ad- 
dressed in order to formulate a management strat- 
egy for this species. Other questions concern the 
survival of the caudex during the interfire period 
and during fires, and the environmental conditions 
that promote flowering and favor seedling estab- 
lishment. These biological constraints can be used 
to answer two key management questions; how 
long should the fire interval be, and how large 
should controlled burns be? 

If the caudex of S. Jayneae can survive both the 
canopy closure of the interfire period, and fire, we 
predict it will respond similarly to W. reticulata; 
abundant flowering shoots will emerge from the 
caudex, seed production will be increased several 
fold, and seedlings will establish in the fire’s ash. 
Under this scenario, precise fire intervals and areas 
are not critical management components for popu- 
lation survival as long as some areas burn some- 
time. However, if the caudex dies out during the 
interfire period or is killed by fire, and there is no 
seed bank, S. layneae will function as a fugitive 
species. It will require open patches of chaparral, 
near an existing reproducing population, to which 
the current crop of seeds can disperse and set seed 
before fire or canopy closure occurs. This is an en- 
tirely different fire survival strategy than the three 
other species described above and is similar to that 
of Furbish’s lousewort (Pedicularis furbishiae) 
(Menges 1990). The disturbance/successional niche 
for Furbish’s lousewort is defined by a river hy- 
drology in which patches of dense shrub thickets 
and trees are removed by scouring ice flows and 
bank slumping. The lousewort recolonizes the new- 
ly opened patch from water-born seeds. In this 
model, disturbance in the form of fire or ice flows 
opens up the regeneration niche (Grubb 1977) al- 
lowing recolonization by wind or water-dispersed 
seed from surviving populations. Populations die 
out as succession progresses, so species survival 
requires Ongoing disturbance. If this is the model 
for S. layneae, survival of the species will depend 
on both fire to create the regeneration niche and a 
supply of seeds to colonize the patch, either dis- 
persing from neighboring populations, or from 
sown seed collected from nearby populations. Fur- 
thermore, the genetic structure found in S. /ayneae 
supports this ‘fugitive’ model where loss of popu- 
lations due to catastrophic fires or lack of fires frag- 
ments populations and fosters genetic drift in the 
remaining populations. 


156 


Ideally, instead of focusing on the preservation 
of each extant population of each imperiled species 
as they occur today, the requirements for continued 
existence of all imperiled species will be incorpo- 
rated into an integrated management plan. This plan 
would include fire management, control of invasive 
exotic plants, protection of preserve boundaries, 
prevention of short-interval fires, and strategies to 
ensure not only species survival but maintain pat- 
terns of genetic diversity laid down through millen- 
nia of microevolution. 


ACKNOWLEDGMENTS 


The authors thank Dr. Michael Baad, California State 
University, Sacramento; Al Franklin of the Bureau of 
Land Management; field assistant Nadia Hoshovsky; Jean- 
nette Martinez and Joyce Havsted of the Spartina Lab at 
UC Davis for their assistance, two anonymous reviewers, 
and the Goethe Foundation for funding support. 


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MADRONO, Vol. 49, No. 3, pp. 158-164, 2002 


WOOD ANATOMY AND SUCCESSIVE CAMBIA IN SIMMONDSIA 
(SIMMONDSIACEAE): EVIDENCE FOR INCLUSION IN 
CARYOPHYLLALES S.L. 


SHERWIN CARLQUIST 
Santa Barbara Botanic Garden, 1212 Mission Canyon Road, 
Santa Barbara, CA 93105, U.S.A. 


ABSTRACT 


Simmondsia chinensis (Link) Schneider, endemic to dry areas of California, Arizona, and adjacent 
Mexico, is the sole species of Simmondsiaceae. Wood anatomy and cambial activity of this species are 
analyzed in view of the transfer of the taxon from Buxaceae or Euphorbiales to Caryophyllales s.l. The 
wood contains features considered primitive in dicotyledons: tracheids, diffuse parenchyma, and rays that 
are both multiseriate and uniseriate. These features are shared with Agdestidaceae, Rhabdodendraceae, 
and Stegnospermataceae, families considered basal to most of the Caryophyllales s.s. (“‘core Caryophyl- 
lales”’). Simmondsia has nonbordered perforation plates and successive cambia, features that occur widely 
in Caryophyllales s.1. Wood anatomy of Simmondsia is congruent with placement of the genus in Cary- 
ophyllales s.l. Simmondsia wood features reflect the desert habitat both quantitatively and qualitatively. 
Terminology and ontogenetic concepts of Simmondsia secondary xylem, successive cambial action, and 
periderm are contrasted with those in a previous study, and the problems of analysis of woods with 


cambial variants are discussed. 


INTRODUCTION 


Simmondsia chinensis, contrary to its species 
name, iS native to limited areas of arid hills and 
low mountain ranges of southern California, south- 
ern Arizona and adjacent portions of Mexico 
(Munz 1973). The genus has frequently been placed 
in Euphorbiales (see Goldberg 1986, Table I, for 
the treatments of Simmondsiaceae by 11 leading 
phylogenists). Simmondsiaceae has been placed in 
the family Buxaceae within the order by many 
workers. The unisexual flowers and tricoccoid fruits 
were suggestive of this relationship for many sys- 
tematists. However, Simmondsia was unique within 
Euphorbiales in having successive cambia. Succes- 
sive cambia have evolved in several groups inde- 
pendently (e.g., Menispermaceae, Gnetales); con- 
sequently, the presence of successive cambia was 
probably not considered a feature of prime taxo- 
nomic value. 

Recent phylogenies based on molecular data 
have opened possibilities in taxonomic interpreta- 
tion with regard to Simmondsia and some other 
genera with successive cambia. These newer views 
have effectively supplanted earlier thinking, so var- 
ious earlier phylogenies are not compared here. An 
expanded Caryophyllales (“‘caryophyllids”’) was 
proposed by Williams et al. (1994). Most notably, 
Droseraceae and Nepenthaceae were added, with 
Dilleniaceae the first branch on this clade. In sub- 
sequent phylogenetic constructions (Nandi et al. 
1998; Soltis et al. 2000), families added to the ex- 
panded Caryophyllales include Ancistrocladaceae, 
Asteropeiaceae, Dioncophyllaceae, Frankeniaceae, 
and Tamaricaceae. Santalales are considered the 
outgroup for Caryophyllales s.l. In all of the phy- 


logenies utilizing molecular data, Buxaceae are not 
adjacent to Caryophyllales s.l., so the concept that 
Simmondsia belongs to Buxaceae is not supported. 
The present study is designed to compare wood and 
stem anatomy of Simmondsia to that of Caryophyl- 
lales s.1. that may be related. That purpose was also 
basic to the recent study of Rhabdodendraceae 
(Carlquist 2001a). Successive cambia occur in Car- 
yophyllales s.l. in Agdestidaceae, Amaranthaceae, 
Aizoaceae, Barbeuiaceae, Basellaceae, Caryophyl- 
laceae, Chenopodiaceae, Nyctaginaceae, Phytolac- 
caceae, Plumbaginaceae, Polygonaceae, and Steg- 
nospermataceae. The only families of Caryophyl- 
lales s.s. (‘core Caryophyllales’’) that lack succes- 
sive cambia are Cactaceae, Didieriaceae, and 
Portulacaceae. Thus, the presence of successive 
cambia in Simmondsia alone is a reason to compare 
Simmondsia to Caryophyllales. 

There have been previous accounts of Simmond- 
sia wood, notably those of Solereder (1885, re- 
ported as Brocchia), Van Tieghem (1897), Bailey 
(1980), and Carlquist (1982a). The present study 
goes beyond those studies by utilizing scanning 
electron microscopy (SEM) and by careful attention 
to ontogenetic phenomena (and associated termi- 
nology). Better understanding of successive cambia 
in Simmondsia will lead to a more accurate picture 
of successive cambia in dicotyledons and Gnetales. 


MATERIAL AND METHODS 


Stems of Simmondsia chinensis from a large 
shrub cultivated in the Santa Barbara Botanic Gar- 
den were fixed in 50% aqueous ethanol. Stems 
were taken in January and June, 2001, in order to 
compare degree of meristematic activity in the lat- 


2002] CARLQUIST 
eral meristem. The stem of S. chinensis is hard 
enough to be sectioned, without softening, on a 
sliding microtome, but thin sections contain frac- 
tures. Some sliding microtome sections were 
stained with safranin and used for permanent slides. 
Some tangential sliding microtome sections were 
dried between clean glass slides, sputter coated 
with gold, and viewed with a Bausch & Lomb Nan- 
olab SEM. Portions of “‘bark’’ (tissue exterior to 
vascular bands) were fixed in 50% aqueous ethanol, 
softened for three days at 60°C in 8% ethylene di- 
amine, infiltrated, embedded with paraffin, and sec- 
tioned according to the method of Carlquist 
(1982b). These sections were stained with a safra- 
nin-fast green combination. Macerations were pre- 
pared with Jeffrey’s Fluid (equal parts of 10% chro- 
mic acid and 10% nitric acid) and stained with saf- 
ranin. The stems studied were between one and 
three cm in diameter. 

Vessel lumen diameter rather than outside vessel 
diameter was measured; for vessels oval in transec- 
tion, long and short chords were averaged. The ves- 
sel density recorded was based upon scans that did 
not include conjunctive tissue. If conjunctive tissue 
were to be included, the number of vessels per mm? 
would be about 50% lower. Both earlywood and 
latewood were included in the computation of 
quantitative vessel data. Terms are in accordance 
with the [AWA Committee on Nomenclature (1964) 
and Carlquist (2001b). The term “‘successive cam- 
bia’ follows the usage of Schenck (1893) and 
Pfeiffer (1926). 


RESULTS 
Secondary Xylem 


Growth rings inconspicuous, but evident on the 
basis of earlywood vessel diameter (Fig. 1); early- 
wood is usually not initiated at the beginning of 
each vascular band, but at some point within vas- 
cular bands. Vessels are virtually all solitary (Fig. 
1); mean number of vessels per group, 1.04. Mean 
vessel lumen diameter, 21 fm. Mean number of 
vessels per mm’, 260 wm. Mean vessel element 
length, 163 wm. Mean vessel wall thickness, 2.8 
wm. Perforation plates simple, nonbordered (Fig. 3, 
top). Helical thickenings present on vessel walls, 
some in the form of pairs of thickenings parallelling 
helices of pits (Fig. 3). Pit cavities of lateral wall 
vessel pits about 3 ym in diameter. Imperforate tra- 
cheary elements all tracheids, densely covered with 
fully bordered circular pits about 3 4m in diameter 
(Fig. 4). Mean tracheid wall thickness, 3.2 wm. Ax- 
ial parenchyma sparse, diffuse, composed of cells 
that are not subdivided. Rays uniseriate to wide 
multiseriate (Fig. 2); uniseriate rays are more com- 
mon than multiseriate rays. Most uniseriate rays are 
a single cell in height (and thus not easily seen in 
Fig. 2). Mean height of multiseriate rays, 212 pm. 
Mean width of uniseriate rays, 74 sm. Mean ray 
cell wall thickness, 1.5 im, walls lignified. Ray cell 


: SIMMONDSIA 


159 


wall pits simple. Ray cells predominantly procum- 
bent; square and upright cells relatively uncommon. 
Starch abundant in ray cells (Fig. 5). Secondary 
xylem nonstoried. 


Conjunctive Tissue and Cambial Action 


Conjunctive tissue is composed of bands five to 
ten cells thick radially (Fig. 1: tangential bands of 
thin-walled radial cells, containing phloem strands, 
intervening between tangential bands of dark sec- 
ondary xylem). As seen in radial section (Fig. 6), 
cells are mostly upright, a few square (and thus 
contrast with ray cells of the secondary xylem). 
Cell walls of conjunctive tissue are lignified and 
about 1.5 wm in thickness, but thin-walled nonlig- 
nified cells present on bark side of phloem strands 
(Fig. 7, above crushed phloem). Functional phloem 
cells, adaxial to the dense crushed phloem strand, 
are somewhat compressed due to sectioning rather 
than polygonal. What portion of the phloem cells 
that are not crushed are functional could not be de- 
termined. Continued production of secondary phlo- 
em is possible because earlier-formed phloem is 
progressively crushed. Solitary rhomboidal crystals 
are occasional in conjunctive tissue (Figs. 6, 8). 
Periclinal divisions can be seen in young conjunc- 
tive tissue (Fig. 9), but number of cell layers in the 
meristematic zone of the lateral meristem between 
the most recently initiated vascular cambium and 
the previous vascular band varies (e.g., Fig. 8). 

Vascular cambia (pointers at right edge, Figs. 
7-9) actively produce secondary phloem and sec- 
ondary xylem: the secondary phloem occurs as iso- 
lated strands, whereas the secondary xylem mostly 
occurs as unbroken cylinders that extend around the 
stem. The first tracheids produced by a cambium 
are polygonal in outline, and their alignment in ra- 
dial rows is sometimes obscure because during 
maturation, the interfaces of the several faces shift. 
As divisions wane in each vascular cambium, a few 
thin-walled tracheids in radial rows are produced 
(Fig. 7, below phloem). Cambial activity produces 
phloem for a prolonged period, probably for several 
years because the abaxial portion of secondary 
phloem strands consist of numerous crushed phlo- 
em cells (Fig. 7, gray mass in center of photograph) 
and phloem cells produced by the cambium are in 
radial rows. Also, crushed phloem cells are always 
abaxial to apparently functional secondary phloem 
cells (Figs. 1, 7). 


Phellogen and Lateral Meristem Action 


The outer layers of the relatively mature stems 
studied here lack the cortical fiber strands reported 
by Bailey (1980) in the relatively young stems he 
studied. Such fiber strands had been shed from the 
stem I studied. Phellem, which consists of cells 
filled with dark-colored contents, is present on the 
stem surface. The phellem cells are narrower than 
cells in what is here termed diffuse lateral meristem 


160 MADRONO [Vol. 49 


ri DM 


Fics. 1-5. Stem sections of Simmondsia chinensis. 1. Transection (abaxial side above), showing alternation of bands 
of secondary xylem (each with associated phloem strands) and conjunctive tissue; vascular bands are not annual in 
extent; the pointers demarcate part of the earlywood of one year (above pointers) from part of the latewood of the 
preceding year (below pointers), several vascular bands are produced per year. 2. Tangential section of secondary 
xylem; a few large multiseriate rays are apparent (center), uniseriate rays are mostly inconspicuous because of their 
very small size. 3-5. SEM photographs from a tangential section of secondary xylem. 3. Portion of inner surface of a 
vessel, showing nonbordered perforation plate (top) and helical thickenings. 4. Outer surface of a tracheid, showing 
bordered nature and density of pits. 5. Starch grains in ray cells. Figs. 1, 2, magnification scale above Fig. 1 (divisions 
= 10 um; Figs. 3—5, scale bar in each figure = 5 wm). 


2002] CARLQUIST: SIMMONDSIA 161 


Fics. 6-9. Sections of stem of Simmondsia chinensis. 6. Radial section of conjunctive tissue to show cell shape and 
(center) rhomboidal crystal (secondary xylem at extreme left, secondary phloem at extreme right). 7. Strand of secondary 
phloem and surrounding tissues (conjunctive tissue above, secondary xylem below); crushed secondary phloem in abaxial 
part of phloem strand (pointer indicates site where vascular cambium was prior to cessation of active division. 8, 9. 
Transections of lateral meristem zone at periphery of stem; pointers at left in each indicate offset between outermost cells 
of the radial lateral meristem (radial files) and the innermost periderm cells; pointers at right indicate vascular cambium 
location. 8. Lateral meristem zone in which a vascular cambium has recently formed (no secondary xylem or phloem 
elements identifiable yet); rhomboidal crystal at bottom center. 9. Lateral meristem zone in which vascular cambium has 
yielded a vessel (extreme left) and some tracheids as well as some secondary phloem (right). Diagonal arrows denote 
recent divisions in the radial files of the lateral meristem zone; horizontal arrow (lower right) denotes a cell plate (obscured 
by cell contents adherent to it) that indicates a recent division in conjunctive tissue that is still somewhat meristematic. 
Figs. 6, 7, magnification scale above Fig. 6 (divisions = 10 pm). Figs. 8, 9, scale above Fig. 8 (divisions = 10 wm). 


162 


(it could also conceivably be termed secondary pa- 
renchyma). The diffuse lateral meristem cells are in 
radial rows; the term “‘diffuse’’ is used because di- 
visions do not occur synchronously in a single layer 
as in a vascular cambium, but are randomly dis- 
tributed throughout the meristematic zone (see 
Carlquist 1999a). A lateral meristem that consists 
of a single layer was observed in Barbeuia (Carl- 
quist 1999b) and also occurs in Nyctaginaceae 
(Carlquist unpublished data; data in earlier papers 
vary in interpretation and will be discussed in a 
later paper). More numerous divisions were ob- 
served in the stems collected in June than in those 
collected in January. The radial rows of lateral mer- 
istem cells are offset from the periderm which con- 
sists of a single layer of phellogen (narrow, like the 
phellem cells, but devoid of dark-colored com- 
pounds). In some places, there is a layer of paren- 
chyma between the phellogen and the lateral mer- 
istem files. No phelloderm cells were identified un- 
equivocally. The offset between the radial rows of 
the diffuse lateral meristem and the periderm is in- 
dicated by a pointer at the left in Fig. 8 and Fig. 9. 
The entirety of the periderm is illustrated in Figs. 
8 and 9, which were selected to show primarily 
lateral meristem and vascular cambia. 

Origin of vascular cambia occurs within the ra- 
dial files of cells produced by the diffuse lateral 
meristem. Although only a small portion (for rea- 
sons of clarity) could be illustrated, study of the 
entirety of sections validates this interpretation. The 
origin of a vascular cambium (Fig. 8, pointer at 
right) can be distinguished from divisions of the 
lateral meristem because divisions of the vascular 
cambium form a single meristematic layer of divi- 
sions that are synchronous in tangential bands 
around the stem. Each vascular cambium soon pro- 
duces secondary xylem internally (adaxially) and 
secondary phloem abaxially (Fig. 9, pointer at left; 
vessels and a few tracheids in secondary xylem). 
The terminal products of a vascular cambium (Fig. 
7) are described above. 


CONCLUSIONS 
Phylogenetic Position 


The occurrence of successive cambia is a char- 
acter widespread in Caryophyllales s.s. (“core Car- 
yophyllales’’) so its occurrence in families now 
added to an expanded Caryophyllales—Rhabdoden- 
draceae (Carlquist 2001a) and Simmondsiaceae is 
not surprising. ““Caryophyllales: s.s.’’ corresponds 
to the betalain-containing families plus Achatocar- 
paceae, Barbeuiaceae, and Molluginaceae, and the 
genera Limeum and Lophiocarpus (Clement et al. 
1994). Simmondsia has characters generally consid- 
ered primitive in dicotyledons: presence of tra- 
cheids, presence of diffuse axial parenchyma, and 
presence of both multiseriate and uniseriate rays 
(Metcalfe and Chalk 1950: xlv, “fibres with dis- 
tinctly bordered pits; Kribs 1935, 1937). All of 


MADRONO 


[Vol. 49 


these features are present in Rhabdodendraceae 
(Carlquist 2001a), now placed at the base of Car- 
yophyllales s.l., and in genera now placed at or near 
the base of Caryophyllales s.l. (Soltis et al. 2000). 
Simmondsia is placed by Hoot et al. (1999) and 
Soltis et al. (2000) near the base of Caryophyllales 
s.s. The other genera with the primitive features 
listed above include Agdestis (Carlquist 1999c) and 
Stegnosperma (Carlquist 1999a); Barbeuia has tra- 
cheids, but not the other character states mentioned 
above (Carlquist 1999b). 

One feature of possible ordinal significance is the 
presence of nonbordered perforation plates. These 
have been demonstrated in most Caryophyllales s.s. 
(see Carlquist 1999a, b, 2000). Nonbordered per- 
foration plates are newly reported here for Sim- 
mondsia (Fig. 3), and have recently been reported 
for some Caryophyllales s.l. such as Rhabdoden- 
draceae (Carlquist 2001a). Nonbordered perforation 
plates may be a symplesiomorphy in Caryophylla- 
les s.l. according to the above data and other ob- 
servations (Carlquist 2001b). 


Ecology 


Simmondsia is a desert shrub with only slight 
succulence in the leaves (Bailey 1980); not surpris- 
ingly, it has xeromorphic wood. The Mesomorphy 
Ratio (vessel diameter times vessel element length 
divided by vessel diameter) was reported to be 27.8 
for Simmondsia by Carlquist and Hoekman (1985). 
A very similar value (24.4) can be derived from the 
present data if conjunctive tissue is not excluded. 
The similarity of the two reports is even closer if 
one notes that outside vessel diameter, rather than 
lumen diameter was used by Carlquist and Hoek- 
man (1985). 

Tracheids are conductively safe (excellent at con- 
fining embolisms to a single cell) compared to ves- 
sel elements. Fiber-tracheids and libriform fibers, 
by contrast, are nonconductive (see discussion in 
Carlquist 2001b). The presence of tracheids in Sim- 
mondsia is a feature of value in a xeromorphic hab- 
itat. Because of the presence of tracheids, vessel 
grouping in Simmondsia is virtually nil (1.04), in 
agreement with the correlation for dicotyledons as 
a whole claimed by Carlquist (1984). The value of 
tracheids in promoting conductive safety exceeds 
the value of vessel grouping (Carlquist 2001b). 


Ontogeny and Terminology 


The terminology in papers and books that deal 
with successive cambia is remarkably diverse, but 
more significantly, different interpretations often 
underlie the terms used. The present paper is not a 
proper venue for a review of this situation. How- 
ever, the paper by Bailey (1980) on Simmondsia is 
appropriate for comparison in view of the ontoge- 
netic interpretations as well as mature structures de- 
tailed in both the present paper and Bailey’s. 

In the present interpretation, a diffuse lateral 


2002] 


meristem forms outside of the vascular cylinder, 
near the stem periphery. This lateral meristem pro- 
duces radial files of cells, producing parenchyma 
cells with primary walls, cells which remain rela- 
tively meristematic judging from recent divisions to 
be found in this region. Within the lateral meristem 
zone, a new vascular cambium is formed (usually 
while the preceding vascular cambium is still ac- 
tively producing secondary xylem and phloem). In 
Bailey’s (1980) interpretation, the zone I have 
termed lateral meristem is called conjunctive tissue 
(despite its lack of lignified secondary walls as 
found in conjunctive tissue in older parts of the 
stem). Bailey (1980) uses the term “‘extrafascicular 
cambium”’ for what I term the vascular cambium 
in each of the concentric vascular bands. 

Bailey (1980) claims that “‘the phellogen is ac- 
tually a region of transition where the peripheral 
conjunctive parenchyma of previous extrafascicular 
cambia undergoes further cellular subdivision; a 
true phellogen is lacking.” In the relatively young 
stems illustrated by Bailey (1980), phellogen might 
well be formed from cortical parenchyma as it is in 
many dicotyledons, but my studies indicate the ex- 
istence of a self-perpetuating phellogen, as so fre- 
quently described in dicotyledons. This phellogen 
is distinguished from the lateral meristem (outer- 
most conjunctive parenchyma of Bailey) not only 
by its tangentially narrower cell diameter but by an 
offset between the periderm files and the files of 
cells in the radial parenchyma (Figs. 8, 9). These 
two differences would be difficult to explain if peri- 
derm were ontogenetically continuous with the files 
of cells of the lateral meristem. 

The vascular cambia produce strands of second- 
ary phloem externally and cylinders of secondary 
xylem internally. In my interpretation, quite ordi- 
nary rays are produced by each cambium. In Bai- 
ley’s (1980) interpretation, “‘conjunctive tissue ini- 
tials produce raylike structures of conjunctive tis- 
sue; true vascular rays are absent.’’ This interpre- 
tation has not, to the best of my knowledge, been 
offered in any genera with successive cambia other 
than in Bailey’s study of Simmondsia. 

The differences in interpretation detailed above 
show that careful analyses of successive cambia 
and other cambial variants still need to be under- 
taken. The diversity of interpretations and terms for 
the anatomical phenomena is still considerable. The 
number of different interpretations and terms within 
instances of successive cambium occurrence seems 
unlikely to be matched by an equal diversity of on- 
togenetic mechanisms. Rather, cellular arrange- 
ments have been viewed differently by different 
workers. In part, the diversity of interpretations and 
terminology may derive from microtechnical con- 
siderations. Thin sections such as can be cut with 
a rotary microtome, are desirable for revealing cell 
lineages and histological details clearly. The hard- 
ness of many stems with successive cambia has, on 
the contrary, led to preparation of sliding micro- 


CARLQUIST: SIMMONDSIA 


163 


tome sections in which soft tissues do not section 
well or are too thick for cell development sequenc- 
es to be revealed clearly. The use of rotary micro- 
tome sections of material that has been chemically 
softened to a suitable degree seems the best solu- 
tion to this dilemma. Embedding in resin or plastic 
is an alternative microtechnical possibility. 


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. 1982b. The use of ethylene diamine in softening 

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. 1984. Vessel grouping in dicotyledon woods: sig- 

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. 1999a. Wood and stem anatomy of Stegnosperma 

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. 1999b. Wood anatomy, stem anatomy, and cam- 

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440. 

. 1999c. Wood anatomy of Agdestis (Caryophyl- 

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. 2001a. Wood and stem anatomy of Rhabdoden- 

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. 2001b. Comparative wood anatomy, 2nd ed. 

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AND D. A. HOEKMAN. 1985. Ecological wood anat- 
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CLEMENT, J. S., T. J. MABRY, H. WYLER, AND A. S. DREID- 
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Kriss, D. A. 1935. Salient lines of specialization in the 
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164 MADRONO 


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METCALFE, C. R. AND L. CHALK. 1950. Anatomy of the 
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MAbRONO, Vol. 49, No. 3, pp. 165-168, 2002 


A NEW SUBSPECIES OF NAVARRETIA LEUCOCEPHALA 
(POLEMONIACEAE) FROM VERNAL POOLS IN EASTERN WASHINGTON 


CURTIS R. BJORK 
Marion Ownbey Herbarium, Washington State University, Pullman, WA 99163 
cbjork @ wsu.edu 


ABSTRACT 


Navarretia leucocephala Benth. subsp. diffusa is newly described from numerous populations in a 
distinctive vernal pool landscape in the Channeled Scablands of Eastern Washington. Its affinities appear 
closer to subspecies of N. leucocephala in the California floristic province, than subspecies minima, which 
is the only other member of the N. leucocephala group known to grow within the range of the Flora of 
the Pacific Northwest (Hitchcock and Cronquist 1973). The range of subspecies diffusa closely corre- 
sponds with the central of three channels of the glacial Spokane Flood events and is segregated from 
populations of subspecies minima by expansive Palouse loess uplands, where vernal pools are absent. 
Just as no plants of subspecies diffusa were found outside this central flood channel, no plants of sub- 
species minima were found within the range of subsp. diffusa. Plants of N. leucocephala subsp. diffusa 
are easily distinguished from subspecies minima in Washington by the fewer flowers within the much 
more openly diffuse heads, and by the conspicuously puberulent calices, fewer seeds per capsule, longer 
corolla lobes and higher filament insertion. It is distinguished from other subspecies by the consistently 


puberulent calices, low seed number and the relatively simple outer inflorescence bracts. 


Key words: Navarretia, Polemoniaceae, vernal pools, Columbia Plateau, Channeled Scablands 


The Navarretia leucocephala Benth. complex in- 
cludes four species in North America: N. fossalis 
Moran, N. leucocephala Bentham, N. myersii PS. 
Allen & A. G. Day (two subspecies) and N. pros- 
trata (A. Gray) E. Greene, and one in South Amer- 
ica: N. involucrata Ruiz & Pavé6n (Day 1993a). Na- 
varretia leucocephala is defined as having five sub- 
species: leucocephala, bakeri (H. Mason) A. G. 
Day, minima (Nuttall) A. G. Day, pauciflora (H. 
Mason) A. G. Day, and plieantha (H. Mason) A. 
G. Day (Day 1993a). Two of the subspecies are 
widespread; subspecies /eucocephala occurs in the 
Great Valley of California and in southwestern 
Oregon (Day 1993b), and subspecies minima is 
found from California to north-central Washington, 
western Idaho and to Utah. During a study of vernal 
pools of the Columbia Plateau, populations of Na- 
varretia leucocephala encountered in one subre- 
gion of the Columbia Plateau vernal pool province 
stood out as being highly distinctive, leading to the 
morphological comparison with the other subspe- 
cies of N. leucocephala presented here. 


TAXONOMY 


Navarretia leucocephala Benth. subsp. diffusa 
Bjork subsp. nov.—TYPE: USA, Washington 
State, Lincoln County, Swanson Lakes Wildlife 
Management Area. Floors of vernal pools on 
scabland basalt flows. 680 m. T24N R34E S4 
NEA/4. June 18, 1997 Curtis R. Bjérk 3229 (Ho- 
lotype, WS, Isotype WTU). 


Planta annua, erecta vel decumbens, 0.5—4.7 cm 
lata; caulis centralis 1.0—13.4 cm alta, ad medium 


0.3—0.7 mm crassus; capitula hemispherica, (1) 4— 
10 (15) mm lata, diffusa, plerumque ramificans, flo- 
ribus (1) 5—20 (30); bracteae exteriorae lobis (0) 2— 
6 (8); calyx pilis brevis crispis plerumque recur- 
vatis, lobi plerumque integri; corolla inclusa, alba, 
lobi lineares; stamina in sinibus lobarum corollum 
inserta, inclusa, stigma superantibus; semina (1) 2 
G6): 

Plants annual, erect to decumbent, 0.5—4.7 cm 
wide; central stem 1.0—13.4 cm high, 0.6 + 0.1 mm 
thick at midlength; heads hemispheric, (1) 4—10 
(15) mm wide, diffuse and generally branching, 
with (1) 5—20 (30) flowers; outer bract lobes (0) 2— 
6 (8); calyx conspicuously puberulent with crisped, 
mostly recurved hairs, lobes mostly entire; corolla 
included, white, lobes linear; stamens included, in- 
serted near corolla-lobe sinuses, surpassing the stig- 
ma; seeds (1) 2 (3). 


Paratypes. U.S.A. Washington, Lincoln County: 
adjacent to Knack Road, 3.5 miles SSW of Telford, 
T25N R35E S29 NE % of NW %, Bjork 3228 (WS), 
18 June 1997; Swanson Lakes Wildlife Manage- 
ment Area, 1 mile West of Florence Lake, June, 
1999, Bjork s.n. (WS); Swanson Lakes Wildlife 
Management Area, 2 miles East of refuge head- 
quarters, T25N R34E S35 NE %, 19 June 1997, 
Bjork 3250 (WS); 3.4 miles SSE of Swanson 
Lakes, along Seven Springs Dairy Road, 13 June, 
1998, Mark Fishbein 3439 (WS); Large vernal pool 
along Swanson Schoolhouse Road, T25N R34E 
S31 NW % of NW %, Bjork 6161 (WS). 


Comparative morphology. Plants of this distinc- 
tive group of populations are here named Navar- 
retia leucocephala subspecies diffusa and are clear- 


166 MADRONO 


ly distinguished from all other subspecies of N. leu- 
cocephala by their consistently and conspicuously 
puberulent calyx, their generally much more diffuse 
heads and their relatively simple outer bracts (Table 
1). It is placed within N. leucocephala rather than 
with any other species or in its own specific rank 
due to the similarity of plant architecture and the 
cymose inflorescence to those of subspecies N. leu- 
cocephala (in contrast especially to N. myersii and 
N. prostrata) (Crampton 1954), and due to the low 
seed number (in contrast with N. fossalis). Addi- 
tionally, it differs from the South American species 
N. involucrata by flower color and the simpler 
leaves and bracts. The subspecies of N. leucoce- 
phala with the greatest morphological similarity to 
subsp. diffusa appears to be subsp. pauciflora of 
Northern California. Both have few seeds per cap- 
sule (usually 2 in both subspecies), few flowers per 
head, high filament insertion and corollas with nar- 
row throats and linear lobes. Specimens of subspe- 
cies diffusa are distinguished from those of subspe- 
cies pauciflora by the often broader (to 15 mm, 
though still few-flowered) and more diffuse heads, 
the consistently white flowers (versus the generally 
bluish flowers of subspecies pauciflora), the sim- 
pler outer bracts and the taller stature (central stem 
length to 10 cm). Also similar to subspecies diffusa 
is subspecies bakeri of Northern California and 
Southwestern Oregon. The fewer flowers per head 
and fewer seeds per capsule distinguish subsp. dif- 
fusa from subsp. bakeri. Subspecies diffusa differs 
from subsp. leucocephala in the included corollas 
and the higher filament insertion. Subspecies dif- 
fusa is distinguished from subsp. plieantha by the 
white versus bluish flowers, fewer flowers per head 
and by the lower seed number. 

Subspecies diffusa differs from subsp. minima by 
the simpler bracts, fewer flowers per head, lower 
seed number, lesser central stem length and width, 
narrower head width, higher filament insertion, ca- 
lyx length and the fewer calyx lobe divisions (Table 
2). Plants of subspecies diffusa are also distin- 
guished from subsp. minima by the nearly simul- 
taneous maturation of flowers within a head. Unlike 
in subsp. diffusa, there are typically some flowers 
in subsp. minima heads that bear mature seeds 
while others are not yet in anthesis. The heads of 
subsp. diffusa are openly branching, so much so 
that the calyx bases are often clearly seen within. 
The calyx lobes of subsp. diffusa are unequal and 
longer in relation to the tube, ranging from 0.8—1.5 
x the tube length, in contrast to those of subsp. 
minima, which are usually subequal in length and 
0.5-1.0 X the tube length. Additionally, one to 
three of the ribs leading to the lobes in subsp. dif- 
fusa maintain their width and herbaceous tissue to 
the base of the calyx tube. The ribs are often wider 
than the intervening membranes, which is appar- 
ently unique within section Navarretia (Day 1993b) 
and the N. leucocephala complex. The calyx tube 
in subsp. diffusa is conspicuously puberulent 


MORPHOLOGICAL COMPARISONS OF THE SUBSPECIES OF NAVARRETIA LEUCOCEPHALA AND N. L. SUBSP. DIFFUSA SuBSP. Nov. Outlier values are shown in parentheses. 


Data are in part from Day 1993b, with additional observations from herbarium specimens (WS, OSC). 


TABLE 1. 


diffusa 


plieantha pauciflora 


bakeri 
2-10 
13-24 
30—60 


minima 


2-11 


(4) 7-16 (20) 


leucocephala 


1-16 (10) 
(1) 4-10 (15) 


(1) 5-20 (30) 


(O) 2-6 (8) 


1-3 
15-20 
20-80 


2-22 
9-36 
15—80 


Central stem length (cm) 
Head width (mm) 
Flowers per head 
Outer bract lobes 


6-12 
6-30 
4-12 
0-10 
1.7-3.5X 


0.8-3.0 
gen. + equal 


none 


20-80 


6-10 
4-8 


9-16 
8—30 
1.2—-2.5X 
1.0-2.3X 


4—10 


(O) 4—16 (30) 


A-14 
(0) 4-19 


0-6 
1.0—2.2x 


1.5—3.0X 
gen. + equal 
gen. none 
dense 
white 


Bract lobe 2° divisions 


1.0—2.0x 


1.5-3.0X 
1.0—2.0X 
gen. unequal 


(1) 2-3 


1.5-3.0X 
0.6-3.0X 
gen. + equal 


gen. none 
gen. sparse 


white 


Bract length/head width ratio 


Calyx tube/lobe length ratio 


gen. + equal 


none 


Calyx lobe relative lengths 
Calyx lobes divided 

Calyx pubescence 

Corolla color 


gen. sparse 
gen. bluish 
+ linear 
incl. 


gen. sparse 
gen. bluish 
+ linear 
incl. 


gen. sparse 
gen. white 
+ jinear 
incl. 


gen. sparse 


white 


+ linear 
incl. 


+ oblong 


incl. 


+ ovate 
excl. 


Corolla lobe shape 


Corollas included/exserted 


Filament insertion 
Seed number 


at sinuses at sinuses 


in throat at sinuses at sinuses 


in throat 


1—2 (3) 


(2) 3-5 (6) 2—4 2-3 


AF) 


[Vol. 49 


2002] BJORK: 
wa | 
A) yy awh WGA on. 
\\ > i Xi OK: A 
es i pe 
a. AG og 
Wyo 7 4 
Ad) WY 
Sls 
yim 
Bia. 1. 


NAVARRETIA LEUCOCEPHALA SUBSP. DIFFUSA 


167 


<V), i We l 
Ni 
y 


Illustration of Navarretia leucocephala subsp. diffusa. A. Mature plant in anthesis. B. View of a seed, showing 


slightly roughened surface. C. Side view of a calyx. D. Corolla, opened from the side and showing pistil and relative 
positions of anthers and stigma. Note also the single veins entering the corolla lobes. 


throughout with crisped, usually recurved hairs. 
The mouth of the calyx is less strongly pubescent 
in subsp. diffusa, despite the external pubescence, 
than in subsp. minima, and the hairs are shorter and 
thinner in width. The corolla lobes of subsp. diffusa 
are narrower and less rounded at the apex than in 
subsp. minima. Subsp. diffusa has simpler leaves, 
often lacking lobes. The stems of subsp. diffusa are 
typically reddish, which is not usually the case with 
subsp. minima, and the stem pubescence differs 
from subsp. minima in being sparser, more crisped 
and less strongly recurved. Additionally, subsp. dif- 
fusa stems often bear scattered patches of glandular 
hairs. These differences are consistent throughout 
the ranges of both subspecies, and no obvious in- 
termediates were found. 


TABLE 2. 


Ecology and distribution. The range of subsp. 
diffusa closely corresponds to a broad, basin-like 
expanse of basalt flows surrounded by loess hills to 
the east and west, and sand and gravel deposits to 
the south. This basin was formed by massive floods 
during the last ice age. The repeated sudden failure 
of an ice-dam in Montana released enormous 
amounts of water onto the Columbia Plateau, scour- 
ing away the thick loess deposits and exposing the 
basalt bedrock over large areas (Bretz 1969). The 
flood waters gouged out three main channels, and 
it is the central of the three, in an area of approx- 
imately 550 km? where subsp. diffusa is found to 
the exclusion of subsp. minima. Other vernal pool 
landscapes occupy the eastern and western of the 
three flood channels. There, subsp. minima grows 


QUANTITATIVE MORPHOLOGICAL COMPARISONS BETWEEN SUBSP. MINIMA AND SUBSP. DIFFUSA. Means are given 


with one standard error and were tested on the null hypothesis of no difference in one-way ANOVAs. Measurements 
were taken from 50 plants of subsp. minima collected from 7 locations in Spokane, Adams, Klickitat and southwest 
Lincoln counties, and 60 plants of subsp. diffusa collected from 5 vernal pools in two locations in central Lincoln 
County. Asterisks indicate statistical significance (** p < 0.01; *** p < 0.001). 


Subsp. minima Subsp. diffusa 


Plant width (cm) 4.6 + 0.6 35 +04 
Central stem length (cm) 6-5) 5 0:4 40 + 0.3% 
Central stem width (mm) 0.8 + 0.1 0.6 + 0.1*** 
Heads per plant (oe ems wf 49 + 0.6 
Head width 2-2 O:3 7.0 + 0.3*+* 
Flowers per head 23-5 3 A:6 10.1 + 0.8++* 
Outer bract divisions 14 = OA 4.5 + 0.2*** 
Calyx length 55.2001 4.8 + 0.1*# 


168 MADRONO 


in the absence of subsp. diffusa. Thus, the two sub- 
species of Navarretia leucocephala in Washington 
are geographically isolated from each other by the 
loess hills, possibly leading to genetic isolation. 

Navarretia leucocephala subsp. diffusa is found 
exclusively in vernal pool basins like all the mem- 
bers of the N. leucocephala group (Spencer 1997). 
It is submerged at germination (in early to mid 
spring on the Columbia Plateau) and it grows and 
flowers when emergent (late April to early July dur- 
ing normal years). Subspecies diffusa is abundant 
in many hundreds of vernal pools, from those as 
small as 1 m* and no deeper than | dm to those as 
wide as 50,000 m? occupying basins as deep as 1 
m. The plants were never found along intermittent 
streams or in any other wetland type. The greatest 
density of individuals is usually where there is little 
competition from taller plants, though high densi- 
ties are sometimes found under dense canopies of 
Deschampsia danthonioides (L.) Beauv. on pool 
margins. Common associates are Polygonum poly- 
galoides subspp., Psilocarphus spp., Bousiduvalia 
spp., Alopecurus saccatus Vasey and Downingia 
yina Applegate. Surrounding plant communities are 
complex and highly diverse mosaics of numerous 
shrub, forb and graminoid codominants on lithosol, 
mima mounds and wetland soils. 

No major ecological differences were apparent 
between the sites in which subsp. diffusa and subsp. 
minima occurred in Washington. However, in ca- 
sual observations, a difference was noted in 
drought response among the taxa that may be sig- 
nificant. Despite the similarly dry and hot condi- 
tions of the ranges of subspecies diffusa and mini- 
ma, plants of subsp. diffusa were more able to with- 
stand the drought and produce seed than plants of 


[Vol. 49 


subsp. minima. A large proportion of the seedlings 
of subsp. minima died during an early three-week 
drought and high temperatures in April, 1998, 
while very few drought-killed individuals of subsp. 
diffusa were seen until late June of that year. 


ACKNOWLEDGMENTS 


Thanks are due to The Nature Conservancy of Wash- 
ington and the Swanson Lakes Wildlife Management Area 
for making the discovery of this taxon possible, Stan 
Spencer, Alva Day, Peter Dunwiddie and Mark Fishbein 
for their helpful comments. 


LITERATURE CITED 


Bretz, J. H. 1969. The Lake Missoula floods and the 
Channeled Scabland of Washington, new data and in- 
terpretations. Journal of Geology 77:503—543. 

CRAMPTON, B. 1954. Morphological and ecological con- 
siderations in the classification of Navarretia (Pole- 
moniaceae). Madrono 12:225—238. 

Day, A. G. 1993a. New taxa and nomenclatural changes 
in Allophyllum, Gilia, and Navarretia (Polemoni- 
aceae). Novon 3:331—340. 

1993b. Navarretia. Pp. 844-849 in James C. 

Hickman (ed.), The Jepson manual: higher plants of 

California. University of California Press, Berkeley, 

CA. 

. 1995. Sessile-flowered species in the Navarretia 
leucocephala group (Polemoniaceae). Madrofio 42: 
34-39. 

HiItTcHcock, C. L. AND A. CRONQuIST. 1973. Flora of the 
Pacific Northwest: an illustrated manual. University 
of Washington Press, Seattle, WA. 

MASson, H. L. 1946. Five new species of Navarretia. Ma- 
drono 8:196—200. 

SPENCER, S. AND J. M. Porter. 1997. Evolutionary diver- 
sification and adaptation to novel environments in 
Navarretia (Polemoniaceae). Systematic Botany 22: 
649-668. 


MADRONO, Vol. 49, No. 3, pp. 169-177, 2002 


RESURRECTION OF A CENTURY-OLD SPECIES DISTINCTION IN 
CALAMAGROSTIS 


BARBARA L. WILSON! AND SAMI GRAY 
Department of Botany and Plant Pathology, Oregon State University, 
Corvallis, OR, 97331 


ABSTRACT 


We investigated genetic diversity and taxonomic status within Calamagrostis breweri Thurber s. /., a 
rare alpine grass species comprising two allopatric races apparently differing in chromosome number. We 
analyzed isozymes and morphology of both races. The two races differ in leaf width and involution, leaf 
vein number, certain morphometric characters of the inflorescence, and habitat. Isozyme band patterns 
also distinguished the northern and southern forms, and revealed little variation within populations. Be- 
cause the northern and southern forms are genetically, ecologically, and morphologically distinct and 
apparently differ in chromosome number, they should be distinguished taxonomically as separate species. 
The southern form is here named C. muiriana sp. nov. in honor of John Muir, whose writings played in 
important role in the conservation of C. muiriana habitats. The name Calamagrostis breweri Thurber is 


retained for the northern form. 


Key words: Calamagrostis breweri, grass, polyploidy, isozymes, Calamagrostis muiriana sp. nov., rare, 


alpine, Poaceae 


The name Calamagrostis breweri Thurber is ap- 
plied to two small, glaucous, subalpine grasses that 
differ in chromosome number, isozymes, range, 
ecology, leaf anatomy, and details of morphology. 
The diversity within C. breweri sensu lato, has 
been discovered repeatedly (Kearney 1898; Nygren 
1954; S. Nugent personal communication; G. L. 
Stebbins personal communication to C. W. Greene). 
The one previous attempt to segregate the popula- 
tions nomenclaturally (Kearney 1898) foundered on 
a misinterpretation of type specimens and gave the 
northern taxon two names (C. breweri and C. lem- 
moni Kearney) while leaving the southern taxon 
nameless. Here we rectify that omission. 


MATERIALS AND METHODS 


Plants of C. breweri sensu lato were collected 
during 1995-1997, sampling both the northern 
and southern forms (Table 1). Collection sites in 
Alpine, Mono, and Tuolumne Counties, Califor- 
nia, were chosen because they appeared to be the 
Same populations sampled during an earlier study 
of chromosome numbers (Nygren 1954). Herbar- 
ium specimens prepared as vouchers for these 
collections were deposited at the Oregon State 
University herbarium (OSC). We maintained at 
least 30 individuals from each population (except 
20 from Mt. Jefferson) in the Oregon State Uni- 
versity Greenhouse. In addition, we examined 
specimens from the herbaria at California Acad- 


‘Current address: Institute for Applied Ecology, 227 
6th Street, Corvallis, OR 97333. E-mail: wilsonb@bcc. 
orst.edu. 


emy of Science, Oregon State University, Uni- 
versity of California at Berkeley, Humboldt State 
University, Rancho Santa Ana Botanic Garden, 
and the Jepson Herbarium. We also borrowed the 
isotype specimen of C. breweri from Harvard 
University and the type specimen of C. lemmoni 
from the Smithsonian Institution. 

Sixteen specimens from 14 northern populations 
and 23 specimens from at least 17 southern popu- 
lations were scored for morphological traits. Spec- 
imens used for this study are asterisked in the Ap- 
pendix; all were collected in the wild. The nine 
morphological characters scored were leaf width 
(measured near the center of leaves of innovations), 
panicle length, panicle width, outer glume length, 
inner glume length, first lemma length, callus beard 
hair length, awn length, and anther length. Results 
were analyzed by 2-tailed t-tests, using Statview 
(Abacus Concepts 1988). 

Leaf cross sections were cut by hand from the 
center portion of mature innovation leaves (those 
not on culms) from all 16 northern specimens with 
innovation leaves, plus 19 of the southern speci- 
mens. Unstained sections were examined under a 
compound microscope at 100 and drawn free- 
hand. 

For isozyme analysis, leaf tissue was ground in a 
tris buffer, pH 7.5 (Soltis et al. 1983), using | g poly- 
vinylpyrrolidone-40 per 25 ml. Wicks prepared from 
Whatman 3 mm chromatography paper were soaked 
in the resulting slurry and stored at —70°C. Methods 
of electrophoresis follow the general methodology of 
Wendel and Weeden (1989). All enzymes were re- 
solved on 12% starch gels. A histidine citrate buffer, 
pH 5.7 (Soltis et al. 1983) was used to resolve glyc- 


170 MADRONO [Vol. 49 


eraldehyde-3-phosphate dehydrogenase (G3PDH), 


oO 
3 2 ea 2 Fe é é malate dehydrogenase (MDH), phosphogluconate de- 
iB) = a eae = a hydrogenase (6PGD), and phosphoglucomutase 
S sam ESN NIN eats (PGM). A tris citrate buffer, pH 7.2 (Soltis et al. 1983) 
LON Zaza was used to resolve isocitrate dehydrogenase (IDH). 
BISHnaRRS A lithium borate buffer, pH 8.3 (Soltis et al. 1983) 
5 ate tum was used to resolve glutamate-oxaloacetate transam- 
inase (GOT), isocitrate dehydrogenase (IDH), and tri- 
osephosphate isomerase (TPI). Superoxide dismutase 
wi 5 (SOD) resolved as clear bands on gel slices stained 
§ é pha g for TPI. Enzyme stain recipes followed Wendel and 
s|/eseuag Weeden (1989). For quality control, nearly all speci- 
S 2 2 5 = B00 mens were run and stained two to eight times for each 
Of SSes buffer/enzyme combination. 
Some isozyme samples were prepared from wild- 
collected leaves, and some from leaves grown by 
5 plants held in the greenhouse for as much as three 
= 5 fe é years. When samples were prepared from both 
Slelus os wild-collected and greenhouse-grown leaves of the 
S = g 3 Ss E same individual, the isozyme patterns were identi- 
cal. Therefore, all samples were pooled for analy- 
sis. 
are Ame The southern Calamagrostis sampled are as- 
elEEcc EE sumed to be tetraploid and the northern ones are 
sie 5 50 = < assumed to be hexaploid, based on Nygren (1954) 
556606 and the herbarium specimens Stebbins 5005 and 
Stebbins 5006, which are vouchers for chromosome 
counts. Both tetraploid and hexaploid individuals 
Ya exhibit normal meiosis and may reproduce sexually 
oot Swun (Nygren 1954). However, because of the compli- 
in a a a2 a a cated banding patterns observed, and because of 
silee rs ono lack of crossing studies to determine inheritance of 
A Bb 2 B B0 bands in these species, we were unable to identify 
L<nZO<L< specific alleles and loci for some enzymes. There- 
Soph ee fore, a phenotypic instead of genotypic analysis 


was performed. Results from these eight enzyme 
stains were treated as nine enzymes, because cys- 
tolic and plastid forms of GOT were readily distin- 
guished on gels. Bands and patterns were analyzed 
in Popgene version 1.21 (Yeh et al. 1997), using 
haploid settings. Phenotypic diversity measures 
were calculated from both band presence/absence 
and multi-band patterns. For presence/absence data, 
phenotypic diversity was measured by a polymor- 
phic index (PI.) based on frequency of occurrence 
of each band. PI. = sum of f(1 — f), where f = the 
frequency of a band in a population (Chung et al. 
1991). For multi-band patterns, phenotypic diver- 
sity measures included: (1) number of patterns 
found in each population, (2) percent of stains that 
yield more than one pattern, (3) average number of 
patterns per stain in each population, and (4) Shan- 
non-Weaver Diversity Index values (Shannon and 
Weaver 1949), based on frequency of each pattern 


Voucher specimen collector, number 
Sami Gray and Barbara L. Wilson 8269 
Sami Gray and Barbara L. Wilson 8258 


Barbara L. Wilson s.n. 
Sami Gray and Barbara L. Wilson 8044 
Sami Gray and Barbara L. Wilson 8901 


1995-1997 COLLECTION SITES FOR ALPINE CALAMAGROSTIS. 
Mike Roantree s.n. 


88 
eal sss 5 iS 3 in each population. Phenotypic relationships among 
eas S S S as 3 populations were determined by calculating Hed- 
es GISSSSESE rick’s phenotypic identities (Hedrick 1971) for band 
Eb SRST SRSTSTS data and for multi-band pattern data. 


2002] 


Fic. 1. 


Leaf cross sections of Calamagrostis breweri and 
C. muiriana. A—C. Calamagrostis muiriana. D-—E Cala- 
magrostis breweri. 


RESULTS AND DISCUSSION 
Morphology 


Foliage characteristics distinguished northern 
and southern forms. In the southern form, leaves of 
the innovations (tillers) were involute and extreme- 
ly thin (0.2—0.35 [—0.4] mm wide) with three (rarely 
four) veins (Fig. 1A—C). In the northern plants, 
such leaves were wider ([0.35] 0.4—0.6 mm wide 
when rolled, 0.9—1.1 mm wide when flat) with sev- 
en or more veins (Fig. 1D—F). These foliage dif- 
ferences between the taxa remained conspicuous 


WILSON AND GRAY: CALAMAGROSTIS 


171 


over years of greenhouse growth. As reported by 
Nygren (1954), southern and northern C. breweri 
also differed in morphology of the flag leaf (up- 
permost leaf on the flowering culm). Northern 
plants had about ten veins in flag leaves, while 
southern plants characteristically had five. Differ- 
ences in leaf size and shape may be attributable to 
the difference in chromosome number between the 
northern and southern populations; higher ploidy 
level is often associated with wider but shorter 
leaves (Tal 1980). 

The two forms also differed in inflorescence 
characters, with statistically significant differences 
in panicle length, panicle width (which is, however, 
strongly influenced by maturity), length of outer 
glume, and length of longest callus beard hairs (Ta- 
ble 2). Although morphological differences be- 
tween northern and southern populations were 
small, they were consistent. 


Isozyme Analysis 


Isozyme band patterns for the enzymes GOTS, 
PGD, and TPI readily distinguished southern from 
northern populations (Table 3). With the exception 
of Mt. Hood and Mt. Jefferson, all populations 
were distinguished by MDH patterns (Table 3). All 
isozyme bands and patterns observed in the invari- 
ant Mt. Jefferson population were also observed in 
Mt. Hood populations, even though the Mt. Jeffer- 
son sample included four subpopulations. Little iso- 
zyme variation was observed within populations, 
and two populations (Mt. Jefferson and Mt. Dana) 
were invariant (Table 4). 

Hedrick’s phenotypic similarity based on fre- 
quencies of isozyme band patterns (Table 5) was 
used to estimate similarity of the Calamagrostis 


TABLE 2. COMPARISON OF CERTAIN MORPHOLOGICAL CHARACTERS IN C. MUIRIANA FROM C. BREWERI. 


Sample 

Characteristic (units) Taxon size Mean (s.d.) p= Range 
Leaf width (mm) C. breweri 15 0.46 + 0.018 0.0001 0.40.6 
C. muiriana 20 O27 == 0:02 0.2—0.4 

Leaf vein number C. breweri 14 OOD TA X 107% 7-9 

C. muiriana 19 Bel O07 2, 3—4 
Panicle length (cm) C. breweri 16 FO= 020 0.0001 5.7-8.4 
C. muiriana 2D, AOS 1.9—7.5 
Panicle width (cm) C. breweri 16 2.2°= O52 0.0343 0.7—5.2 
C. muiriana 21 1.4 + 0.18 0.4-—3.0 
Length of outer glume (mm) C. breweri 16 AWS =O. 02 0.0005 3.1-—4.9 
C. muiriana 25 5-008 3.0—4.4 
Length of inner glume (mm) C. breweri 16 3.4 + 0.09 0.0778 3.3—4.5 
C. muiriana 2S 351 <=70:08 3.0—4.5 
Lemma length (mm) C. breweri 16 335 OO 0.3338 2.6—4.0 
C. muiriana 23 3.2006 2.6—3.8 
Callus beard hair length (mm) C. breweri 16 0.7 + 0.06 0.0001 0.3—1.2 
C. muiriana 25 Oe 0.02 0.3-0.6 
Lemma awn length (mm) C. breweri 16 A: Gee OAS 0.5693 3.4-5.5 
C. muiriana 23 4.5 + 0.09 3.3-5.9 
Anther length C. breweri 11 DAY SOLD 0.1782 1.3—2.6 
C. muiriana 18 1.8 + 0.09 0.9—2.3 


172 


MADRONO 


[Vol. 49 


TABLE 3. FREQUENCIES OF VARIABLE ISOZYME PATTERNS IN CALAMAGROSTIS MUIRIANA (TIOGA AND DANA POPULATIONS) 
AND C. BREWERI (HOOD, JEFF, CARSON, AND EDDY POPULATIONS). In addition, enzymes GOTE G3PDH, IDH, PGM, and 
SOD were invariant across all tested populations. Blank means frequency = zero. 


Hood 


0.036 
0.964 


Dana 


1.000 


Enzyme Pattern 


GOTS 
GOTS 
MDH 
MDH 
MDH 
MDH 
MDH 
MDH 
MDH 
MDH 
MDH 
MDH 
PGD 
PGD 
PGD 
Tell 
TPI 
WAL 


Tioga 


O27 
0.028 
0.971 
0.029 


1.000 


0.500 
0.500 
1.000 simi 
1.000 
1.000 1.000 
1.000 


QABrFORMraHrTOAMoOeS aS 


populations. Material from Mt. Dana resolved poor- 
ly and no data was obtained for three enzymes (Ta- 
ble 3). Each similarity between the Mt. Dana pop- 
ulation another population is based on only six of 
the nine enzymes assayed. The two southern pop- 
ulations formed one cluster and the four northern 
populations formed a second (Fig. 2). 


Chromosome Numbers 


We relied on previously reported chromosome 
counts for these grasses, and collected from popu- 
lations in Mono, Tuolumne, and Alpine Counties, 
California, that are the same populations or very 
close to the populations sampled in a study of Cal- 
amagrostis chromosome counts (Nygren 1954). 
The thin-leaved southern plants appear to be tetra- 
ploid, as indicated by published chromosome 
counts from Mono, and Tuolumne, Counties (Ny- 
gren 1954) and counts for herbarium specimens 
from Mariposa and Mono Counties (Stebbins 5005 


TABLE 4. ISOZYME PHENOTYPE STATISTICS FOR CALAMA- 
GROSTIS BREWERI AND C. MUIRIANA. N = sample size/en- 
zyme. P = percent polymorphic enzymes. A = average 
number of patterns per enzyme. PI. = polymorphic index 
(see methods). S-W = Shannon-Weaver Diversity Index. 


Population N* ps A* Pell. S-W 

Tioga 32 22% 1.222 0.3093 0.0248 
Dana 24 0% 1.000 0.0000 0.0000 
Hood DI WAG NPA VOX  OMS447/ 
Jeff 18 0% 1.000 0.0000 0.0000 
Carson 29 33% 1.444 0.6175 0.0492 
Eddy Dy 22% 1.333 0.6685 0.0508 
C. muiriana 44 22% 1.222 0.1989 0.0199 
C. breweri 90 44% 2.111 2.0862 0.2784 
Over all 134 44% 2.555 3.0702 0.4600 


Jeff Carson Eddy Band pattern 
001 
1.000 1.000 1.000 Lid 
00100001010001 
ALA MOOLOLOOOL 
0.167 00100001110101 
0.567 GOOLOLOOLL NOLO i 
0.267 00101001010101 
0.028 AO AE MOO OIE OAL OAL 
0.111 GLALOOOL LALO LOAL 
0.861 OO OOOO WO GLa 
1.000 GOLOOOOL LI OLOG 
00100001101100 
Lah a 
1.000 0.286 0.233 O11 
0.714 0.767 001 
OIL 
1.000 0.967 1.000 OOL 
0.033 Lid 


and Stebbins 5006). Only hexaploid plants were 
found in the one northern population (in Alpine 
County) for which chromosome counts were pub- 
lished (Nygren 1954). 


Ecology 


Published accounts and labels from 96 collec- 
tions of the southern and 26 of the northern pop- 
ulations (see Appendix) indicate that the southern 
and northern populations differ in habitat and 
abundance. Nygren (1954) reported that tetra- 
ploids grew in high mountains at and above 3000 
m elevation, while hexaploids grew at 1800 to 
2100 m in the woods. Specimen labels show that 
the altitudinal range of the southern plant is 
2484-3900 m (mean = 3191 + 33 m; n = 73); 
that of the northern form is 1700—2600 m (mean 
= 2158 + 52 m; n = 21). However, that differ- 
ence may be less important than it appears; both 
forms grow near timberline, although the south- 
ern one also grows above it. 

Labels for southern populations frequently list 
the habitat as moist or dry meadows. (The distinc- 
tion may be seasonal; the plants grow in flood- 
plains.) While labels for the northern plants are fre- 
quently uninformative, three mention trees or shade 
(Hitchcock & Martins 5413, Stebbins 5009, Steb- 
bins 7771). Northern plants may grow in full sun 
or partial shade (personal observation), with one 
population seen in full shade (Susan Nugent per- 
sonal communication). 

Herbarium labels report the southern plant as 
“abundant” (Sharsmith 2011), “‘“excellent feed and 
abundant at and above timberline’’ (Hatton H-11), 
“‘large abundance at high altitudes in the Kern .... 
One of the best forage plants of the higher ranges” 
(Morrow 5), and the “‘most abundant grass”’ (Ferris 


2002] 


WILSON AND GRAY: CALAMAGROSTIS 173 


TABLE 5. HEDRICK’S (1971) PHENOTYPIC SIMILARITY BETWEEN POPULATIONS OF CALAMAGROSTIS BREWERI (MT. Hoop, MrT. 
JEFFERSON, CARSON PASS, AND MT. EDDY) AND C. MUIRIANA (TIOGA PASS AND MT. DANA). 


Tioga Dana Hood Jeff Carson Eddy 
Tioga 
Dana 0.86138 
Hood 0.45953 0.37223 
Jeff 0.43944 0.344828 0.83399 
Carson 0.46035 0.355837 0.728548 0.705464 
Eddy 0.46713 0.366033 0.737174 0.713429 0.790985 


9824). Southern plants are community dominants, 
but northern plants are not (personal observation). 
In contrast, northern plants have been described as 
occasional at Mt. Eddy (Whipple 1981) and rare in 
the Lake Tahoe region (Smith 1984) and in Oregon 
(Anonymous 1995). 


Taxonomic History 


Calamagrostis breweri was first described by 
Thurber, who wrote that the leaves were “‘seta- 
ceously involute’’ (Thurber 1880, p. 281). Later de- 
scriptions agreed, calling the leaves ‘“‘usually in- 
volute filiform’? (Hitchcock 1912; Abrams 1940; 
Hitchcock and Chase 1950) or “‘usually involute”’ 
(Munz and Keck 1959), or “flat or inrolled”’ 
(Greene 1993). Setaceously involute leaves are 
characteristic of the southern, not the northern, 
form (personal observation). Thurber based his de- 
scription of C. breweri upon three specimens: 
Brewer 2128 from near the summit of Carson’s 
Pass, Bolander 6098 from the Tuolumne River, and 
Lemmon s.n. from “‘California.”’> Calamagrostis 
breweri was effectively lectotypified by the citation 
of Brewer 2128 as the type (Hitchcock and Chase 
1950). We have examined these specimens. Brewer 
2128 and Lemmon s.n. are individuals of the north- 
ern populations, with relatively wide leaves that 
rolled up as they wilted. Only Bolander 6098 has 


Tioga Pass 


C. muiriana 


Mt. Dana 
Carson Pass 
Mt. Eddy 
Mt. Jefferson 


Mt. Hood 


0.40 0.60 0.80 1.00 


Hedrick’s (1971) genetic similarity 


Fic. 2. Cluster diagram based on Hedrick’s similarities 
of Calamagrostis breweri (Mt. Hood, Mt. Jefferson, 
Carson Pass, and Mt. Eddy populations) and C. muiriana 
(Tioga Pass and Mt. Dana populations), calculated from 
isozyme pattern frequencies. 


truly setaceously involute leaves and represents the 
southern form. 

Kearney (1898) apparently observed the same 
morphological variation within C. breweri that 
prompted this study. He split the taxon into two 
species, C. breweri as the name was commonly ap- 
plied, and a new species, C. Jemmoni, characterized 
by broader leaves. He stated that C. lemmoni was 
“intermediate between C. deschampsioides and C. 
breweri”’ (Kearney 1898). He cited Lemmon s.n. as 
the type specimen. Lemmon s.n. is a broad-leaved 
northern plant, different from the majority of plants 
to which the name C. breweri has been traditionally 
applied. However, both Lemmon s.n. and Brewer 
2128 are broad-leaved northern plants. The name 
C. lemmoni was appropriately relegated to synon- 
ymy with C. breweri by Hitchcock (1912). Kear- 
ney’s attempt to split C. breweri left the northern 
form with two names, and the narrow-leaved south- 
ern plant nameless. Doubtless this confusion re- 
sulted from the fact that leaves of both Lemmon 
s.n. and Brewer 2128 had wilted and rolled up 
tightly before pressing, so that they superficially ap- 
pear to be setaceously involute. 

Nygren (1954) detailed the cytology of C. brew- 
eri and rediscovered the its morphological varia- 
tion, but did not treat it taxonomically. He found 
that plants of two southern populations (near Tioga 
Pass in Mono County and Mt. Dana in Tuolumne 
County) were tetraploids (2n = 28), and plants 
from a northern population (near Carson Pass, Al- 
pine county) were hexaploids (2n = 42). Nygren 
also noted that the tetraploid and hexaploid forms 
could be distinguished by the shape of “‘the upper- 
most leaf of the straw [culm]? (Nygren 1954). 
More recently, Greene (1993) acknowledged the 
range of morphology, describing C. breweri leaves 
as “‘flat or involute.”’ 


CONCLUSIONS 


Kearney (1898) was correct that Calamagrostis 
breweri as traditionally understood includes two 
entities that differ in morphology and habitat. Per- 
haps the differences between the two taxa are due 
to consistent differences in chromosome number; 
ploidy level in itself can affect the morphological 
traits and habitat preference like those that differ- 
entiate the two forms of C. breweri (Tal 1980). 
Ploidy levels can represent barriers to gene flow, 


174 


and therefore populations that differ in chromo- 
some number may be treated as separate species 
(Harlan and de Wet 1971). However, ploidy level 
may mean little taxonomically; grasses of two ploi- 
dy levels may mingle in a population without ob- 
vious morphological or ecological differentiation 
(Hultquist et al. 1997; Keeler et al. 1987). Taxon- 
omists should avoid encumbering the taxonomic lit- 
erature with species that differ only in ploidy level 
(Lewis 1980), but in C. breweri the southern (pre- 
sumably tetraploid) and northern (presumably 
hexaploid) forms exhibit sufficient morphological, 
ecological, leaf anatomical, and isozyme differenc- 
es that they can reasonably be recognized as dif- 
ferent species. 

We name the southern species after John Muir, 
pioneering naturalist and first president of the Sierra 
Club. Muir wrote extensively about the Sierras and 
in particular the Yosemite area. In the summer of 
1869, he spent more than a month in Big Tuolumne 
Meadows “sketching, botanizing, and climbing 
among the surrounding mountains” (Muir 1894, p. 
70). He must frequently have walked or slept on 
this small reedgrass that carpets the floodplain 
meadows of the Yosemite region. 


Calamagrostis muiriana B. L. Wilson and Sami 
Gray sp. nov.—TYPE: USA: California: Tuol- 
umne County: Yosemite National Park; Dana 
Fork, Tuolumne River, Elevation: 9525 feet. 1 
August 1997. B. L. Wilson and S. Gray 8909. 
(Holotype: OSC; isotypes, CAS, NY, RSA, UC, 
US, UTC). 


Gramen pusillum alpinum, maxime similare Cal- 
amagrostidi breweri, sed tetraploideum et foliis an- 
gustioribus trinerviis. Gramen perenne caespito- 
sum, 12—34 cm altum, culmis florentibus folia ex- 
cedentibus. Folia perangusta glauca, vaginis folior- 
um marginibus imbricatis, ligulatis 0.8—2.2 mm 
longis, laminis filiformibus glaucis glabris, 4—12 
cm longis, 0.2—0.4 mm latis, venis ternis. Inflores- 
centia paniculata erecta, pauciflora, atroviolacea, 
1.9-7.5 cm longa, ramis expansis. Spiculae atro- 
violaceae, uniflorae, rachilla sterili pilis albis, glum- 
is aequalibus, 3—4.5 mm longis, atroviolaceis, at- 
tenuatis vel minutae aristatis. Lemma hyalinum vel 
atroviolaceum, 2.5—4.0 mm longa, apicale denti- 
culatum, dentibus quatuor, pusillis, atroviolaceis, 
pilis calli albis 0.3—0.6 mm longis, arista lemmatis 
geniculata, 3.3-6.0 mm longa, infra medium cari- 
nae orienti. Palea hyalina, lemmati aequanti. An- 
therae maturae 0.9—2.5 mm longae, purpuratae. 
Species tetraploidea, chromosomata 28. 

Gramen abundans in pratis alpinis apricis, dis- 
tributum in montibus excelsis Californiae cen- 
tralis. 

A Calamagrostide breweri laminis angustioribus 
venis ternis, paniculis pusillioribus, pilis calli bre- 
vioribus distinguenda. 


Plants cespitose, with intravaginal shoots. Young 
plants densely tufted. Old plants spreading outward 


MADRONO 


[Vol. 49 


while dying in the middle and thus forming rings 
3 dm or more in width. Foliage glaucous, 6-15 cm 
long. Leaf sheaths open, glabrous to retrorsely 
short-pubescent. Ligules 0.8—2.2 mm long, entire, 
rounded apically. Leaf blades involute, well-devel- 
oped leaves 4—12 cm long 0.2—4 mm wide as in- 
volute, (but earliest leaves of innovations with 
blades ca. 0.5 cm long and 0.2—0.4 mm wide), leaf 
blades abaxially scabrous on veins and glabrous or 
scabrous between them, adaxially pubescent; leaf 
blades of innovations with 3 (—5) veins and usually 
7 sclerenchyma bundles; leaf blades of culms with 
5 veins. Leaf tip straight-sided, not prow-shaped. 
Flowering culms taller than the foliage and 12—35 
cm tall. Inflorescence a spreading panicle, few- 
flowered, 1.9—7.5 cm long and 0.4—3.0 cm wide, 
dark purple or rarely straw-colored. Spikelets one- 
flowered, with a sterile rachilla about half as long 
as the floret and covered with long white hairs. 
Glumes equal in length, 3—4.5 mm long, thin-tex- 
tured, purple, often with hyaline margins distally 
(occasionally straw-colored), acute to apiculate, mi- 
nutely awned, or attenuate, rounded on the back or 
keeled distally, glabrous (occasionally scabrous), 
sometimes with hairs on the keel. Lemma 2.5—4.0 
mm long, thin-textured, hyaline or dark purple, gla- 
brous or scabrous, usually with purple on veins dis- 
tally, the veins extending as four short (ca. 0.5 mm 
long) teeth. Callus beard hairs white, 0.3—0.6 mm 
long. Lemma awn purple, arising from below the 
middle of the lemma, 3.3—6.0 mm long, geniculate, 
extending beyond the glumes. Palea hyaline, pig- 
mented along veins distally, about as long as the 
lemma. Mature anthers 0.9—2.5 mm long, purple. 
2n = 28. 


Descriptions of Calamagrostis breweri are, in 
most instances, descriptions of Calamagrostis mui- 
riana. We therefore re-describe Calamagrostis 
breweri sensu stricto here: 


Plants cespitose, with intravaginal (occasionally 
extravaginal) shoots. Young plants densely tufted. 
Old plants spreading outward while dying in the 
middle and thus forming rings ca. 1.5 dm in di- 
ameter Foliage glaucous, 10—20 cm long. Leaf 
sheaths open, scabrous. Ligules 1.7—4.1 mm long, 
entire to erose. Leaf blades flat but readily rolling 
when dry, well-developed leaves (2—) 10-15 cm 
long, 0.4—0.6 mm wide when rolled, the innovation 
leaves 0.9—1.1 mm wide when flat, (earliest leaves 
of innovations only slightly reduced), the culm 
leaves 1.3—1.7 mm wide when flat, leaf blades 
abaxially scabrous, adaxially pubescent; leaf blades 
of innovations with 7—9 veins and 9-11 abaxial 
sclerenchyma bundles; leaf blades of culms often 
with 11 or more veins. Leaf tip prow-shaped. Flow- 
ering culms taller than the foliage and 29-54 cm 
tall. Inflorescence a spreading panicle, few-flow- 
ered, 5.7—8.4 cm long, and 0.7 to 5.2 cm wide, pale 
to dark purple. Spikelets one-flowered, with a ster- 


2002] 


Fic. 3. Calamagrostis muiriana. A. habit. B & C. spike- 
lets. D. floret. E. cross section of leaf blade. FE cross sec- 
tion of leaf sheath. 


ile rachilla about half as long as the floret and cov- 
ered with long white hairs. Glumes subequal in 
length, the lower 3.1—4.9 mm long; the upper 3.3— 
4.5 mm long. Glumes thin-textured, pale to dark 
purple or sometimes greenish, often with hyaline 
margins distally (occasionally straw-colored), acute 
to apiculate, minutely awned, or attenuate, rounded 
on the back or keeled distally, glabrous (occasion- 
ally scabrous), sometimes with hairs on the keel. 
Lemma 2.6—4.0 mm long, thin-textured, hyaline or 
dark purple, glabrous or scabrous, usually with pur- 
ple on veins distally, the veins extending as four 
short teeth. Callus beard hairs white, 0.3—1.2 mm 
long. Lemma awn purple, arising from below the 
middle of the lemma, 3.4—5.5 mm long, geniculate, 
extending beyond the glumes. Palea hyaline, pig- 
mented along veins distally, about as long as the 
lemma. Mature anthers 1.3—2.6 mm long, purple. 
2n = 42. 


WILSON AND GRAY: CALAMAGROSTIS 175 


Calamagrostis breweri 


Calamagrostis muiriana 


Fic. 4. Distribution of Calamagrostis breweri (@) and 
C. muiriana (*) in Oregon and California. 


KEY TO ALPINE REEDGRASSES OF WESTERN NORTH 
AMERICA 


1. Leaves extremely thin and involute, 0.25—0.4 mm 
wide as rolled, with 3 (—4) veins, the tip straight- 
sided; panicle length 1.9—5.7 cm; callus beard hair 
length 0.3—0.6 mm; range south of Sonora Pass 
(Yosemite area and south; Fig. 4) ..... C. muiriana 

1. Leaves narrow, flat in life but readily rolling when 
wilted, 0.4—0.6 mm wide as rolled, with 7 or more 
veins, the tip prow-shaped; panicle length 5.7—8.5 
cm; callus beard hair length 0.3—1.2 mm; range 
north of the Sonora Pass (Carson Pass area north to 
Mt= Hoods @Oreson)i 24 5...... 2: C. breweri s. str. 


ACKNOWLEDGMENTS 


We thank Dr. Aaron Liston for his guidance, and Susan 
Nugent for initiating this study. Heather Laub and Mike 
Roantree collected plants at Mt. Hood and Mt. Jefferson, 
respectively. We thank Dr. Richard Halse, curator of the 
Oregon State University Herbarium, for assistance in ob- 
taining herbarium specimens. Jan Van Wagtendonk and 
Leslie Chow provided the permit necessary to collect in 
Yosemite National Park. The study was supported finan- 
cially by the Mt. Hood National Forest and the Native 
Plant Society of Oregon. We thank Dr. Craig Greene for 
his helpful comments and for making available a copy of 
his dissertation on Calamagrostis. Finally, we thank the 
following institutions for loans of herbarium specimen: 


176 MADRONO 


Rancho Santa Ana Botanical Garden, Humboldt State 
University, California Academy of Science, and the Uni- 
versity of California at Berkeley. 


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GREENE, C. W. 1993. Calamagrostis. Pp. 1243-1246 in J. 
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AND A. CHASE. 1950. Manual of grasses of the 
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berta, Canada. 


APPENDIX 1 


SPECIMENS EXAMINED 
* = specimen included in morophometric study; 
** = type specimen 


Calamagrostis muiriana: USA: California: Fresno 
Co., Baxter Lake, 6 Sep 1959, DeDecker 1119 (CAS); N 
of Kings River, Red Mt. Basin, 16 Aug 1961, Hardham 
8748 (CAS); Silver Pass in Fish Creek Country; Sierra 
Nat’! Forest; 19 Sep 1912, Hatton H-109 (CAS); Death 
Pond, Kaiser Ridge, 12 Aug 1928, Jepson 13229* (UC); 
Bear Cr. Watershed, Mt. Hilgard, 20 Aug 1951, Quibell 
680 (RSA); Bear Cr. Watershed, 2 mi W of Mt. Hilgard, 
21 Aug 1951, Quibell 767 (RSA); Bighorn Lake, head- 
waters N Fk Mono Cr., 2 mi W of Red & White Mt., 31 
Aug 1952, Quibell 1575 (RSA); Bighorn Lake, headwa- 
ters N Fk Mono Cr., 2 mi W of Red & White Mt., 31 Aug 
1952, Quibell 1583 (CAS); Bear Creek, near Florence 
Lake, 7 Sep 1954, Quibell 4985 (OSC, RSA); Upper 
French Canyon Basin (near Bishop), 6 Aug 1955, Quibell 
5387 (RSA); Colby Meadows, 24 Jul 1952, Raven 4677 
(CAS), Laurel Creek, 13 August 1953, Raven 6174 
(CAS); Rose Lake, 8 Aug 1954, Raven 7858 (CAS); 
Bench Lake, 25 Jul 1956, Raven 9846 (CAS); Humphrey’s 
Basin, W of Mt. Humphreys, 12 Aug 1937, Sharsmith 
3165* (CAS, UC); Mono Pass, 118°50’W, 37°25'N, 10 
Sep 1959, Thomas 8144 (CAS); Second Recess, 18 Jul 
1953, Raven 5694 (CAS); Inyo Co., Paiute Pass; 11,300— 
11,409 ft., 22 Jul 1934, Ferris 8877* (CAS, UC); Rock 
Cr. Lake Basin, Mosquito Flat, 14 Jul 1946, Howell 22277 
(CAS); Mono Mesa, 26 Jul 1946, Howell 22730 (CAS); 
Big Pine Lakes, 5th Lake, 5 Aug 1947, Howell 23767 
(CAS, RSA); Rock Cr. Lake Basin (Long Lake), 22 Jul 
1931, Peirson 9398 (RSA); Rock Cr. Lake Basin, 17 Jul 
1931, Peirson 9399 (RSA); Rock Cr. Lake Basin, Ruby 
Falls, 20 Jul 1934, Peirson 12617* (CAS, RSA, UC); 
Coyote Ridge, 7 Aug 1950, Raven & Stebbins 217 (CAS); 
Coyote Ridge, 2 Aug 1950, Raven & Stebbins 227* (UC); 
Mono Pass; Ruby Lake, 16 Jul 1936, Robinson 689 
(RSA); Lone Pine Canyon, E of Mt. Muir, 20 Aug 1937, 
Sharsmith 3310* (UC); Mono Mesa near junction of 
Mono, Inyo, & Fresno Co., 37°28'N, 118°46'W, 9 Sep 
1961, Thomas 9747 (CAS); Madera Co., Lyell Fork of 
Merced River, Yosemite Nat’] Park, 1 Aug 1931, Blasdale 
s.n. (UC); Minarets, 23 Aug 1918, Grant 1585 (CAS); 
Minarets, 23 Aug 1918, Grant 1587 (CAS); Mariposa 
Co., Soda Springs by the Tuolumne, 15 Aug 1894, Cong- 
don s.n. (CAS); Yosemite Nat’! Park, % mi N of Tenaya 
Lake, 18 Jul 1951, Stebbins 5005* (UC); Emeric Lake, 
T2S R24E S8, 15 Sep 1935, Thomas 490* (UC); May 
Lake, 15 Jul 1977, Vale s.n. (CAS); Mono Co., Slate Cr. 
Basin, E of Mt. Conness, 7 Sep 1934, Clausen 922 (CAS); 
Tioga Pass, Yosemite Nat’! Park boundary, 16 Aug 1936, 
Ferris 9824* (CAS, UC); Tioga Pass, just S of entrance 
to Yosemite Nat’] Park, Heller 15461 (CAS); Mono Pass, 
16 Aug 1944, Howell 20631 (CAS); Saddlebag Lake, 


payee 


2002] 


head of Lee Vining Cr., 18 Jul 1968, Howell & True 4432 
(CAS); Saddlebag Lake, head of Lee Vining Cr. 18 Jul 
1968, Howell & True 4435 (CAS): Slate Cr. Basin, 26 
Aug 1937, Keck 4591 (CAS, UC); HM Hall Natural Area, 
Green Lake, N base of Mt. Conness, 27 Aug 1927, Keck 
4610 (CAS); HM Hall Natural Area; saddle between Mt. 
Conness & East Plateau, 27 Aug 1927, Keck 463] (CAS); 
Slate Cr. Valley, Inyo Nat'l Forest, 26 Jul 1954, Krucke- 
berg 3602 (CAS, RSA); Slate Cr., 17 Aug 1954, Munz 
19987 (RSA); Slate Cr., Aug 1954, Munz 20088 (RSA); 
Saddlebag Lake, 18 Jul 1968, Rose 68/51] (CAS, HSC, 
RSA); Tioga Pass, 14 Aug 1930, Stanford 2029 (RSA): 
summit of Dana Plateau; 10 km W of Mono Lake, 19 Jul 
1951, Stebbins 5006* (UC): Mono Basin, Parker Cr. Ba- 
sin, | Aug 1987, Taylor 9210* (RSA, UC); Mono Pass, 
4th Recess, 37°26'N, 118°48’W, 11,000—12,000 ft., 13 Sep 
1954, Thomas & Thomas 4630 (CAS): Mt. Conness Re- 
gion, NE of Steelhead Lake, 11 Sep 1959, Twisselmann 
5700 (CAS); Saddlebag Lake, 16 Aug 1933, Wolf 5334 
(RSA); below Highway 120 near Yosemite E entrance, 11 
Aug 1970, Wood 242 (HSC); near Tioga Pass, TIN R25E 
$30, 22 Aug 1936, Yates 6301 (RSA, UC); Tioga Pass, 
TIN R25E S30, 22 Aug 1936, Yates 6304* (CAS, UC); 
a mile above Conness Lake, Convict Cr. drainage, 26 Aug 
1968, Zufeldt 2 (CAS); Tulare Co., Kern Lake, 11,000— 
12,100 ft. CAP 2074* (UC); Mt. Whitney, 11 Jul 1910, 
Clemens s.n. (CAS); Yosemite Nat’] Park, Mt. Conness, 
19 August 1897, Dudley 1703 (CAS); between Reflection 
Lake & Harrison Pass, 8 Aug 1940, Howell 16048 (CAS); 
Little Five Lakes Basin, 29 Jul 1942, Howell 17379 
(CAS); Sky Parlor Meadow, Chagoopa Plateau, 1 Aug 
1942, Howell 17520 (CAS): Center Basin, 26 Jul 1948, 
Howell 25042 (CAS, RSA); Rock Creek, 21 Jul 1949, 
Howell 25568* (CAS, UC); Mineral King vicinity, Fare- 
well Gap, 20 Jul 1951, Howell 28013 (CAS, RSA); Min- 
eral King Vicinity, White Sheif Region, 21 July 1951, 
Howell 28088 (CAS); Diamond Mesa, 25 Jul 1954, Kehr- 
lein (CAS); Kern Nat'l Forest (Mt. Whitney District), 
Morrow 5 (CAS); Mineral King Game Refuge, Sequoia 
Nat'l Forest, N of Franklin Lakes, 8 Aug 1966, Rice 5/6 
(OSC, RSA): Kern Plateau, Twisselmann et al. 11280 
(RSA); Chicken Spring Lake, Kern Plateau, 31 Jul 1970, 
Twisselmann et al. 17396 (CAS, RSA): Tuolumne Co., 
Tuolumne Meadows, September 1866, (Bolander?) 6098* 
(UC); Tuolumne Meadows, Yosemite Nat’! Park, Babcock 
3627 (UC); Slide Canyon W of Finger Peak near N 
boundary of Yosemite Nat’! Park, 31 July 1934, Barthol- 
omew s.n. (UC); Mt. Lyell Quadrangle, 2 miles W of Mt. 
Gibbs, 6 Aug 1937, Bufford s.n.* (UC); Yosemite Nat’l 
Park, Lake Ireland, 29 Jul 1917, Clemens s.n. (CAS); Mt. 
Dana, 16 Aug 1894, Congdon s.n.* (CAS, UC): Yosemite 
Nat’! Park, Mt. Conness, 9 Aug 1934, DeRoy 115 (CAS); 
Gaylor Lake (upper), Yosemite Nat’l Park, Hall 11864 
(CAS, UC); Tioga Pass, Yosemite Nat’! Park, 13 Sep 
1922, Hall s.n. (CAS); Dog Lake, 6 Aug 1944, Howell 
20034 (CAS); Gaylor Lake, 12 Aug 1944, Howell 20372* 
(CAS, UC); Lyell Fk of Tuolumne River, 25 Jul 1936, Lee 
2334* (UC): Yosemite Nat’! Park, 12 Mar 1909, Lemmon 
1897 (CAS); Tioga Pass, 30 Aug 1957, Rose 57127 (CAS, 
RSA, UC); Lyell Fk. of Tuolumne River, upper Lyell Can- 


WILSON AND GRAY: CALAMAGROSTIS ee 


yon, 25 Jul 1933, Sharsmith 189 (CAS, UC); Dana Mead- 
ows, west base of Mt. Dana, 15 Aug 1933, Sharsmith 
414* (UC); Mt. Dana, west slope, | Aug 1931, Sharsmith 
S00 (UC); Mt. Dana, NW plateau, 10 Sep 1934, Sharsmith 
2100 (CAS, US); Mt. Dana, NW slope, 2 Aug 1933, 
Sharsmith 257B (UC); Gaylor Lake, Yosemite Nat’! Park, 
23 June 1976 Vale & Wagnon s.n. (CAS); Tuolumne 
Meadows, Mt. Lyell Quadrangle, 22 Aug 1936, Yates 
6325* (CAS, RSA, UC): Tuolumne Meadows, Yosemite 
Nat’! Park, 8500—9500 ft., Jul 1902, Hall & Babcock 3627 
(UC). 

Calamagrostis breweri: USA: California: Alpine (?) 
Co., near summit of Carson’s Pass, Brewer 2/28** (GH, 
US); Lake Winnemucca ca. 2 miles south of Carson Pass, 
17 Aug 1996, Gray & Wilson 8270 (OSC); Amador Co., 
Woods Lake, summit of Carson Pass, 20 Jul 1951, Steb- 
bins 5009* (UC); Eldorado Co., Echo Lake, 11 Aug 
1981, Best s.n.* (CAS); E side Wright’s Lake, T12N 
RI6E, 16 Jul 1977, Stebbins 7771* (CAS); Trail from 
Suzie to Heather Lake, 17 Aug 1928, Wolf 3338* (RSA); 
Nevada Co., Basin Peak, 27 Aug 1968, True 4590* 
(CAS); 1 mi SW of Mt. Lola, White Rock Lake, 7920— 
8400 ft., 28 Jul 1976, Trowbridge SOO0* (CAS); 1 mi SW 
of Mt. Lola, White Rock Lake, 28 Jul 1976, Trowbridge 
S074 (CAS); Placer Co., Donner Pass, 10 Aug 1903, 
Heller 7130* (CAS, OSC, UC); Siskiyou Co., English 
Peak, Diamond Lake, 2 Aug 1968, Oettinger 345* (RSA); 
English Peak, Marble Mt. Wilderness Area, Shasta Ridge, 
3 Sep 1969, Oettinger 1613* (RSA, UC); Shasta Ridge, 
N slope of English Peak, 3 Sep 1969, Oettinger 1613 
(HSC); S side Caribou Lake, T37N RIOW S35, 19 Aug 
1980, Renner 2506 (HSC); above Sugar Lake, T40N ROW 
S31, 4 Aug 1969, Sawyer 1803* (OSC); above Sugar 
Lake, 4 Aug 1969, Sawyer 1803 (HSC); W of Little Duck 
Lake, T40N ROW S19, 14 Sep 1972, Smith & Sawyer 
5813 (HSC); Mt. Eddy, north face, T40N R5W S7, 25 Jul 
1976, Whipple 1576 (HSC); Mt. Eddy, north slopes, T40N 
RSW S7, 20 Jul 1977, Whipple 2006 (HSC): near Little 
Crater Lake, T40ON R5W S5&6, 5 Aug 1977, Whipple 
2183 (HSC); 1.45 mile by road north of Parks Creek Sum- 
mit, T41N R6W S33, 16 Aug 1996, Wilson & Gray 8258 
(OSC); Trinity Co., Stuart Fk. drainage, west of Morris 
Lake, T36N RIOW S22, 24 Aug 1997, Ferlatte 1899* 
(HSC, UC); Lower Canyon Cr. Lake, 10 mi W of Dedrick, 
Salmon-Trinity Alps, 11 Jul 1939, Hitchcock & Martins 
5413* (CAS, UC); Echo Lake, T35N ROW S3, 12 Aug 
1994, Rolle 853 (OSC); California, no county, [no lo- 
cation], Lemmon s.n.** (US); [no location], 18 Feb 1909, 
Lemmon s.n. (US); Oregon: Hood River Co., Mt. Hood 
(SE; upper Sahalie Falls), 2 Aug 1924, Henderson 1045 
of 1924* (CAS, OSC); Mount Hood Meadows Ski Area, 
T3S R9E S4, 11 Sep 1995, Laub s.n. (OSC); Mt. Hood, 
Heather Canyon, T2S R9E S24, 1 Aug 1999, Nugent s.n. 
(OSC); Mt. Hood, T2S R9E S34, Poff s.n. (OSC); Mt. 
Hood, T2S R8.5E S13, 4 Sep 1979, Siddall s.n.* (OSC); 
Paradise Park, Mt. Hood, 11 August 1926, Thompson 
1660, Linn Co., Jefferson Park, 16 Aug 1946, Prescott 
s.n., Marion Co., Mt. Jefferson, TIOS R5E S11, 9 Sep 
1992, Roantree, s.n.* (OSC); Mt. Jefferson Wilderness 
Area, SE endof Russel Lake, TIOS R8E S11, 18 Oct 1995, 
Roantree, s.n. (OSC). 


MADRONO, Vol. 49, No. 3, pp. 178-185, 2002 


PLANT ZONATION IN A SHASTA COUNTY SALT SPRING SUPPORTING 
THE ONLY KNOWN POPULATION OF PUCCINELLIA HOWELLII 
(POACEAE) 


LARRY LEVINE 
P.O. Box 4783, Arcata, CA 95518 
1-levine @northcoast.com 


MARY BACCA 
H. T. Harvey & Associates, 3150 Almaden Expressway, Suite 145, San Jose, CA 
mbacca @harveyecology.com 


K. O. FULGHAM 
Department of Rangeland Resources and Wildland Soils, 
Humboldt State University, Arcata, CA 95521-8299 
fulghamk @axe.humboldt.edu 


ABSTRACT 


Three small salt springs adjacent to state highway 299 west of Redding, California, support the only 
known population of the grass, Puccinellia howellii J.1. Davis. The common halophyte grass, Distichlis 
spicata (L.) E. Greene, and Puccinellia each dominate different areas within the springs. In 1991, a 
highway realignment encroached on the Puccinellia, and in partial mitigation, an attempt was made to 
convert a Distichlis-occupied area to Puccinellia. The subsequent re-occupation by Distichlis forced a 
reconsideration of the restoration rationale and methods, and raised concern for the potential of Distichlis 
to replace Puccinellia elsewhere. But a TWINSPAN of systematically chosen samples suggested that the 
two grasses are members of distinct vegetation types associated with different hydrology. Salinity and 
growth monitoring of stands of Puccinellia and Distichlis suggested that the former tends to occupy areas 
continually irrigated by spring discharge, while the latter tends to occupy areas that are less directly 
irrigated, where salt can accumulate during the warm months, but also where precipitation during the 
cool months can lower the salinity enough to temporarily permit the growth of glycophytic annuals. 
Triglochin maritima L. and Juncus bufonius L. were also monitored. Greenhouse tests showed that Puc- 
cinellia is adversely affected by the high salinity typical of the Distichlis areas during summer. The 
association of P. howellii with continual surface flow should be considered when selecting and preparing 
revegetation sites, and when surveying for new populations. 


Key words: Puccinellia howellii, salt spring, salt marsh, Distichlis spicata, vegetation zonation, halophyte, 


endemic 


Adjacent to State Route 299, approximately 32 
km west of Redding, California, near the juncture 
with Crystal Creek Road, and within the boundaries 
of the Whiskeytown Recreational Area, are three 
mineral springs totaling 0.49 hectares (Fig. 1). The 
low grass cover and the bare and rocky areas con- 
trast sharply with the adjacent canyon live oak and 
blue oak woodland. The springs discharge from nu- 
merous points, producing a sheet flow over much 
of the site as the water moves downhill. Salinity 
ranges from 15—35 dS/m (mmho/cm), approximate- 
ly half that of sea water, with a calcium content of 
1—1.8 g/liter. Discharge volume and chemistry re- 
main stable year round (CH2M HILL 1991-1992). 
Though initially alkaline (pH 9—9.6) the water acid- 
ifies (pH 7-5) as it flows away from the discharge 
points (Fulgham et al. 1997), presumably due to the 
influence of decomposing organic matter. Typical 
for this region, precipitation is restricted to a period 
from mid-fall to mid-spring. 

Two halophytic perennial grass species dominate 


the site, usually in separate stands, Puccinellia how- 
ellii J.1. Davis, ““‘Howell’s alkali grass,”’ a cool sea- 
son species endemic to this site, and Distichlis spi- 
cata (L.) E. Greene (including D. stricta (Torr.) 
Rydb.), “‘salt grass,’ a warm season species wide- 
spread in both coastal and inland settings in North 
America. 

Puccinellia howellii was first recognized as a dis- 
tinct species in 1990 (Davis). The closest species 
morphologically is P. pumila (Vasey) A. Hitchc., 
found in coastal marshes from Washington to Alas- 
ka, rarely south to California (Kartesz 1999). Al- 
though commonly referred to as “alkali grass,” 
most Puccinellia are associated with saline habitat 
of neutral pH. Although Puccinellia howellii is in- 
cluded on the California Native Plant Society List 
1B (“‘rare, threatened or endangered in California 
and elsewhere’’), it currently has no state or federal 
legal status (CNPS 2001). 

Puccinellia howellii stands range from sparse to 
dense, and consist either of individual tufts or a 


2002] 


OREGON 


CALIFORNIA 


 WHISKEYTOWN 
ae LAKE 


RED BLUFF 


PACIFIC ° 


OCEAN -% © 10 20 30 40 50 


MILES 


Fic. 1. Location of the study area west of Redding, Shas- 
ta County (Bacca 1995). 


dense, turf-like growth. Plants at maturity can range 
in stature from 2.5 to 20 cm or taller, excluding the 
length of the inflorescences. The tuft form reaches 
the greatest height. The structure of the tufts sug- 
gests that individuals rarely persist longer than sev- 
eral years under field conditions. Triglochin mari- 
tima L. often co-occurs with Puccinellia in the wet- 
ter locations. 

While Puccinellia tends to occupy the central 
portions of the springs, Distichlis is often found at 
the periphery, adjacent to the surrounding non-sa- 
line vegetation. In winter and spring, when Distich- 
lis is dormant, the areas occupied by Distichlis of- 
ten support a sparse to dense cover of the annual 
grass species found outside the springs. At this lo- 
cation, Distichlis appears to spread primarily via 
rhizomes and stolons; little seed production has 
been observed. In contrast, Puccinellia reproduces 
primarily from seed. 

Juncus bufonius L., a glycophytic (non-halo- 
phytic) annual, seasonally occupies some otherwise 
barren areas within the springs. The exotic and po- 
tentially invasive Atriplex rosea L. is present, but 
was uncommon during the data collection portion 
of this study. 

In 1991, a realignment of State Route 299 en- 
croached on the salt springs. The California De- 
partment of Transportation (CalTrans) salvaged 
Puccinellia tufts from the construction zone and 
transplanted them to an adjacent 176 m? area re- 
claimed from a deposit of roadside spoil occupied 
by Distichlis. An aerial photo predating the spoil 
suggested that the area had previously been occu- 
pied by Puccinellia. The site was prepared by re- 
moving approximately % m of soil, which was 
found to be dense with Distichlis rhizomes. After 
transplanting, the Puccinellia tufts quickly became 
established, and at first produced considerable re- 


LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING NT) 


cruitment from seed. However, by the second year, 
Distichlis rhizomes and stolons began to re-enter 
from the periphery. By the third year, Puccinellia 
tufts were dying and not being replaced by seed- 
lings, and by 1998, Distichlis had replaced Pucci- 
nellia in most of the transplant area. This unantic- 
ipated outcome raised questions about the potential 
for Distichlis to replace Puccinellia elsewhere. 
Concurrent with the transplant project, CalTrans 
funded an ecological study of P. howellii, which 
was conducted by the authors between 1993-1995. 
Drawn from that study (Fulgham et al. 1997), this 
paper describes plant zonation within the springs, 
and provides information about the relative habitat 
requirements of the component species. 


METHODS 


Sampling. In May 1993, the visible extent of the 
three springs was systematically surveyed for veg- 
etation analysis. Parallel transects were laid out 
3.05 m apart. Along each transect, an initial sam- 
pling position was selected randomly, and subse- 
quent positions spaced at 3.05 m intervals. Species 
percent cover was estimated using a tripod-mount- 
ed optical point-intercept bar (Cover-Point/ESCO) 
with 20 sighting positions along its 1 m length. At 
each sampling position, the bar was placed perpen- 
dicular to the transect, randomly either left or right. 
Each bar setting constituted a sample. The combi- 
nation of systematic and randomized elements in 
the sampling design ensured a uniform coverage, 
while providing a considerable degree of indepen- 
dence between samples. Because the Distichlis 
growth for the current year had not yet appeared, 
it was surveyed by using the dead growth from the 
previous year, which at the time was still structur- 
ally intact. 

In addition to the collection of species data, the 
surface moisture at each intercept point was re- 
corded in one of four categories, depending on 
whether it appeared dry, moist, saturated (i.e., 
glistening’), or with standing water. 


Vegetation analysis, association. The cover sam- 
ples were analyzed with TWINSPAN (Hill et al. 
1988). Of the 551 samples, only the 394 with =5% 
cover were used. Pseudospecies cut levels were set 
at 5, 15, 33, and 67, and each given equal weight. 
Cover from species found in five or fewer samples 
were combined into a category “‘misc’’. 


Vegetation analysis, moisture. A subset of 235 
vegetation samples was used to compare the sur- 
face moisture associated with each species. As de- 
fined, these were plots with =15% total cover, and 
where one species contributed >50%. In reality, the 
large majority of the samples that met these criteria 
had one species exceeding 66%. To compare the 
relative moisture between samples, the percent cov- 
ers for the moisture categories were weighted and 
summed to form a quasi-continuous moisture in- 
dex. ““Dry”’ cover was weighted by 0.25, “‘moist’’ 


180 


MADRONO 


[Vol. 49 


TABLE 1. TWINSPAN VEGETATION GRouPS, MEAN COVER AND CONSTANCY. Survey area with 551 samples. Analysis 
limited to 394 samples with 25% cover. Tabulation includes species with constancy 25%. n = number of samples. 
Cov = % cover within the group. Con = % constancy. ' Pooled from species “‘misc’’ (species found in <5 samples), 
and ““PLspp”’ (Plantago spp.). 7 Pooled from Bromus diandrus, B. hordaceus, B. rubens, Lolium multiflorum, and 
HOspp (Hordeum spp.). * All cover, including species with <5% constancy. 


TRMA PUHO SCBO/misc DISP/glyco DISP/halo JUBU 
See n=33 n=19 n=19 n=61 n=43 £42.91 =44 
Cov Cov Con Cov Con Cov Con Cov Con Cov Con Cov Con 
Triglochin maritima 1.0 14 100 1 7 
Puccinellia howellii 10.8 1 1D 28 100 6 32 i “23 
Low frequency glycophytes! 0.8 13 OS pee NS) 
Scribneria bolanderi 0.6 8 37 
Annual grasses? es) 1 5 10 63 2 il 
Distichlis spicata 8.6 2 26 54 5 97 29 98 1 16 
Polypogon monspeliensis 0.7 2 tit 6 4] <1 7 
Juncus bufonius 1.4 1 21 D2 20 15 100 
Total cover? De) 15 30 26 66 46 18 


by 0.5, “‘saturated”’ by 0.75, and “‘standing water’”’ 
by 1.0. The resulting composite values ranged from 
25 to 100. A Mann-Whitney test (Minitab 1991) 
was used to compare the moisture index values of 
the Puccinellia and Distichlis samples. 


Species monitoring. Following vegetation anal- 
ysis, groups of four 1 X 2 meter plots were as- 
signed to monitor the biotic and edaphic conditions 
of each of ten vegetation categories of interest, dur- 
ing the period from June 1993 to December 1994. 
To attempt to identify the conditions responsible for 
the variability of the Puccinellia stands, plots were 
assigned to monitor six stature and density combi- 
nations found within the population. The remaining 
sixteen plots were assigned to monitor nearly pure 
stands of the three next largest contributors to cov- 
er, Distichlis, Triglochin, Juncus bufonius, and also 
to barren locations. Initially the plots were random- 
ly assigned to suitable locations identified by the 
systematic sampling; however, this produced a poor 
spatial distribution within and between the springs, 
therefore some plots were reassigned to other ran- 
dom or nearby locations. 

Soil samples were analyzed for texture and rock 
fragment content, and for soluble and exchangeable 
cations, nitrogen, phosphorus, and organic matter. 
Soil moisture, salinity, pH, growth and cover were 
monitored approximately monthly during the peri- 
od. In the case of Puccinellia, additional biotic re- 
sponses were monitored, such as culm and seed 
production and seedling survival. This paper re- 
ports only salinity and cover. Except for salinity, 
and to a lesser extent, soil moisture, little difference 
was noted between the conditions associated with 
the monitored groups. (Among the Puccinellia 
groups, stature was inversely proportional to rock 
fragments.) The plots from the high density/medi- 
um stature ““H3”’’ group will be used to represent 
Puccinellia, because this group seemed to reflect 
conditions favorable for growth, and the vegetation 


in the four plots remained stable for most of the 
duration of the study. 

At each monitoring visit, several soil samples 
were collected from the periphery of each plot and 
pooled to form a composite weighing 300—600 g. 
After air-drying, 100 g portions of the fine-earth 
fraction were used to determine salinity by the sat- 
urated-extract procedure (Roades 1982). The ratio 
of the water content of the saturated paste to that 
of the original soil sample was used to convert the 
electroconductivity to a value presumed equivalent 
to the original field salinity. For specimens contain- 
ing precipitated salt at the time of collection, this 
method produced artificial values exceeding the 
conductivity of a saturated solution (e.g., 226 dS/ 
M at 25°C for NaCl), but nonetheless roughly pro- 
portional to the salt content. 


Greenhouse. In the greenhouse, Puccinellia 
growth was monitored at five levels of salinity, 
ranging from 0 to 80 dS/m, using 4 cm Puccinellia 
plants that had been germinated and reared at 20 
dS/m, a typical field level. The treatment groups 
were adjusted to their target level at a rate of 10 
dS/m per 8 days. Germination was also monitored 
under a similar range of salinities. 


RESULTS AND DISCUSSION 


Vegetation analysis, association. According to 
the point-intercept data, the total vegetative cover 
was 25.2%, with Puccinellia and Distichlis 10.8% 
and 8.6%, respectively. Table 1 lists the cover of 
the major contributors, and summarizes the mean 
cover and constancy for the component species of 
each of the six vegetation groups suggested by 
TWINSPAN. The dendrogram in Figure 2 shows 
the sequence of the divisions. TWINSPAN initially 
segregated two largely monotypic groups from the 
rest of the data, TRMA (Triglochin), and PUHO, 
the latter capturing 94% of the Puccinellia cover. 


2002] 
Level O 

1 

227 PUHO/TRMA 
2 
3 
33 194 
4 19 
TRMA PUHO SCBO/misc 

Fic. 2. 


LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING 181 


394 


167 
123 
104 DISP 44 
61 43 
DISP/glyco DISP/halo JUBU 


TWINSPAN species dendrogram for 394 vegetation samples. Associations: TRMA = Triglochin maritima; 


PUHO = Puccinellia howellii; SCBO/misc = Scribneria bolanderi, Plantago spp., and miscellaneous low frequency 
species; DISP/glyco = higher-density Distichlis spicata sometimes associated with annual grasses; DISP/halo = lower- 
density Distichlis sometimes associated with Puccinellia and Polypogon; JUBU = Juncus bufonius. 


The distinction between TRMA and PUHO, how- 
ever, was not visually apparent in the field, where 
the wetter habitat seemed to support both species 
intermixed. Therefore, it seemed reasonable to re- 
gard the two as a single association, PUHO/TRMA. 

Distichlis was divided between two groups. Ap- 
proximately two-thirds of the cover was placed in 
a group that we labeled DISP/glyco, characterized 
by higher density Distichlis, and often accompanied 
by annual grasses and other glycophytes that are 
active during the rainy season. Puccinellia was no- 
tably absent. During the vegetation survey, we re- 
garded the annual grasses as presumably salt-tol- 
erant contaminants, but we revised this view later 
as we observed Distichlis emerge amid the brown 
stalks of annual grasses in areas beyond the as- 
sumed boundary of the salt spring. In the down- 
slope portions of these peripheral and sometimes 
disjunct zones there were seepages that had not 
been visible. The accumulating mineral stains and 
evaporate indicated that during summer, after the 
cessation of precipitation, salt water was able to 
reach the surface. We inferred that in the central 
Spring area, the presence of glycophyte grasses 
amid the Distichlis might also indicate a salinity 
regime that alternates between fresh in winter and 
saline in summer. 

The remaining % of the Distichlis, labeled DISP/ 
halo, had a lower mean density and a smaller gly- 
cophytic component. Puccinellia and/or Polypogon 
monspeliensis (L.) Desf. were sometimes present. 
Although the latter is not usually regarded as a hal- 


ophyte, it will germinate simultaneously with Puc- 
cinellia when seed of both species are irrigated di- 
rectly by salt spring surface flow (Levine personal 
observation). 

The primarily monotypic JUBU, capturing most 
of the Juncus bufonius, also had a small Puccinellia 
component. The heterogeneous SCBO/misc., in- 
cluding nearly all the Scribneria bolanderi (Thur- 
ber) Hackel and most of the miscellaneous low- 
frequency glycophytes, consisted of the remaining 
samples. 


Vegetation analysis, moisture. The vegetation 
sampling was performed well after the cessation of 
seasonal precipitation, when the surface moisture 
primarily reflected the influence of the spring dis- 
charge, but before the evaporative effects of sum- 
mer heat. Among the 235 samples predominated by 
a single species, the wetter samples were common- 
ly dominated by Puccinellia or Triglochin, the drier 
by Juncus and Scribneria, and Distichlis was inter- 
mediate (Fig. 3). A Mann-Whitney test showed that 
the Puccinellia samples had significantly greater 
surface moisture values than the Distichlis (P < 
0.0001). 


Species monitoring. Possible insights into the re- 
lationship between growth and the salinity regime 
may be provided by examining the data from the 
monitoring plots for the four species (Fig. 4a—h). 
Although the sample size is small, the data suggest 
that areas occupied by different species may also 


182 


MADRONO 


[Vol. 49 


samples with one species predominant 
surface moisture histograms 


f=] SCBO J PUHO[__]DISP 
JUBU WY TRMA 


sample count 


25 35 45 55 
moisture index center mark 


Fic. 3. 


65 15 85 95 


Histogram of surface moisture for the 235 vegetation samples 215% cover, where one species contributed 


more than 50% (the large majority contributing >66%). The dominant species are indicated by the acronyms defined 
in Figure 2. Values range from 25 = “dry” to 100 = “standing water’’. 


differ in the annual pattern of salinity, and by in- 
ference, the amount of irrigation. 

In the Puccinellia and Triglochin plots (Fig. 4a, 
c), saline conditions occurred year-round, and rose 
only moderately during the summer. Apparently, 
the volume of irrigation was sufficient to minimize 
the seasonal influence of precipitation or evapora- 
tion. In contrast, salinity in the Juncus plots and 
most of the Distichlis plots (Fig. 4e, g) was low 
from fall to spring, and became high during the 
summer. The exception was Distichlis plot #3, 
which remained saline through the winter. This plot 
was selected for monitoring because of the atypi- 
cally short stature and low density, and the adja- 
cency of robust Puccinellia, perhaps suggestive of 
the TWINSPAN group, DISP/halo. 

The growth cycles of the four species appear to 
track the change in salinity. Puccinellia (Fig. 4b) 
has a phenology similar to the glycophytic grasses 


of the region. Growth occurs during the cool 
months, and seed is produced before a summer dor- 
mancy. However, regeneration by germination or 
resprouting often begins in advance of the fall 
rains. Triglochin (Fig. 4d) remains active through 
the summer, but declines during the cold months. 
Its succulent foliage is susceptible to damage from 
freezing. Distichlis (Fig. 4f) typically dies back 
completely during the cold months, but its prefer- 
ence for warm weather and its high salt tolerance 
(Ungar 1974; Hansen et al. 1976) enables it to grow 
on sites that during the summer become too saline 
for Puccinellia. Salt build-up in the Juncus bufon- 
ius plots (Fig. 4h) was more extreme than in the 
Distichlis plots. An early-flowering annual, Juncus 
can complete its life cycle within a short period of 
low salinity during winter and spring. 

Though the ability of Distichlis to tolerate high 
salinity is by no means unique, its often greater 


=> 


Fic. 4. Seasonal salinity and cover in sets of monitoring plots. a, b) H3-Puccinellia = high density, medium stature 
Puccinellia howellii. Reported conductivities are the saturated extract values adjusted for field moisture. Samples with 
accumulated salt produced artificial conductivity values that exceeded the level of a saturated solution (e.g., 226 dS/m 
at 20°C for NaCl), but which remained proportional to the salt content. 50 dS/m is approximately equivalent to coastal 
sea water. The unconnected cover values labeled “‘green + brown,” and “‘brown”’ were measured at the end of the 
previous growing season. 


LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING 


Species Monitoring Plots - June 1993 to December 1994 


2002] 
a_H8-Puccinellia Estimated Salinity 
100 
—= PLOT-1 ~*~ PLOT-2 ~*~ PLOT-3 --a- PLOT-4 

80 
60 

£ 

” 

me) 
40 
20 


dS/m 


dS/m 


dS/m 


Ja4-94 Mr6é  My1 JI3 Se4 Oc 29 


fe) 
Jn29-93 Se5 Nod 


Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oct No22 


C __Triglochin Estimated Salinity 


100 


80 


60 


40 


0 
Jn29-93 Se5 NoS5 Ja4-94 Mr6 My1 JI3 Se4 Oc29 
Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oc1 No22 


e Distichlis Estimated Salinity 


400 D-1 121 D-3 103 


80 
60 


40 


Ja4-94 M Myt JIS Se4 Oc29 
No22 


ie) 
Jn29-93 Sed No5 
Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oct 


Juncus Estimated Salinity 
+2 491 J-2 983 J-1 489 


100 


80 


40 


Se4 Oc29 


ie) 
Jn29-93 Sed NoS Ja4-94 Mré6 My1 Ji3 


Au4 Oc2 De4 Fe4 Ap3 Jn7 Au2 Oc1 No22 


b  H8-Puccinellia % Green Cover 
100 


80 
x 


(green+ 
brown) 


60 


40 


0 
Je24-93 Se10 No15 Fe23-94 My3 Jy13 Se22 De21 
Au4 Oc7 De31 Mr31 Jn7 Au15 No3 


d Triglochin % Green Cover 


100 


80 


60 


40 


eres 


Fe23-94 My3 Jy13 Se22 De21 


ie) 
Je24-93 Se10 No15 


Au4 Oc7 De31 Mr31 Jn7 Au15 No3 
f Distichlis % Green Cove 


100 — — 
80 


60 


40 


0 = " Pres = “- = - 
Je24-93. Se10  Noi5 Fe23-04 My3 Jy13 Se22.—s- De? 


Au4 Oc7 De31 Mr31 Jn7 Au15 No3 
h Juncus % Green Cover 


100 
80 
60 

=< 


40 
(brown) 


Jy13 Se22 De21 


Fe23-94 My3 
Au4 Oc7 De31 Mr31 Jn7 Au15 No3 


6 . 
Je24-93 Se10 No15 


183 


184 MADRONO 


capacity compared with other species has been not- 
ed in other settings. For example, in New England, 
Distichlis rapidly re-colonized bare patches of 
coastal salt marsh that were created by shading 
from wrack (Bertness et al. 1992; Shumway 1995). 
These areas then become hypersaline due to in- 
creased evaporation. Their studies suggest that Dis- 
tichlis can invade and occupy patches too saline for 
the establishment and persistence of discrete indi- 
viduals by transporting along its runners water and 
carbon from ramets outside the hypersaline zone. 

The distribution of Distichlis within the salt 
springs seems to reflect not only its relatively high 
salt tolerance, but also a limited capacity for im- 
mersion. On both coasts Distichlis is a component 
of high marsh vegetation, where it is not subject to 
daily flooding (Macdonald and Barbour 1974; Sil- 
berhorn 1982; Macdonald 1988). In inland settings 
it is associated with only moderate moisture (Ungar 
1974). The detrimental effect of inundation on Dis- 
tichlis, and the associated abiotic factors, was dem- 
onstrated in a Louisiana tidal marsh undergoing 
vegetation decline associated with relative sea level 
rise. Webb et al. (1995) transplanted sod blocks of 
four species level with the marsh surface and raised 
20 cm higher, equivalent to the level of a nearby 
healthy marsh. All species produced less biomass 
at the ambient level, but the greatest reduction was 
in those typically associated with high marsh hab- 
itat, a twenty-fold difference in the case of Distich- 
lis. The poor growth at the ambient level was as- 
sociated with negative substrate redox potential at 
both 2 cm and 15 cm depth. The elevated plantings 
had uniformly higher potentials, with oxidized con- 
ditions at 2 cm. 

The comparatively larger number of plots as- 
signed to monitor Puccinellia, and their division 
into stand categories, helped provide examples of 
the variation in growing conditions. The timing of 
Puccinellia germination appeared related to sum- 
mer salinity. Seed in locations directly inundated 
by surface flow germinated as early as mid-sum- 
mer, while areas with high summer salinity pro- 
duced their major germination pulse in November 
or later. Locations with visible salt accumulation, 
including those near the evaporative edge of sur- 
face flows, seemed more prone to seedling mortal- 
ity, and to the failure of tufts to regenerate in the 
fall. 

The two Puccinellia plots located in the trans- 
plant area provided an example of salinity-related 
mortality, and evidence that the pattern of seepage 
can shift over time. During the first summer of 
monitoring, these high-density/high-stature stands 
had a measured salinity similar to Distichlis plots, 
and visible salt accumulation. At the same time, the 
plots were being invaded by Distichlis. The follow- 
ing fall, the Puccinellia tufts did not resprout, and 
though recruitment was heavy, little germination 
was recorded within the plot itself. In effect, the 
Puccinellia had regenerated, but had shifted in po- 


[Vol. 49 


sition, and was no longer centered within the plot 
boundaries. Toward the end of the monitoring pe- 
riod, plot H3-3 provided a similar example of non- 
regeneration due to shifting irrigation (Fig. 4b), 
however, the visible salt accumulation within the 
plot was not reflected in the soil samples collected 
from the periphery. Dense germination occurred 
outside and adjacent to the plot boundary, where 
surface flow was evident. 


Greenhouse. The salinity preferences of Pucci- 
nellia are suggested by its behavior in the green- 
house. The greatest growth was seen in the group 
that remained at 20 dS/m. Growth at 60 dS/m was 
stunted, and 80 dS/m proved lethal. In the field, 
summer levels of 60 dS/m and higher were com- 
monly found in the Distichlis and Juncus plots, and 
also in the plots of low-density/low-stature Pucci- 
nellia. The amount and rate of germination was in- 
versely proportional to salinity, consistent with field 
observations that germination is delayed in areas 
that that accumulated salt during the previous sum- 
mer. 


CONCLUSIONS 


The large majority of the Puccinellia cover was 
found in monotypic stands. The Distichlis stands 
were divided into two groups. The larger portion, 
two-thirds of the total cover, contained no Pucci- 
nellia, but often supported annual grasses during 
the cool season. The presence of Puccinellia in the 
remaining one-third and in the Juncus stands, might 
indicate locations where habitat conditions were in- 
termediate or in flux. 

The vegetation at the salt springs can be divided 
into two groups, based on the relative influence of 
spring discharge. A saline-winter/saline-summer di- 
vision, typified by Puccinellia and Triglochin, tends 
to be wet year-round. The habitat remains saline 
during the rainy season, excluding glycophytes dur- 
ing the period of active Puccinellia growth. Salinity 
rises only moderately during the summer, permit- 
ting late-summer to early-fall germination by Puc- 
cinellia. A fresh-winter/saline-summer division, 
typified by Distichlis and Juncus, is more likely to 
occur at the periphery or at elevated areas within 
the springs, where the influence of the discharge is 
less direct. During the summer, wicking and evap- 
oration promote hypersalinity, but winter precipi- 
tation dilutes the salt sufficiently to permit the 
growth of glycophytes. The areas occupied by these 
vegetation types may shift somewhat over time in 
response to changes in the pattern of seepage and 
surface flow. 

This study concludes that a major requirement 
for Puccinellia, apart from suitable salinity, is a 
volume of spring discharge sufficient to maintain 
salinity during both the wet and dry seasons. In this 
region of relatively high precipitation, few salt 
springs may meet this requirement, but qualifying 
sites merit attention because they may harbor un- 


2002] 


documented populations or could become candidate 
sites for outplanting. The evidence that Puccinellia 
and Distichlis have different hydrologic preferences 
provides useful information for managing the only 
known population of Puccinellia in the presence of 
Distichlis. For instance, Distichlis might be exclud- 
ed by lowering the grade sufficiently so that surface 
flow occurs year-round. This approach is currently 
being explored by CalTrans within the area of their 
transplant project. 


ACKNOWLEDGMENTS 


We wish to extend our appreciation to Sharon Stacey 
of CalTrans District 2, Redding, and to Gretchen Ring and 
Jennifer Gibson of the Whiskeytown National Recreation- 
al Area, for their assistance in this study, and for their 
continued commitment to the conservation of Puccinellia 
howellii. 


LITERATURE CITED 


Bacca, M. 1995. Control strategies to inhibit saltgrass 
(Distichlis spicata) encroachment upon Howell’s al- 
kali grass (Puccinellia howellii) at an inland miner- 
alized spring area in Shasta County, California. M.S. 
thesis. Humboldt State University, Arcata, CA. 

BERTNESS, M. D., L. GOUGH, AND S. W. SHUMWAY. 1992. 
Salt tolerances and the distribution of fugitive marsh 
plants. Ecology 73:1842-—1851. 

CALIFORNIA NATIVE PLANT SOCIETY (CNPS). 2001. Inven- 
tory of rare and endangered plants of California, sixth 
edition. Rare Plant Scientific advisory Committee, D. 
Tibor, convening editor. California Native Plant So- 
ciety, Sacramento, CA. 

CH2M HILL, 1991-1992. Reports of inorganic analyses 
of water samples supplied by CalTrans, lab numbers 
31306, 32495, 32650, 32818, 32948, 33641. Avail- 
able at CalTrans, Redding, CA. 

Davis, J. I. 1990. Puccinellia howellii (Poaceae), a new 
species from California. Madrono 37:55-—58. 

FULGHAM, K. O., L. LEVINE, AND M. Bacca. 1997. Aut- 
ecological Study of Puccinellia howellii: contract 
02E326 final report. Available at CalTrans, Redding, 
CA. 

HANSEN, D. J., P- DAYANANDAN, P. B. KAUFMAN, AND J. D. 


LEVINE ET AL.: PLANT ZONATION IN SHASTA COUNTY SALT SPRING 185 


BROTHERSON. 1976. Ecological adaptations of salt 
marsh grass, Distichlis spicata (Gramineae) and en- 
vironmental factors affecting its growth and distri- 
bution. American Journal of Botany 63:635—650. 

Hitt, M. O., C. T: FE TER BRAAK, O. E R. VAN TONGEREN, 
AND J. B. BirKs. 1988. TWINSPAN. Modification by 
J. B. Birks of a FORTRAN program originally writ- 
ten by M.O. Hill, 1979. Available from U.K. Mirror 
Service. September 22, 2002. http://www.mirror.ac. 
uk/collections/Mensa-micros/local/msdos/science/ 
cornell_ecology/cep2.zip 

KARTESZ, J. T. 1999. A synonymized checklist and atlas 
with biological attributes for the vascular flora of the 
United States, Canada, and Greenland. /n J. T. Kartesz 
and C. A. Meacham (eds.), Synthesis of the North 
American Flora, Version 1.0. North Carolina Botan- 
ical Garden, Chapel Hill, NC. 

MACDONALD, K. B. 1988. Coastal salt marsh. Pp. 263—294 
in M. G. Barbour and J. Major (eds.), Terrestrial veg- 
etation of California. California Native Plant Society, 
Sacramento, CA. 

AND M. G. BARBourR. 1974. Beach and salt marsh 

vegetation of the North American Pacific coast. Pp. 

175-235 in R. J. Reimold and W. H. Queen (eds.), 

Ecology of halophytes. Academic Press, New York, 

NY. 


MIniTaBs. 1991. MINITAB version 8.2, statistical software. 
Minitab, Inc., State College, PA. 

RoaDEs, J. D. 1982. Soluble salts. Pp. 167—180 in A. L. 
Page (ed.), Methods of soil analysis, Part 2, Chemical 
and microbiological properties, 2nd ed. American So- 
ciety of Agronomy, Inc., and Soil Science Society of 
America, Inc., Madison, WI. 

SHUMWAY, S. W. 1995. Physiological integration among 
clonal ramets during invasion of disturbance in a New 
England salt marsh. Annals of Botany 76:225—233. 

SILBERHORN, G. M. 1982. Common plants of the Mid- 
Atlantic coast. Johns Hopkins University Press, Bal- 
timore, MD. 

UnGar, I. A. 1974. Inland halophytes of the United States. 
Pp. 235-306 in R. J. Reimold and W. H. Queen (eds.), 
Ecology of halophytes. Academic Press, New York, 
NY. 

WEBB, E. C., I. A. MENDELSSOHN, AND B. J. WILSEY. 1995. 
Causes of vegetation dieback in a Louisiana salt 
marsh: a bioassay approach. Aquatic Biology 51: 
281-289. 


MADRONO, Vol. 49, No. 3, pp. 186-188, 2002 


SPHAGNUM BALTICUM IN A SOUTHERN ROCKY MOUNTAIN IRON FEN 


DAVID J. COOPER 
Department of Earth Resources, Colorado State University, 
Fort Collins, CO 80523 
dcooper @rm.incc.net 


RICHARD E. ANDRUS 
Environmental Studies Program, Binghamton University, Binghamton, NY 13902 


CHRISTOPHER D. ARP 
Department of Biology, Utah State University, Logan, UT 84322 


Sphagnum balticum (Russow) C. Jensen is a 
widespread Holarctic peat moss of raised and blan- 
ket bogs, occurring partly or completely submerged 
in pools, in floating mats and on hummocks. It is 
known from arctic and subarctic Greenland, North 
America (Crum and Andersen 1981; Crum 1984), 
Scandinavia, the British Isles, Russia and northern 
China. It is one of the rarest plants in the United 
Kingdom (UK Biodiversity Group 1998). Until 
now, the known range of S. balticum in North 
America extended south to about 57 degrees north 
latitude in western and central Canada and it is un- 
known in the lower 48 United States. Therefore, it 
is striking to discover this species in the San Juan 
Mountains in southwestern Colorado disjunct by 
more than 2000 km from the main range of the 
species in northern Canada and Alaska (Sphagnum 
balticum, Cooper #2281, COLO, BING). 

During a regional analysis of iron fens, we found 
Sphagnum balticum in the Chattanooga iron fen 
(2990 m elevation, Latitude 37°50'N, Longitude 
107°43’W) south of Red Mountain Pass where it is 
the most abundant moss in shallow pools, growing 
with the sedges Carex aquatilis Wahlenberg and C. 
utriculata Boott. It is easily separated in the field 
from the only other Sphagnum in section Cuspidata 
present in the area, S. angustifolium (Russow) C. 
Jens. by a laxer habit and conspicuous lingulate and 
spreading stem leaves. The fen is almost complete- 
ly covered by Sphagnum mats and hummocks, with 
abundant S. angustifolium, S. russowii Warnst., S. 
fuscum (Schimp.) Klinggr. and S. fimbriatum Wils. 
(nomenclature follows Crum 1984). We found S. 
girgensohnit Russow, another new Colorado re- 
cord, in a different iron fen. 

The water and soil in most Sphagnum-dominated 
peatlands is acidic. The origin of the acids has been 
linked to Sphagnum cation exchange capacity (Cly- 
mo 1963; Clymo and Hayward 1982; Glaser 1987), 
atmospheric acid deposition (Gorham 1967), bio- 
logical uptake of nutrient cations by plants (Mitsch 
and Gosselink 1994), and the buildup of organic 
acids by decomposition (Gorham et al. 1984). 
These processes control autochthonous production 
and accumulation of acids in ombrogenous peat- 


lands (bogs). Fens in the Rocky Mountains, how- 
ever are soligenous (formed on slopes) or limno- 
genous (formed on lake margins), and are season- 
ally flushed with abundant snow melt water (Coo- 
per and Andrus 1994). Because strong flushing 
occurs, the pH of surface and ground water is con- 
trolled by the chemistry of watershed surface and 
ground waters, not autochthonous acid production. 

Warm and dry summers with long rainless peri- 
ods and a large evaporative demand are character- 
istic of the continental climate of the southern 
Rocky Mountains. Peatlands occur only where a 
continuous supply of ground water maintains pe- 
rennially saturated soil conditions. The most com- 
mon peatlands are transitional fens with slightly 
acid waters in mountain ranges of granite, rhyolite 
or metamorphic bedrock (Cooper and Andrus 1994) 
and extreme rich fens with circumneutral or basic 
waters in areas of, limestone, dolomite, shale, and 
basalt bedrock (Cooper 1996). 

Colorado iron fens appear to be geochemically 
unique and occur in areas with highly mineralized 
outcrops, such as occur on Red Mountain Pass. 
While iron fens are hydrologically similar to other 
Colorado fens, with ground water discharge per- 
manently saturating valley bottom wetlands, the 
source water flows through fractured iron pyrite- 
rich bedrock and talus, oxidizing pyrite and form- 
ing sulfuric acid. 

Chattanooga iron fen surface water has a pH 
from 3.8 to 4.4, similar to bogs and poor fens in 
northern Minnesota (Glaser 1987). Acid drainage 
from historical metal mines in Colorado produces 
similar low pH waters, and pollutes some Colorado 
wetlands (Arp et al. 1999), however iron fens are 
natural ecosystems. Although geochemical process- 
es external to the peatland generate the acids in iron 
fens, the fen flora is limited to acid tolerant species 
that also occur in poor fens and bogs. 

Bog waters have low ion concentrations because 
precipitation is the primary water supply. In con- 
trast, iron fens have high ion concentrations be- 
cause sulfuric acid produced in the watershed 
leaches and mobilizes metal and base cations from 
rock. Concentrations of Ca?* in Chattanooga fen 


2002] 


surface water range from 14—20 mg/liter, and SO, 
averages 117 mg/liter. Ca** concentration in bogs 
range from 0.5 to 2.0 mg/liter (Glaser et al. 1981, 
1990), in poor fens from 2.0—5.0 mg/liter (Sjérs 
1963: Glaser et al. 1981), transitional fens in the 
Rocky Mountains from 2.0—7.0 mg/liter (Cooper 
and Andrus 1994), and rich fens from 5.0—30.0 mg/ 
liter (Glaser et al. 1981, 1990). Thus, the Chatta- 
nooga iron fen has Ca** concentration most similar 
to a rich fen. High iron concentrations (0.2—6.3 mg/ 
liter) precipitate onto fen organic matter forming 
terraced bog iron ore (limonite) deposits (Harrer 
and Tesch 1959), which are characteristic of iron 
fens. Limonite terraces perch the water table and 
form extensive networks of pools and ponds that 
are uncommon in most other Colorado fens. 

The presence of plants, such as Sphagnum bal- 
ticum, that are widely disjunct from their main 
ranges has always intrigued biogeographers. Did 
people, migratory birds or other vectors transport 
these plants? Are they the last vestiges of plant 
populations that established in the southern Rocky 
Mountains from the north during the Pleistocene as 
suggested by Hooker and Gray (1880)? Or do they 
reflect much older plant evolutionary and devel- 
opmental patterns as suggested by Weber (1965)? 

It is unlikely that animals, people or wind would 
have dispersed Sphagnum balticum, and Colorado 
has numerous other boreal montane bryophytes and 
lichens that exist in tiny populations disjunct from 
their main ranges in the boreal and subarctic re- 
gions of North America. Examples include Cladina 
stellaris (Opiz) Brodo found in a few 10 m? patches 
on the margin of a fen in the Tarryall Range, 
Sphagnum platyphyllum (Braithwaite) Warnstorf in 
two small wetlands in the Sawatch Range, S. con- 
tortum Schultz in one small wetland in the Front 
Range, Paludella squarrosa (Hedwig) Bridel in a 
couple of tiny alpine wetlands on Guanella Pass, 
and Scorpidium scorpioides (Hedwig) Limpricht in 
a few acres of calcareous fen in South Park (Cooper 
1991; Weber and Wittmann 1996). The large num- 
ber of species persisting as localized populations, 
many found in only one fen, or on one mountain 
slope, suggests that these populations are the ves- 
tiges of what may have been wider distributions 
along the Cordillera that are much older than the 
Pleistocene. Many Colorado fens have basal '*C 
dates of 10,000—12,000 years BP (Cooper 1990; 
Chimner and Cooper 2002), with peat body initia- 
tion soon after the melting of Pleistocene glaciers. 
Boreal montane species could have found stable re- 
fugia in the southern Rocky Mountains and per- 
sisted through the Holocene. 

It is striking that new populations of widespread 
and well-known boreal montane plants are still be- 
ing found in Colorado, a state with modern floristic 
manuals (e.g., Weber and Wittmann 2001), numer- 
ous botanists, and a rich history of botanical inves- 
tigation. It indicates that additional species are like- 
ly to be discovered. Rare plants, such as S. balti- 


COOPER ET AL.: Sphagnum balticum in the Sourthern Rocky Mountains 187 


cum, occur in very specialized habitats, such as iron 
fens, that are difficult to find. Careful inventories 
are necessary to make certain that logging, mining, 
and recreation and water developments do not in- 
advertently destroy the only populations of species 
occurring in the southern Rocky Mountains, and 
that may have existed here for many millennia. It 
also indicates the sensitivity of these habitats and 
populations to hydrologic variability that could ac- 
company human induced climate changes. Since 
populations of these species are very small and the 
circumstances of survival of each species are likely 
to be unique, there is no chance for species replen- 
ishment. 


ACKNOWLEDGMENTS 


This research was funded by a grant from the State of 
Colorado Natural Areas Program. We thank Janet Coles, 
Kathy Carsey, Gay Austin and Paula Lehr for their assis- 
tance in the field, and for Dr. W. A. Weber for discussions 
on biogeography. The identification of S. balticum was 
verified by Dr. Kjell Flatberg, University of Trondheim, 
Norway, and we thank him for his assistance. We also 
thank three reviewers for comments, which improved this 
manuscript. 


LITERATURE CITED 


Arp, C. D., D. J. COOPER, AND J. D. STEDNICK. 1999. The 
effects of acid rock drainage on Carex aquatilis leaf 
litter decomposition in Rocky Mountain fens. Wet- 
lands 19:665—674. 

CLymo, R. S. 1963. Ion exchange in Sphagnum and its 
relation to bog ecology. Annals of Botany 27:309-— 
324. 

CLymo, R. S. AND P. M. HAywarpD. 1982. The ecology of 
Sphagnum. Pp. 229-289 in A. J. E. Smith (ed.), Bry- 
ophtye ecology. Chapman and Hall, London, U.K. 

CHIMNER, R. A. AND D. J. COOPER. 2002. Modeling carbon 
accumulation in fens using the century ecosystem 
model. Wetlands 22:100—110. 

Cooper, D. J. 1990. The ecology of wetlands in Big 
Meadows, Rocky Mountain National Park, Colorado: 
the correlation of vegetation, soils and hydrology. Bi- 
ological Report 90(15). U.S. Department of the In- 
terior, Fish and Wildlife Service, Washington, DC. 

. 1991. The habitats of three boreal fen mosses 

new to the southern Rocky Mountains of Colorado. 

The Bryologist 94:49—50. 

. 1996. Water and soil chemistry, floristics and 

phytosociology of the extreme rich High Creek fen, 

in South Park, Colorado, U.S.A. Canadian Journal of 

Botany 74:1801—1811. 

AND R. E. ANpDRuS. 1994. Patterns of vegetation 
and water chemistry in peatlands of the west-central 
Wind River Range, Wyoming, U.S.A. Canadian Jour- 
nal of Botany 72:1586—1597. 

Crum, H. 1984. North American flora: Sphagnopsida, 
Sphagnaceae. New York Botanical Garden, New 
York, NY. 

AND L. E. ANDERSEN. 1981. Mosses of eastern 
North America. Columbia University Press, New 
York, NY. 

GLASER, P. H. 1987. The ecology of patterned boreal peat- 
lands of northern Minnesota: a community profile. 


188 MADRONO 


Biological Report 85(7.14). U.S. Department of In- 

terior, Fish and Wildlife Service, Washington, DC. 

, G. A. WHEELER, E. GORHAM, AND H. E. WRIGHT, 

JR. 1981. The patterned mires of the Red Lake Peat- 

land, northern Minnesota: vegetation, water chemis- 

try, and landforms. Journal of Ecology 69:575-—599. 

, J. A. JANSSENS, AND D. I. SIEGEL. 1990. The re- 
sponse of vegetation to chemical and hydrological 
gradients in the Lost River Peatland, northern Min- 
nesota. Journal of Ecology 78:1021—1048. 

GORHAM, E. 1967. Some chemical aspects of wetland 
ecology. Committee on Geotechnical Research, Na- 
tional Research Council on Canada, No. 90, pp. 20— 
38. 


, 8. E. BAYLEY, AND D. W. SCHINDLER. 1984. Eco- 
logical effects of acid deposition on peatlands: a ne- 
glected field in “‘acid-rain” research. Canadian Jour- 
nal of Fisheries and Aquatic Science 41:1256—1268. 

HARRER, C. M. AND W. J. TESCH, JR. 1959. Reconnaissance 
of iron occurrences in Colorado. U.S. Department of 


[Vol. 49 


the Interior, Bureau of Mines, Information Circular 
7918. 

HOOKER, J. D. AND A. GRAY. 1880. The vegetation of the 
Rocky Mountain region and a comparison with that 
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Territories 6:1—62. 

Mitscu, W. J. AND J. G. GOSSELINK. 1994. Wetlands, 2nd 
ed. Van Nostrand Reinhold, New York, NY. 

Syors, H. 1963. Bogs and fens on Attawapiskat River, 
northern Ontario. Bulletin of National Museum of 
Canada 186:45—103. 

UK Bropiversity Group. 1998. Tranche 2 action plans, 
plants and fungi. English Nature, London, U.K. 
WEBER, W. A. 1965. Plant geography of the southern 
Rocky Mountains. Pp. 453—468 in H. E. Wright, Jr. 
and David G. Frey (eds.), The Quaternary of the Unit- 
ed States. Princeton University Press, Princeton, NJ. 

AND R. C. WITTMANN. 2001. Colorado flora: east- 

ern slope, revised edition. University Press of Colo- 

rado, Niwot, CO. 


MADRONO, Vol. 49, No. 3, pp. 189-192, 2002 


EVIDENCE OF A NOVEL LINEAGE WITHIN THE PONDEROSAE 


ANN M. PATTEN! AND STEVEN J. BRUNSFELD? 
Department of Forest Resources, University of Idaho, Moscow, ID USA 83843 


ABSTRACT 


Phylogenetic analysis of the DNA of a putative portion of the nuclear NADH-specific nitrate reductase 
gene revealed the existence of a Pinus jeffreyi lineage that gave rise to P. washoensis and the North 
Plateau race of P. ponderosa var. ponderosa. These data are consistent with Lauria’s (1991) hypotheses 
that the North Plateau race is genetically distinct from the other races of the species, and that this race 


should be considered conspecific with P. washoensis. 


Pinus subsection Ponderosae is an economically 
important and well-represented group across much 
of western North America. However, the species in 
this group have been the source of considerable tax- 
onomic disagreement (Lauria 1991, 1997; Kral 
1993; Rehfeldt 1999). Taxonomic treatments and 
inferences about the evolution of the Ponderosae 
have been based on a large number of different data 
sets, including quantitative morphological charac- 
ters (Peloquin 1984; Rehfeldt et al. 1996; Rehfeldt 
1999), terpene chemistry (Mirov 1961; Smith 1964, 
1967, 1977; von Rudloff and Lapp 1991), isozymes 
(Niebling and Conkle 1990), crossability (Critch- 
field 1984), provenance analysis (Wells 1964; van 
Haverbeke 1986), and the fossil record (Stockey 
1984; Axelrod 1986). Some of these studies have 
produced conflicting data or additional uncertainty 
because of the omission of key species or varieties 
(Lauria 1991). 

The primary objective of this limited study was 
to test two of Lauria’s hypotheses: (1) the North 
Plateau race of ponderosa pine (Pinus ponderosa 
Douglas ex Lawson and C. Lawson) is a distinct 
genetic entity relative to the other races of this spe- 
cies (Lauria 1991); and (2) Washoe pine (Pinus 
washoensis Mason and Stockwell) and the North 
Plateau race of ponderosa pine are conspecific 
(Lauria 1997). Sampling thus focused primarily on 
the five geographic races of ponderosa pine (Smith 
1977; Conkle and Critchfield 1988), Washoe pine, 
and Jeffrey pine (Pinus jeffreyi Grev. and Balf.). 
Three other members of Ponderosae, Pinus arizon- 
ica Engelm. and Martinez, Pinus durangensis Mar- 
tinez, and Pinus engelmannii Carr., were included 
for comparison of relative genetic divergence. Pi- 
nus coulteri D. Don from subsection Sabinianae 
was also included because of its high crossability 
with Jeffrey pine (Zobel 1951) and the close rela- 
tionship between subsections Sabinianae and Pon- 
derosae exhibited in a chloroplast DNA analysis 


' Current address: The Institute of Biological Chemis- 
try, Washington State University, Pullman, WA 99164- 
6340. 


* Corresponding author. E-mail: sbruns @uidaho.edu. 


(Kupkin et al. 1996). Pinus contorta Dougl. ex 
Loud. was included as an outgroup. Most taxa were 
represented by two samples; however four samples 
from the North Plateau race and one sample from 
each of the three recognized populations of Washoe 
pine were included (Table 1). This study was con- 
ducted concurrently with a larger ecological genet- 
ics study (Rehfeldt 1999) of Washoe pine, Jeffrey 
pine, and ponderosa pine. 

DNA was isolated from needle tissue according 
to Lodhi et al. (1994). PCR products were gener- 
ated using primers designed to target a region cod- 
ing for the two hinges that connect the internal 
heme domain to the amino and carboxy terminal 
domains of the nuclear NADH-specific nitrate re- 
ductase (NADH-NR) gene (Zhou et al. 1995; Patten 
1999). Manual sequencing of the PCR product was 
conducted using the USB Sequenase Kit (Amer- 
sham). A BLAST search of the GenBank data base 
did not reveal a match between our sequences and 
those reported as NADH-NR. However, the se- 
quence and structure of nitrate reductase is not 
known for any gymnosperm. Furthermore, the PCR 
primers are targeted to an area known to contain 
introns (Zhou and Kleinhofs 1996), lessening the 
chance of similarity to pine. Thus, to be conser- 
vative, the sequences might best be considered 
anonymous, although the parsimonious distribution 
of synapomorphies strongly suggests the sequences 
are orthologous (Fig. 1). Paralogous PCR products 
would likely exhibit a more random distribution 
with respect to taxonomic classification and geog- 
raphy (see below). 

Because there were variable amounts of missing 
data at the termini of the PCR products, a 287-bp 
fragment was used in the final phylogenetic anal- 
ysis. The sequences were analyzed using PAUP 
version 3.1.1 (Swofford 1993). The branch and 
bound exact algorithm was used and two most par- 
simonious trees were recovered. These differed 
only in the resolution of one dichotomy. Both trees 
had a length of 25 and a consistency index of 0.90. 
A consensus tree was constructed and all branches 
of zero length were collapsed (Fig. 1). 

This phylogenetic analysis offers novel insights 


190 MADRONO [Vol. 49 
TABLE 1. 27 SAMPLES USED IN PHYLOGENETIC ANALYSIS OF NORTH AMERICAN PONDEROSAE. 

Sample Location Collection GenBank # 
P. contorta Latah Co., ID A. Patten #95-14 AF06764 
P. coulteri/A Black Mtn., CA A. Patten #95-10 U77801 
P. coulteri/B Black Mtn., CA A. Patten #95-9 U77802 
P. jeffreyi/A Nevada Co., CA USFS U77803 
P. jeffreyi/B Nevada Co., CA USFS U77804 
P. ponderosa/NP-ID Idaho Co., ID A. Patten #95-4 U77805 
P. ponderosa/NP-OR Benton Co., OR G. Rehfeldt #94-9 U77810 
P. ponderosa/NP-WA1 Kittitas Co., WA G. Rehfeldt #94-7 U77806 
P. ponderosal[NP-WA2 Puget Sound, WA G. Rehfeldt #94-8 U77819 
P. washoensis/BP Babbitt Peak, CA USES #55 U77807 
P. washoensis/MR Mount Rose, NV USES #27 U77808 
P. washoensis/[WM Warner Mtns., CA USES #48 U77809 
P. ponderosa/RM-A Daggett Co., UT USES #206 U77 sil 
P. ponderosa/RM-B Fergus Co., MT C. Baldwin #95-12 U77820 
P..ponderosa/PAC-A Washoe Co., NV USFS #60 U77814 
P. ponderosa/PAC-B Nevada Co., CA USES #71 U77815 
P. ponderosa/PAC-C Illinois River, OR C. Baldwin #95-2 U77818 
P. ponderosa/SCA-A Black Mtn., CA A. Patten #95-8 U77816 
P. ponderosa/SCA-B Fraser Park, CA C. Baldwin #95-13 U77817 
P. ponderosalSW-A Lincoln Co., NM USFS #356 U77813 
P. ponderosa/SW-B Graham Co., AZ USES #10-300 U77812 
P. arizonica/US-A Graham Co., Az USES #1-9 WT7S22 
P. arizonica/US-B Cochise Co., AZ USES #4-119 U77821 
P. arizonica/MX Chihuahua, MX USFS #25-100 U77823 
P. durangensis Durango, MX USFS #29-400 U77824 
P. engelmannii/A Cochise Co., AZ USES #5-108 UZ7825 
P. engelmannii/B Chihuahua, MX USFS #23-200 U77826 


as well as support for previously proposed relation- 
ships within the Ponderosae. The most intriguing 
finding is the existence of a P. jeffreyi lineage, 
which contains P. jeffreyi in a basal position and 
P. washoensis and the North Plateau race of P. pon- 
derosa as derived taxa (Fig. 1). Based on this anal- 


P. contorta 

P. coulteri/A 

P. coulteri/B 

P. jettreyi/ A 

P. ponderosa /NP - ID 
P. washoensis /BP - CA 
P. ponderosa /NP - WA1 
P. washoensis /MR - NV 
P. washoensis /WM - CA 
P. ponderosa /NP - OR 
P. jettreyi/B 

P. arizonica/US - B 


* 


* All other samples have zero branch length 


Fic. 1. Strict consensus tree for 27 samples of Pinus 
using branch and bound search and bootstrap of 1000 rep- 
licates in PAUP 3.1.1. Numbers above the branches rep- 
resent unambigious base substitutions, bracketed numbers 
represent indels. Numbers below the branches represent 
bootstrap values and decay indices, respectively. 


ysis, P. washoensis and the North Plateau race sam- 
ples share a common ancestor with sample A of P. 
Jeffreyi, and this monophyletic group is separated 
from sample B of P. jeffreyi by four derived nucle- 
otide substitutions. The marked differentiation be- 
tween the P. jeffreyi samples was unexpected be- 
cause they come from the same seed provenance in 
the Sierra Nevada. However, extant P. jeffreyi has 
been shown to possess rich intrapopulational ge- 
netic diversity (Furnier and Adams 1986). Future 
phylogenetic studies involving P. jeffreyi need to 
have considerably greater intra- and interpopula- 
tional sampling. Although the P. jeffreyi lineage de- 
scribed in this paper is novel, previous literature 
supports a close relationship among Jeffrey pine, 
Washoe pine, and the North Plateau race. Lauria 
(1991) observed that the purple color of immature 
ovulate cones of these three taxa is unique among 
the Ponderosae. All other members of this subsec- 
tion, including the geographically-proximal Pacific 
race of ponderosa pine, exhibit green to greenish- 
yellow ovulate cone color (Critchfield 1984). Mirov 
(1967) noted the similarity of ovulate cone structure 
between Washoe and Jeffrey pines. 

The results of the phylogenetic analysis also sug- 
gest that Washoe pine and the North Plateau race 
of ponderosa pine form a robust clade (bootstrap 
100%), within which both taxa are polyphyletic 
(Fig. 1). The putative NADH-NR sequences from 
these samples appear to represent four alleles dis- 
tributed randomly among six populations. Due to 


2002] 


the small sample size, it cannot be determined how 
these alleles are structured within or among popu- 
lations or species. Expanded studies using addition- 
al informative DNA regions and statistically-signif- 
icant intrapopulation sampling are needed. Regard- 
less, the random distribution of alleles is consistent 
with the close relationship or even conspecific sta- 
tus previously proposed between Washoe pine and 
the North Plateau race (Wells 1964; Haller 1965; 
Critchfield 1984; Niebling and Conkle 1990; Lauria 
1991, 1997; Brayshaw 1996; Rehfeldt 1999). Prov- 
enance tests by Wells (1964) showed that Washoe 
pine was more similiar to the North Plateau race 
than it was to the Pacific race of ponderosa pine. 
Critchfield (1984) proposed that Washoe pine could 
be a recent derivative of the North Plateau race. 
This view is consistent with a close relationship 
inferred from isozyme data (Niebling and Conkle 
1990). Washoe pine and the North Plateau race 
were found to have a genetic distance of 0.004, a 
value nearly identical to the genetic distance found 
among the three recognized populations of Washoe 
pine. Furthermore, Rehfeldt’s (1999) quantitative 
analysis of adaptive traits determined that Washoe 
pine and the North Plateau race of ponderosa pine 
are very closely related. Based on his and previous 
research, Rehfeldt (1999) concluded that these taxa 
should be considered synonymous. 

The high level of divergence of the Washoe/ 
North Plateau clade (seven synapomorphies and 
two deletions) suggests an origin involving small 
population size and isolation. The divergent Wash- 
oe pine/North Plateau clade eventually came to oc- 
cupy the Willamette Valley and the region approx- 
imating the current extent of the maritime climate 
east of the Cascade crest. Members of this clade 
dispersed as far east as the Continental Divide, 
where an abrupt genetic transition is evident. Latta 
and Mitton (1999) found a steep east-west cline in 
cpDNA and mtDNA, consistent with secondary 
contact between diverged taxa. Similarly, Critch- 
field (1984) previously proposed that Washoe pine 
expanded over the Pacific Northwest only to be lat- 
er absorbed by the North Plateau race of ponderosa 
pine. 

This study includes a single tree from a popula- 
tion on the Fort Lewis plains of the Puget Sound. 
This population is isolated from the North Plateau 
race by the Cascade mountains. Our DNA sequence 
from this individual is identical to that of Ponde- 
rosae found in the Rocky Mountains, Sierra Madre, 
and Sierra Nevada, suggesting that ponderosa pine 
from the Puget Sound region could be a relictual 
population of formerly widespread P. ponderosa 
s.l. This is in no way conclusive, as a single DNA 
marker from a single specimen may not be repre- 
sentative of a population. Nevertheless, it does sug- 
gest that the genetics of Ponderosae in the Puget 
Sound area needs to be investigated in detail as 
numerous studies have hypothesized that the Puget 
Sound region was a glacial (Pleistocene) refugium 


PATTEN AND BRUNSFELD: JEFFREY PINE LINEAGE ON 


for numerous plants and animals (e.g., Harris 1965; 
Steinhoff et al. 1983; Heusser 1985; Soltis et al. 
1997). 

This study does not support the traditional vari- 
etal classification of ponderosa pine (e.g., Conkle 
and Critchfield 1988). Variety scopulorum, the 
Rocky Mountain form, is not distinct in our anal- 
ysis from the Pacific and the Southern California 
races of var. ponderosa. The principal finding of 
this phylogenetic analysis is the existence of a P. 
Jeffreyi lineage that gave rise to Washoe pine and 
the North Plateau race of var. ponderosa, a race that 
appears to be fundamentally distinct from the re- 
mainder of P. ponderosa s.l. Our results cannot 
confirm conspecificity of Washoe pine and the 
North Plateau race of ponderosa pine, but do indi- 
cate a very close relationship of these taxa to each 
other and to Jeffrey pine. This study did not lend 
insights into the relationships among the south- 
western and Mexican species of subsection Pon- 
derosae, suggesting relatively little genetic diver- 
gence of these taxa compared to the P. jeffreyi lin- 
eage. However, morphological and ecological data 
(e.g., Peloquin 1984; Rehfeldt et al. 1996) indicate 
the existence of significant patterns of genetic di- 
vergence in the Ponderosae of the southwestern 
U.S. and Mexico. We hope that this note provides 
the impetus for a more detailed phylogenetic study 
of the subsection Ponderosae. Larger sample sizes 
and multiple, rapidly-evolving DNA segments 
should be analyzed to test the results reported here. 


ACKNOWLEDGMENTS 


We thank G.E. Rehfeldt for providing samples and in- 
valuable guidance, Pam Soltis for assistance with the phy- 
logenetic analysis and her generous advice during the pro- 
ject, and Calib Baldwin for collecting several samples. 
This research was supported by funds provided by the 
Intermountain Research Station, Forest Service, U‘S. 
Dept. of Agriculture, a grant from the Stillinger Trust 
Fund, University of Idaho, and through funding received 
from the NSF-Idaho EPSCoR program under NSF Co- 
operative Agreement #OSR-9350539. 


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MADRONO, Vol. 49, No. 3, p. 193-197, 2002 


NOTEWORTHY COLLECTIONS 


ALASKA 


ALLIARIA PETIOLATA (Bieb.) Cavara & Grande (BRAS- 
SICACEAE).—City and Borough of Juneau, Alaska, in 
the landscaped area next to a parking lot near the inter- 
section of Village and Wittier streets, 58°18'05’N, 
134°24'53"W, 6 June 2001. One mature, flowering plant 
was found by P. Johnson and removed before it set seed. 
It is unclear how this plant arrived at this urban area. It 
is unlikely that it was introduced by the landscaping ac- 
tivities since there had been no recent additions of plants 
or soil. 

Previous knowledge. Native to northern Europe. Com- 
monly called garlic mustard because of the characteristic 
smell of its leaves when crushed, it is a highly competitive, 
aggressive herbaceous invader that forms dense understory 
populations. Present in 34 USA states (http:// 
plants.usda.gov), particularly in the eastern USA (Rollins, 
R.C., The Cruciferae of Continental North America. 1993. 
Stanford University Press) and the four Canadian provinces 
of British Columbia, New Brunswick, Ontario, and Quebec 
(http:/infoweb.magi.com/~ehaber/ipcan.html). The previ- 
ously known British Columbia collections were near Van- 
couver and Vernon (E. Haber personal communication). 

Significance. First record in Alaska. The previously 
known sites in British Columbia are approximately 1300 
km to the southeast. Alliaria petiolata is one of the most 
pestiferous non-native invasive species of forest understo- 
ries in the USA and Canada. It can form monospecific 
stands, exclude native communities, and be essentially im- 
possible to eradicate once it is established. While it was 
originally believed this plant was the only individual 
growing in Alaska, P. Johnson subsequently found a large 
population of plants nearby. Although the early detection 
of this plant has been important, it is unclear whether the 
population can be successfully eradicated. The specimen 
has been placed in the Herbarium at the University of 
California, Davis (DAV). 


—Barry A. Rice, The Nature Conservancy, Wildland 
Invasive Species Team, Department of Vegetable Crops 
and Weed Sciences, University of California, Davis, CA 
95616. 

—PHILLIP JOHNSON, P.O. Box 22898, Juneau, AK 
99802. 


CALIFORNIA 


DROSERA ALICIAE Hamet (DROSERACEAE).—Mendo- 
cino county, CA, 39°15’N, 123°45’W, elevation 160 m, 2 
November 1997. A few hundred meters west of Albion 
Little River Road, just south of the County Airport. A 
single spreading colony of plants was found in wet de- 
pressions and Sphagnum mounds in a pine/cypress pygmy 
forest. 

Previous knowledge. Native to South Africa, Drosera 
aliciae is a plant commonly grown by carnivorous plant 
enthusiasts. 

Significance. It has not previously been collected in 
California, and is probably a new introduction for North 
America. This plant was introduced by horticulturists with 


a number of other carnivorous plant taxa. It was repro- 
ducing both vegetatively and by seed. A more complete 
discussion is given under the Drosera capensis collection 
description. A specimen has been placed in the Herbarium 
at the University of California, Davis, #MR971101. 


DROSERA CAPENSIS L. (DROSERACEAE).—Mendocino 
county, CA, 39°15’N, 123°45’W, elevation 160 m, 2 No- 
vember 1997. A few hundred meters west of Albion Little 
River Road, just south of the County Airport. Large col- 
onies of plants were found growing in wet depressions 
and Sphagnum mounds in a pine/cypress pygmy forest. 

Previous knowledge. Native to South Africa, Drosera 
capensis is a common greenhouse weed in collections 
where carnivorous plants are grown. It is not listed in the 
various floristic works of California. The Jepson Manual 
notes the presence of D. linearis Goldie in Mendocino 
County, but this is probably an erroneous reference to ob- 
servations of D. capensis. A North American Drosera 
species, D. linearis does resemble D. capensis, but D. li- 
nearis is extremely difficult to cultivate and is nearly ab- 
sent from most collections. As such it is unlikely any hor- 
ticulturists have ever planted out D. linearis in California. 
D. linearis grows in Canada from Labrador west to On- 
tario, and in the US in Minnesota, Wisconsin, Michigan, 
and Maine. No plants of D. linearis were found in the 
area. 

Significance. This cluster of wildland locations has been 
used by carnivorous plant horticulturists for introduction 
experiments since the 1970s (P. D’Amato, Carnivorous 
Plant Newsletter, 1988, 17: 15-21). Plants from nearly 
every Carnivorous genus have been planted over the years, 
but most died within a few years. Non-native species that 
persisted and were spreading by seed or vegetative means 
were Drosera aliciae, D. binata, D. capensis, D. capillar- 
is, D. filiformis, D. intermedia, D. nitidula X occidentalis, 
Sarracenia flava, S. leucophylla, S. minor, S. purpurea, S. 
rubra, many interspecific Sarracenia hybrids, and Urtri- 
cularia subulata. Additional species present but which 
may be waifs were Dionaea muscipula, Drosera burman- 
nil, D. slackii, Pinguicula lusitanica, and Utricularia gib- 
ba. None of these plants are included in Californian flo- 
ristic works except for Darlingtonia californica, Drosera 
capensis (incorrectly listed in The Jepson Manual as D. 
linearis), D. filiformis, Sarracenia purpurea, and Utricu- 
laria gibba. Darlingtonia californica was abundant al- 
though it is not native to this location—its nearest natural 
occurrence is in central Trinity County (J. H. Rondeau, 
Carnivorous Plants of California, 1991, unpublished man- 
uscript). Many of the clumps of this plant were heavily 
damaged by infestations of greenhouse thrips, a condition 
not seen in natural populations. The only thriving species 
were Darlingtonia californica, Drosera capensis, and 
Utricularia subulata. Although it is unlikely any of these 
plants will spread from these isolated plantings, Drosera 
capensis or Utricularia subulata (which reproduce both 
vegetatively and by copious seed production) would be 
difficult to eradicate if they invaded other high quality 
natural habitats. The only carnivorous plant native to the 
site is Drosera rotundifolia. Those familiar with the site 
believe the Drosera rotundifolia is being displaced by the 
exotic species (C. Gardner personal communication). 


194 


Specimens have been placed in the Herbarium at the Uni- 
versity of California, Davis, #MR971103. 


UTRICULARIA SUBULATA L. (LENTIBULARIACEAE).— 
Mendocino county, CA, 39°15'N, 123°45’W, elevation 
160 m, 2 November 1997. A few hundred meters west of 
Albion Little River Road, just south of the County Air- 
port. Large colonies of plants were found in wet depres- 
sions and water drainages in a pine/cypress pygmy forest. 

Previous knowledge. This is a widespread species found 
on every continent except Antarctica. In the United States 
it is found on the Atlantic and Gulf coasts from Massa- 
chusetts to Florida to Texas, and inland to Arkansas and 
Tennessee (P. Taylor, The Genus Utricularia: a Taxonomic 
Monograph, 1989, Kew Bulletin Additional Series XIV). 
It is a common greenhouse weed in collections of carniv- 
orous plants. 

Significance. A first collection for California. The plants 
were growing in densely matted clumps and were repro- 
ducing vegetatively and by seed. Both cleistogamous and 
chasmagomous flowers were present. Unless deliberately 
spread, it is unlikely this plant will escape from these 
plantings, but if it did it would be difficult to eradicate. 
Utricularia gibba (probably introduced) was also present 
in flower. A more complete discussion is given under the 
Drosera capensis collection description. A specimen of 
U. subulata has been placed in the Herbarium at the Uni- 
versity of California, Davis, #MR971102. 


—Barry A. RICE, The Nature Conservancy, Wildland 
Weeds Management and Research, Department of Vege- 
table Crops and Weed Sciences, University of California, 
Davis, CA 95616. 


OREGON 


AGROSTIS HOWELLIT Scribn. (POACEAE).—Linn Co., 
rare, N-sloping bench in a moist Acer circinatum-Carex 
deweyana community, Coburg Hills, 25 km NE of Eu- 
gene, T15S R2W S39, elev. 450 m, 3 Oct 1995, Brainerd 
42 (OSC) (! K. L. Chambers 1996 OSC). 

Previous knowledge. Previously known as a narrow en- 
demic from a few sites on the south side of the Columbia 
River Gorge in northern Oregon. 

Significance. Circa 165 km SSW of previously docu- 
mented populations. 


CAREX SCIRPOIDEA Michx. subsp. STENOCHLAENA (Holm) 
A. Love & D. Léve (CYPERACEAE).—Lane Co., drip- 
ping cliff with Salix sitchensis, Agrostis, above Forest Ser- 
vice Road 19, near Cougar Dam, T16S RSE S31, elev. 
520 m, 15 Jul 1998, Newhouse 98027 (MICH, OSC, 
WTU) (! A. A. Reznicek 2001 MICH). 

Previous knowledge. Ledge sedge ranges from Alaska 
south to Washington and Montana. 

Significance. First record for this subspecies in Oregon. 


DAPHNE LAUREOLA L. (THYMELAEACEAE).—Lane 
Co., Laurelwood golf course, Eugene, elev. 215 m, 13 Feb 
1998, Newhouse 98002 (OSC); Hawkins Heights, Eugene, 
elev. 215 m, 20 Feb 1998, Newhouse 98003 (OSC); Ma- 
sonic Cemetery, Eugene, elev. 150 m, Aug 1997, New- 
house 97051 (OSC). 

Previous knowledge. Spurge-laurel is native to Europe, 


MADRONO 


[Vol. 49 


and adventive in British Columbia and western Washing- 
ton, where it is bird-disseminated. 

Significance. First report as an escape from cultivation 
in Oregon. 


GALIUM PEDMONTANUM (Bellardi) All. (RUBI- 
ACEAE).—Benton Co., common in disturbed meadow 
0.5 km NW of Pigeon Butte, elev. 80.m, T13S R5W S32, 
5 Jun 1993, Zika 12025 (OSC, WTU). 

Previous knowledge. Mountain crosswort is native to 
the Mediterranean, and adventive in Idaho, Montana, and 
in the southeastern United States. 

Significance. First Oregon report; discovered by Robert 
Frenkel in 1992. 


PETASITES FRAGRANS (Vill.) C. Pres] (ASTERACEAE).— 
Benton Co., steep forested bank on W shore of Willamette 
River, Corvallis, elev. 60 m, T12S R5W S2, 3 Feb 1999, 
Zika 13717 (OSC, US, WTU); same site, 15 Mar 2000, 
Zika 14848 (OSC). 

Previous knowledge. Winter heliotrope is native to N 
Africa, and occasionally planted as an ornamental in west- 
ern Oregon. 

Significance. First report of an escape from cultivation 
in Oregon. 


—BRUCE NEWHOUSE and RICHARD BRAINERD, Salix As- 
sociates, 2525 Potter, Eugene, OR 97405; and PETER E 
ZIKA, Herbarium, Dept. of Botany, Box 355325, Univ. of 
Washington, Seattle, WA 98195-5325. 


OREGON 


ACAENA NOVAE-ZELANDIAE Kirk (ROSACEAE).—Coos 
Co., Randolph Road, near Route 101, 6 km N of Bandon, 
common weed on sandy banks, roadbeds, dikes, and cran- 
berry fields, with Crepis capillaris, Hypericum boreale, 
Juncus planifolius, J. canadensis, Poa annua, elev. 52 m, 
T28S R14W S4, 7 Sep 1999, Zika 14247 (OSC, WTU); 
Curry Co., Gold Beach, 13 Aug 1951, Jenkins s.n. (OSC); 
adventive on Azalea Lane, Wedderburn, elev. 30 m, T36S 
R1IS5W S25, 7 Jun 2000, Stansell 3196 (OSC); lawn weed, 
Route 101, Gold Beach Ranger Station, Gold Beach, T37S 
R1I5W S1, 17 Aug 2000, Stansell 3201 (OSC). 

Previous knowledge. Biddy-biddy is native to New Zea- 
land, and occasionally cultivated as a ground cover. It 
readily spreads via barbed fruits. Acaena is classified as a 
noxious weed in California, where it is found on disturbed 
ground along the coast. In Oregon it has been observed 
at several sites in addition to the ones vouchered, includ- 
ing a large population at Cape Blanco lighthouse in Curry 
Co. 

Significance. First report for Oregon. We first observed 
Acaena in June 1992 at the U.S. Forest Service office in 
Gold Beach, where lawn mowers scattered the seeds and 
led to an increase in the population. The Oregon Dept. of 
Agriculture has made several unsuccessful attempts to ex- 
tirpate the species with herbicides, starting in 1997. 


—Davip Pivorunas, Navy Region Southwest Natural 
Resource Office, 33000 Nixie Way, Bldg. 50, Suite 333, 
San Diego, CA 92147-5110; VEVA STANSELL, P.O. Box 
6077, Pistol River, OR 97444-1575; and PETER FE ZIKA, 
Herbarium, Dept. of Botany, Box 355325, Univ. of Wash- 
ington, Seattle, WA 98195-5325. 


2002] 


OREGON 


CERASTIUM PUMILUM Curtis (CARYOPHYLLA- 
CEAE).—Jackson Co., pasture with vernal pools, Route 
234 E of Sams Valley, 12 May 1974, Chambers 3974 
(OSC, WTU); Multnomah Co., silty shore, delta of Sandy 
River, elev. 4 m, 16 Apr 1992, Zika 11470 (OSC, WTU), 
Zika 11480 (WTU). 

Previous knowledge. Dwarf mouse-ear is native to Eu- 
rope, and naturalized in eastern North America as well as 
British Columbia. In the Pacific Northwest often growing 
among and confused with C. semidecandrum L. 

Significance. First report for Oregon. 


COTONEASTER DIVARICATUS Rehder & E.H. Wilson (RO- 
SACEAE).—Lane Co., bird-sown in thicket with Quercus 
garryana, Toxicodendron, Morse Ranch Park, Eugene, 30 
Apr 1998, Love 9816 (OSC). 

Previous knowledge. Spreading cotoneaster is native to 
central China, and cultivated as an ornamental in the Pa- 
cific Northwest. 

Significance. First collection of an escape from culti- 
vation in Oregon. 


COTONEASTER INDURATUS J. Fryer & B. Hylm6 (ROSA- 
CEAE).—Lane Co., thickets, near Willow Creek, West 
Eugene, elev. 122 m, 28 May 1992, Zika 11593 (WTU); 
same population, 8 Jul 1997, Zika 13231 (WTU). 

Significance. First reports for hard cotoneaster as an es- 
cape from cultivation. Specimens identified by Jeanette 
Fryer. 


SORBUS CALIFORNICA Greene (CAPRIFOLIACEAE).— 
Klamath Co., Rim above Crater Lake, 12 Aug 1919, 
Sweetser s.n. (ORE): same site, elev. 2150 m, 13 Jul 1929, 
Wynd 1533 (ORE); Wizard Island, Crater Lake, 28 Jun 
1934, Applegate 8977 (OSC); Phantom Ship, Crater Lake, 
elev. 1885 m, Zika 12516 (OSC). 

Significance. First collections for Oregon. All sites are 
in Crater Lake National Park. 


VERONICA VERNA L. (SCROPHULARIACEAE).—Union 
Co., sunny opening, Route 82 beside Wallowa River, 6 
km SE of Minam, elev. 825 m, 2 Jun 1961, Mason 1142 
(ORE): dirt road, Miller Flat, W shore of Wallow River, 
elev. 750 m, 21 May 1994, Zika 12182 (OSC); Wallowa 
Co., dirt road 11 km W of Enterprise, elev. 1220 m, 24 
Apr 1961, Mason 807AA (OSC): 7 miles basalt cliff, Wal- 
lowa Falls, elev. 1525 m, 24 Jun 1962, Mason 5051 
(OSC): Buck Creek near Imnaha River, elev. 580 m, 7 
May 1991, Zika 11089 (OSC). 

Previous knowledge. Spring speedwell is native to Eu- 
rope. Crins et al. (Michigan Botanist 26: 161-166, 1987) 
discuss how to separate it from V. arvensis. Mason (1980, 
Guide to the Plants of the Wallowa Mountains of North- 
eastern Oregon, Museum of Natural History, Univ. of 
Oregon, Eugene) reported her collections of V. verna as 
V. triphyllos. The latter has lower leaves palmately lobed, 
and bracts shorter than fruiting pedicels. Veronica verna 
has pinnately lobed lower leaves and bracts longer than 
the fruiting pedicels. 

Significance. First documentation in Oregon. 


W ASHINGTON 


CARDAMINE FLEXUOSA With. (BRASSICACEAE).— 
Grays Harbor Co., moist ground, edge of building, near 


NOTEWORTHY COLLECTIONS 195 


mouth of Boone Creek, elev. 5 m, 13 Dec 2001, Zika 
16733 (WTU); King Co., wet sunny ditch, Burke Gilman 
Trail 2.4 km N of Matthews Beach, Lake City, Seattle, 
elev. 10 m, 30 Aug 2001, Zika 16467 & Jacobson (WTU);: 
weed in garden bed, Madison Park, Seattle, elev. 10 m, 
31 Aug 2001, Zika 16474 (WTU); San Juan Co., wet 
ground in shade of Alnus rubra, Mineral Point, San Juan 
Island, elev. 5 m, 27 Oct 2001, Zika 16708A (WTU). 

Significance. First report for Washington for this Eur- 
asian native. 


CERASTIUM PUMILUM Curtis (CARYOPHYLLA- 
CEAE).—Island Co., dunes near W shore of Cranberry 
Lake, Whidby Island, elev. 3 m, 20 May 2000, Zika 
I5000A (WTU):; King Co., cracks in asphalt sidewalk, 
Montlake, Seattle, elev. 15 m, 20 May 2000, Zika 14998 
(WTU);: San Juan Co., Turn Point, San Juan Island, 4 Apr 
1992, Atkinson 307 (WTU): sand, Spencer Spit, Lopez 
Island, elev. 2 m, 21 May 2000, Zika 15005 (WTU). 

Significance. First report for Washington. 


COTONEASTER DIVARICATUS Rehder & E.H. Wilson (RO- 
SACEAE).—King Co., bird-sown in thickets, with Cory- 
lus cornuta, Gaultheria shallon, arboretum, near Union 
Bay, Seattle, elev. 20 m, 15 Sep 1999, Zika 14332 & 
Jacobson (WTU); same population, 26 Oct 2000, Zika 
15609 (WTU; dupl. det. by Bertil Hylm6). 

Significance. First collection of an escape from culti- 
vation. 


COTONEASTER LUCIDUS Schltdl. (ROSACEAE).—Colum- 
bia Co., spreading and naturalized in Tucannon River bot- 
tomland, TON R41E S30, elev. 1045 m, 27 Jun 1989, 
Urban 89-001 (OSC). 

Previous knowledge. Shiny cotoneaster is native to Si- 
beria and Mongolia. It is occasionally cultivated in the 
Pacific Northwest, often under the misapplied name C. 
acutifolius Turcz. 

Significance. First collection of an escape from culti- 
vation. 


COTONEASTER NITENS Rehder & E.H. Wilson (ROSA- 
CEAE).—King Co., thickets, partial shade, campus of 
Univ. of Washington, Seattle, elev. 25 m, 26 Oct 1999, 
Zika 14660 & Jacobson (WTU); same population, 6 Nov 
2000, Zika 15645 (WTU). 

Previous knowledge. Few-flowered cotoneaster is native 
to western China, and an uncommon ornamental in west- 
ern Washington. 

Significance. First collection of an escape from culti- 
vation. 


COTONEASTER SALICIFOLIUS Franch. (ROSACEAE).— 
King Co., spreading from cultivation to thickets, Kubota 
Gardens, Rainier Beach, Seattle, elev. 50 m, 3 Nov 1999, 
Zika 14704 (WTU); bird-sown in thickets, campus of 
Univ. of Washington, Seattle, elev. 25 m, 7 Nov 1999, 
Zika 14708 (WTU): bird-sown, partial shade, Madrona, 
Seattle, elev. 50 m, 2 Aug 2000, Zika 15187 (WTU): 
cracks in concrete wall, ship canal near Portage Bay, Se- 
attle, elev. 6 m, 7 Nov 2000, Zika 15655 (WTU). 

Previous knowledge. Willow-leaved cotoneaster is na- 
tive to western China, and planted as an ornamental in 
western Washington for its brilliant autumn fruits. Amer- 
ican robins (Turdus migratorius) and American crows 
(Corvus brachyrhynchos) eat the fruit and disperse the 
seed. 


196 


Significance. First collections for Washington as an es- 
cape from cultivation. 


COTONEASTER TENGYUEHENSIS J. Fryer & B. Hylm6 (RO- 
SACEAE).—King Co., thickets near Washington Park, 
Seattle, elev. 35 m, 15 Sep 2000, Zika 15482 (WTU); 
slope near small creek, Washington Park arboretum, Se- 
attle, elev. 20 m, 2 Nov 2000, Zika 15630 (WTU). 

Previous knowledge. Tengyueh cotoneaster is native to 
SW China, and an uncommon ornamental in western 
Washington. 

Significance. First report of an escape from cultivation 
in Washington. 


CREPIS SETOSA Haller f. (ASTERACEAE).—Clark Co., 
grassy roadside near Loop Road, elev. 10 m, 24 Jul 2000, 
Zika I5114A & Weinmann (WTU); lawn weed by soccer 
field, NE18th St., elev. 90 m, 14 Sep 2001, Zika 16558 
(WTU). 

Significance. First collections in Washington for this 
southern European native. 


CYPERUS DIFFORMIS L. (CYPERACEAE).—Franklin 
Co., sandy E shore of free-flowing Columbia River, elev. 
100 m, TION R28E S1, 2 Oct 2001, Zika 16671 (EIU, 
MICH, WS, WTU). 

Significance. First report for Washington for this Asian 
native. 


FRAXINUS PENNSYLVANICA Marsh. (OQLEACEAE).—Grant 
Co., low ground between dunes, near Potholes Wildlife 
Area, SW of Moses Lake, elev. ca. 330 m, 14 Jun 2001, 
Zika 16258 (WTU); King Co., wet thicket, Madrona Park, 
W shore of Lake Washington, Madrona, Seattle, elev. 5 
m, 2 Aug 2000, Zika 15184 (WTU). 

Previous knowledge. Green ash is native to eastern 
North America, west to Montana, and planted as an or- 
namental in the Pacific Northwest. 

Significance. First record for Washington escaping from 
cultivation and naturalizing. 


GALIUM PEDMONTANUM (Bellardi) All. (RUBI- 
ACEAE).—Klickitat Co., disturbed meadow, Conboy Na- 
tional Wildlife Refuge, elev. 570 m, 15 Jun 2001, Rodman 
508 et al. (WTU). 

Significance. First report for Washington for this Eu- 
ropean native. 


GERANIUM PYRENAICUM Burm. f. (GERANIACEAE).— 
King Co., gravel alleys and waste ground, Madrona, Se- 
attle, elev. 95 m, 14 May 2000, Zika 14969 (OSC, WS, 
WTU). 

Previous knowledge. Hedgerow cranesbill is native to 
Europe, grown in gardens, and known as a weed in Cal- 
ifornia and eastern North America. 

Significance. First report in Washington as an escape 
from cultivation. 


HIERACIUM LACHENALII C.C. Gmel. (ASTERACEAE).— 
King Co., sunny roadside, Route 410, Greenwater, 14 Jun 
2001, Walker s.n. (WTU); Skamania Co., north ridge of 
Table Mountain, 25 Jun 2000, Arnett s.n. (WTU); Sno- 
homish Co., logged area, Perry Creek trail, 5 Aug 1962, 
Kruckeberg 5515 (WTU); common on roadside, Route 2 
east of Index, elev. 300 m, 5 Jun 2000, Zika 15095d 
(WTU). 

Previous knowledge. Hieracium lachenalii s. str. (syn. 
H. acuminatum Jord.) is native to Europe and adventive 


MADRONO 


[Vol. 49 


in eastern North America. All reports of H. vulgatum Fries 

from Washington are H. lachenalii, except one collection 

of true H. vulgatum from Pacific Co. (Maxwell 215 WTU). 
Significance. First collections for Washington. 


HIERACIUM MURORUM L. (ASTERACEAE ).—Pierce Co.., 
roadside and adjacent forest, Route 706 at Westside Road, 
Mt. Rainier National Park, elev. 640 m, 14 Aug 1999, Biek 
2 (WTU). 

Previous knowledge. Wall hawkweed is native to Eu- 
rope, and adventive in eastern North America. Reports of 
H. atratum Fries from Washington belong here. Hieracium 
murorum has been collected as a weed in Portland, 
Oregon (Ornduff 6196 OSC, WTU). 

Significance. First collection for Washington. 


HIERACIUM SABAUDUM L. (ASTERACEAE ).—King Co., 
roadside, Interstate 90, 16 km E of North Bend, elev. 420 
m, 20 Sep 2001, Brunskill s.n. (WTU); Skagit Co., road- 
side, Cain Lake Road near Alger, 3 Sep 1996, Lantz s.n. 
(WTU); Whatcom Co., E Lake Samish Road, ca. 10 km 
S of Bellingham, 2 Sep 1990, Burnett 280 (WTU); Inter- 
state-5, near S end of Samish Lake, elev. 30 m, 9 Sep 
2000, Zika 15465 (WTU). 

Previous knowledge. Savoy hawkweed is native to Eu- 
rope and adventive in eastern North America and British 
Columbia. Reports of H. laevigatum Willd. from Wash- 
ington belong here. 

Significance. First report for Washington. 


HYPERICUM MACULATUM Crantz subsp. OBTUSIUSCULUM 
(Tourlet) Hayek (CLUSIACEAE).—King Co., crack in 
concrete sidewalk, Montlake, Seattle, elev. 25 m, 22 Jul 
2001, Zika 16393 (WTU). 

Previous knowledge. Dotted St. Johnswort is native to 
Europe, and adventive in southern British Columbia. In 
Seattle it is spreading from an introduction in a “wild- 
flower”’ seed mix. 

Significance. First report for Washington. 


JUNCUS PATENS E. Mey. (SUNCACEAE).—Clark Co., 
shade of Fraxinus, Lackamas Creek floodplain, elev. ca. 
65 m, 22 Mar 2001, Zika 15799 (WTU); low ground near 
SE Ist Street, Grass Valley, elev. ca. 70 m, 22 Mar 2001, 
Zika 15802 (WTU). 

Previous knowledge. Native in the Willamette Valley of 
Oregon, 20 km to the S. “Reported but not seen from 
Washington”’ (Hitchcock, Cronquist, and Ownbey, 1969, 
Vascular Plants of the Pacific Northwest, Part 1, Univ. of 
Washington Press). 

Significance. First collections to document this native 
in Washington. 


MOENCHIA ERECTA (L.) P. Gaertn., B. Mey. & Scherb. 
(CARYOPHYLLACEAE).—Pierce Co., dry prairie rem- 
nant, with Lepidium heterophyllum, N of Muck Creek, 
elev. ca. 120 m, 4 Jun 2001, Zika 16157 & Weinmann 
(WTU); dry prairie remnant, Route 507, 5 miles NE of 
Roy, elev. ca. 120 m, 4 Jun 2001, Zika 16166 & Wein- 
mann (WTU). 

Previous knowledge. Upright chickweed is native to 
Europe, and adventive in Oregon and British Columbia. 

Significance. First report for Washington. 


PHOTINIA DAVIDIANA (Decne.) Cardot (ROSACEAE).— 
King Co., bird-sown epiphyte in tree, near Lake Washing- 
ton, Martha Washington Park, Seattle, elev. 10 m, 6 Jun 
2001, Zika 16184 & Jacobson (UBC, WTU); Kitsap Co., 


2002] 


with Alnus rubra, pondshore, Bloedell Reserve, N end of 
Bainbridge Island, Puget Sound, elev. 30 m, 15 Nov 1999, 
Zika 14724 & Jacobson (WTUV). 

Significance. First collections of garden escapes for this 
native of China. 


PHOTINIA VILLOSA (Thunb.) DC. (ROSACEAE).—King 
Co., moist ground, with Alnus rubra, Rubus spectabilis, 
Union Bay, Seattle, elev. 5 m, 29 Sep 2000, Zika 15524 
(WTU); rare adventive, Volunteer Park, Seattle, elev. 130 
m, 8 Nov 2000, Zika 15670 & Jacobson (WTU). 

Previous knowledge. Oriental redtip is native to E Asia, 
and known as a garden escape in the eastern United States. 
Both King Co. sites are adjacent to ornamental plantings, 
and the species was apparently spread by frugivorous 
birds. 

Significance. First report from Washington as an escape 
from cultivation. 


PYRACANTHA COCCINEA M. Roem. (ROSACEAE).—King 


NOTEWORTHY COLLECTIONS 197 


Co., near shore of Portage Bay, Seattle, elev. 5 m, 27 Aug 
1999, Zika 14136 & Jacobson (WTU); San Juan Co., 3.5 
km SE of Sportsman Lake, San Juan Island, 24 Oct 1999, 
Zika 14641 (WTU). 

Significance. First Washington report as an escape from 
cultivation. 


STACHYS ARVENSIS (L.) L. (LAMIACEAE).—King Co., 
weed on gravel roadside, near S shore of Steele Lake, 
Federal Way, 3 Apr 2001, Zika 15866 (WTU). 

Significance. First collection in Washington for this Eu- 
ropean native. 


VERONICA VERNA L. (SCROPHULARIACEAE).—Asotin 
Co., Route 129, Buford Cr., elev. 600 m, 30 May 1991, 
Zika 11135 (OSC, WTU, WS); Chelan Co., N shore Lake 
Chelan, elev. 335 m, 7 Jun 1998, Zika 13427 (WTU). 

Significance. First report for Washington. 


—PETER FE ZiIkA, Herbarium, Dept. of Botany, Box 
355325, Univ. of Washington, Seattle, WA 98195-5325. 


MADRONO, Vol. 49, No. 3, p. 198, 2002 


REVIEW 


A cactus odyssey: Journeys in the wilds of Bolivia, 
Argentina, and Peru. By JAMES D. MAUSETH, ROB- 
ERTO KIESLING, AND CARLOS OSTOLAZA. 2002. Tim- 
ber Press, Portland, OR. 306 pp. ISBN 0-88192- 
526-8. 


This wonderful book presents an engrossing ac- 
count of the authors’ botanical fieldwork in all 
manner of habitats throughout much of South 
America Over a seven-year period. The relaxed nar- 
rative style, beautiful photographs, and liberal in- 
terjection of humor make this an entertaining read 
for just about anyone. Indeed, approachability by 
the layperson is a stated goal of the work, and one 
that is met very admirably. For botanists, the book 
will be even more absorbing, as intriguing aspects 
of taxonomy, morphology, anatomy, ecology and 
physiology of cacti are highlighted throughout. As 
a cactus freak, I found the work completely en- 
thralling. 

I am very inspired by the authors’ philosophy, 
stated clearly at the beginning and throughout the 
work: that cooperation, openness and sharing of 
findings among biologists is beneficial to all. It is 
quite refreshing to view this opinion in print. The 
fruits of this philosophy are evident in the quality 
and number of publications resulting from the field- 
work documented here (Kielsing 1995; Mauseth 
and Kielsing 1997; Ostolaza 1997; Mauseth 1999, 
2000; among others). 

Seven chapters are included: one on cactus bi- 
ology, and two each on the fieldwork conducted in 
Bolivia, Peru, and Argentina, respectively. The in- 
troduction to cacti is broadly written and accessible, 
resulting in a concise yet thoughtful description of 
the family, its evolution, ecology, and taxonomy. 
My one minor criticism of the book is in this intro- 
duction, where the movement of an ocean current 
is described somewhat inaccurately (p. 22). The cir- 
cumglobal southern ocean current (Bartholomew 
1958, Plate 2) precludes movement of water from 
the Atlantic to the Pacific between South America 
and Antarctica. This is the most minor of criticisms, 
however. 

Subsequent chapters entwine absorbing vignettes 
of all manner of cacti in a chronological framework 
of field experience. The discovery of each new tax- 
on on the journey is used to highlight one or more 
interesting facets of cactus biology. For example, 
the appearance of Melocactus includes a discussion 
of cephalia, or the sight of Prosopis (Fabaceae) and 
Prosopanche (Hydnoraceae) touches off a discus- 
sion of mistletoes found in cacti. The authors con- 
sistently relate the spotlighted theme for that taxon 


to other plants, often drawing parallels between the 
plants encountered and other cacti more familiar to 
the North American reader. The authors have done 
a tremendous job here; I cannot emphasize enough 
how captivating and diverse a portrait of cacti is 
presented in this way. The biological material pre- 
sented is written accessibly, but without apparent 
oversimplification. I also appreciate the time spent 
describing those aspects of cactus biology that are 
not easily understood. The authors’ repeated ap- 
peals for students to study these phenomena are a 
welcome incorporation in the work. 

Equally appealing is the description of the va- 
garies of the field. The less-than-ideal road condi- 
tions, rough accommodations, cold food, unsym- 
pathetic authorities and vehicle breakdowns of 
fieldwork are related with the dry wit that I have 
associated with Mauseth since taking his plant anat- 
omy course in 1997. Here, his sense of humor is 
unerring. Anyone who has spent time afield will 
find themselves at least smiling but probably laugh- 
ing outright at the dead-to-rights depiction of being 
constantly outpaced by the sun. 

The photographs are all excellent, and well cho- 
sen to illustrate particular points. South America is 
a beautiful, wild, diverse place as pictured here, and 
the cacti are flat-out gorgeous. My own favorites 
include figures of Browningia candelaris and Azur- 
eocereus. Many wonderful landscape and habitat 
shots are included, as well as several graphical de- 
pictions of life in the field. 

This book has something for everyone, and I rec- 
ommend it to all. I am in the middle of my second 
reading, and it has not lost an ounce of my interest. 


—M. PATRICK GRIFFITH, Rancho Santa Ana Botanic Gar- 
den, 1500 N. College Avenue, Claremont, CA 91711. 
E-mail: michael.patrick.griffith @cgu.edu. 


LITERATURE CITED 


BARTHOLOMEW, J. 1958. The Times atlas of the world: 
mid-century edition, Vol. 1. Times Publishing Com- 
pany, London, U.K. 

KIELSING, R. 1995. Argentine notocacti of the genus Par- 
odia. Cactus and Succulent Journal (U.S.) 67:14—22. 

MAUuSETH, J. D. 1999. Comparative anatomy of Espostoa, 
Pseudoespostoa, Thrixanthocereus, and Vatricania 
(Cactaceae). Bradleya 17:33—43. 

MAusETH, J. D. 2000. Theoretical aspects of surface-to- 
volume ratios and water-storage capacities of succu- 
lent shoots. American Journal of Botany 87:1107— 
NS), 

MAUuSETH, J. D. AND R. KIELSING. 1997. Comparative anat- 
omy of Neoraimondia roseiflora and Neocardenasia 
herzogiana (Cactaceae). Haseltonia 5:37—S0. 

OSTOLAZA, C. 1997. Cactus del sur de Cajamarca y del 
valle del Rio Sana. Quepo 11:57—68. 


MADRONO, Vol. 49, No. 3, p. 199, 2002 


ANNOUNCEMENT 


BIENNIAL GRADUATE STUDENT MEETING AND 
ANNUAL BANQUET 


15 FEBRUARY 2003 AT THE UNIVERSITY OF 
SAN DIEGO 


The California Botanical Society’s Biennial 
Graduate Student Meeting and Annual Banquet will 
be held on Saturday, 15 February 2003 at the Hahn 
University Center of the University of San Diego. 
Graduate students everywhere who are initiating, 
conducting, or finishing research projects in any 
area of botany (e.g., ecology, evolution, conserva- 
tion, floristics, morphology, development, etc.) are 
encouraged to attend the meeting (and banquet) and 
to give short oral presentations on their research 
plans or findings. The venue is an ideal opportunity 
for students to gain experience giving presentations 
in the standard format of scientific meetings, to 
meet students involved in botanical research from 
other institutions, and to learn more about botany 
in general. Presentations will be judged by student 
peers and awards for best papers in proposed re- 
search, research-in-progress, and completed re- 
search will be presented at the evening banquet. 


Abstracts of all presentations will be published on- 
line at the California Botanical Society web-site 
(www.calbotsoc.org). 

Our speaker for the annual banquet will be Dr. 
Jon Rebman, Curator of the Herbarium at the San 
Diego Natural History Museum, who will present 
an after-dinner lecture entitled “‘Discoveries on a 
Floristic Frontier: Baja California.’’ Dr. Rebman’s 
botanical explorations of remote regions of Baja 
California are exemplary of binational collabora- 
tion between the US and Mexico and have yielded 
many exciting findings, including plants previously 
unknown to science and new insights into cactus 
biology and evolution. We look forward to a fas- 
cinating evening of highlights from Dr. Rebman’s 
field research in beautiful and rugged desert ranges, 
such as the Sierra de la Giganta, where he will be 
involved in a major expedition this fall. 

The graduate student meeting and annual ban- 
quet are open to CBS members and non-members 
alike; anyone interested in the meeting and/or ban- 
quet is encouraged to attend. Registration infor- 
mation for the meeting and banquet will be forth- 
coming. 


MADRONO, Vol. 49, No. 3, p. 200, 2002 


ERRATUM 


STuTZ, H. C., M. R. STUTZ, AND S. C. SANDERSON. 
2001. Atriplex robusta (Chenopodiaceae), a new 
perennial species from northwestern Utah. Madro- 
no 48:112—115. 


The name “Atriplex robusta’ H. C. Stutz, M. R. 
Stutz, & S. C. Sanderson (Madrono 48:112. 2001) 
is illegitimate, having already been used (A. robusta 
Speg. In Gand.) 


Atriplex tridentata Kuntze var. robusta H. C. 
Stutz, M. R. Stutz, & S. C. Sanderson, var. 
nov.—TYPE: USA, Utah, Tooele Co., 1 mi W 
of Knolls, T15 R13W S15, shoulder of highway 
I-80, 1280 m elevation, 16 Sep 1977, H. C. Stutz 
$141 (Holotype: BRY; Isotypes, BRY, CA, CAS, 
GH, MO, NY, RM, UC). 


Frutices caespitosi, 40—80 cm alti. Caules erecti 
vel ascendentes, ramosi a basi ad apicem, dense 


furfuraceus, 1-8 mm diam., fragilis. Folia oblonga, 
ascendentia usque appressa, dense furfuraceae; fo- 
lia ephemera verna et aestiva 15—30 mm longa, 5— 
10 mm lata; folia serotina aestiva et hiberna 3—10 
mm longa, 2—5 mm lata, anatomia foliaris Kranz- 
typi. Plantae dioeciae, raro monoeciae. Flores stam- 
inati sessiles, ad brevi-ramulus axillares in angusti 
paniculas terminales; perianthium campanulatum, 
5-partitum ad medium, dense furfuraceum, segmen- 
tis ovatis usque ellipticis, 2 mm longis, | mm latis; 
stamina 5, filamentis | mm longis, antheris ca. 2 
mm longis, | mm latis. Flores pistillati solitarii, ses- 
siles, in pleurumque sine foliis confertas paniculas 
terminales. Bracteae fructiferae furfuraceae, com- 
pressae, urceolatae, latissimae infra media, 5 mm 
latae, 7-8 mm longae, exappendiculatae, cum 3—10 
marginalibus dentibus, 0.5—2 mm longis, qui me- 
dianus longissimus. Utriculus orbiculatus, pericar- 
pio membranceo pellucido. Semena 5 mm diam., 
testa membranacea, brunnea; radicula supera. 


Volume 49, Number 3, pages 137—200, published 17 December 2002. 


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jf 


M a5 é VOLUME 49, NUMBER 4 OCTOBER-—DECEMBER 2002 


MADRONO 


A WEST AMERICAN JOURNAL OF | IND 


va SN Wis 


ai i wlll | 
EW al 


| 


CONTENTS FLOWERING PATTERNS AND REPRODUCTIVE ECOLOGY OF MAMMILLARIA GRAHAMII 

(CACTACEAE), A COMMON, SMALL CACTUS IN THE SONORAN DESERT 

HOT CB LEST ON ON RTI Aa aes Re Pr Ra Oe ne 201 
THE EFFECT OF FIRE AND COLD TREATMENTS ON SEED GERMINATION OF ANNUAL 

AND PERENNIAL POPULATIONS OF ESCHSCHOLZIA CALIFORNICA 

(PAPAVERACEAE) IN SOUTHERN CALIFORNIA 

Arlee M. Montalvo, Laura J. Feist-Alvey, and Catherine E. Koehler ...... 207 
TEMPERATURE LIMITATIONS FOR CULTIVATION OF EDIBLE CACTI IN CALIFORNIA 

Park S. Nobel, Erick De la Barrera, David W. Beilman, Jennifer H. 


ID OMCTIN: GING Mate ONG Is ZU essa athe SP ea con a ees os ad sc ooaggpsaaasaseasedendes 228 
Six New SPECIES AND TAXONOMIC REVISIONS IN MEXICAN GAUDICHAUDIA 

(MALPIGHIACEAE) 

SUCVEI ME VCSSUD r,s Sence nce dscneck saeco see Me uae Op ao soon eye UR Pn hnn sehen vaeindeauese 237 


RETICULATE ANCESTRY IN MEXICAN GAUDICHAUDIA (MALPIGHIACEAE) ANALYZED 

WITH RAPD's AND SOUTHERN HYBRIDIZATION 

Steven L. Jessup’ .... 2075 IY. ACE s AZ LAPS oy aev en cnnserensnnenaseee 256 
LONG-TERM POPULATION DyNAMICS OF NATIVE NASSELLA (POACEAE) 

BUNCHGRASSES IN CENTRAL CALIFORNIA 

Jason G. Hamilton, James R. Griffin, and Mark R. Stromberg ................ 274 


A NEw SPECIES OF PRUNUS (ROSACEAE) FROM THE MOJAVE DESERT OF CALIFORNIA 


BREA IO A EAU ME A Nae ses MY sn one MUN Sea A eee ove ve ve ntneneceoeeee 285 
A NEw CEANOTHUS (RHAMNACEAE) SPECIES FROM NORTHERN BAJA CALIFORNIA, 

MEeExIco 

Steve Boyd anal dor FE. Keeley! iso ON. cascccnes oe p ew wae evenseesseceeseserenes 289 


COLLINSIA ANTONINA IS EVOLUTIONARILY DISTINCT FROM C. PARRYI 
(SCROPHULARIACEAE SENSU LATO) 


BriceG, Baldwin and W. SCOtt AVRIBTUSIET ......<...0cssssoesecsvesecseeveeeseevsseoues 295 
BOOK REVIEW FIELD GUIDE TO LIVERWORT GENERA OF PACIFIC NORTH AMERICA, BY W.B. SCHOFIELD 

Sa Eat SN TN OGG eh tee a ee eae dot a sleds vesteSouW aeiGe nab vbsvniade Usavenvenies 298 

ANNOUNCEMENTS Perse GREP ORE BOR VOLUME A 2 occ caevesessuvevsetnccavcnctutancunsiecncersssevreecseccneeses 299 
BNE, SEP CRUE EUR) VOW LINID AO ..5.cccccbesssccUel oeetsesnctudtonviwavsdéuccsiuvadeveosvavenvesesesseoenens 300 

ESTE TEES STEED 1 27 SO i ee 301 

I egien Sam N NDEI ie ys as gsc Seo un Cee cen cu cideavevovanrecbuvedkadddussvosUviversessvvesneseee 302 

ie abate MEAT TPE OY Feet APIA i, 2. A coescchcescovesoceendenatevncndaeusessvivesewssssecceusnosseete 305 

Par EE OBE INES HOR. V OLLIE A oo oo acc cso reakeeneeaceovactsscesivsserootcssecesalsnccsoonecees il 

aE Sa MANU PRN Esc tee icc tac neva caw oc cusevncecsesnvonewvni es dovenevisdveersbneeeen- iV 


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This paper meets the requirements of ANSI/NISO.Z39.48-1992 (Permanence of Paper). 


MabpDRONO, Vol. 49, No. 4, pp. 201—206, 2002 


FLOWERING PATTERNS AND REPRODUCTIVE ECOLOGY OF 
MAMMILLARIA GRAHAMII (CACTACEAE), A COMMON, SMALL CACTUS 
IN THE SONORAN DESERT 


JANICE E. BOWERS 
U.S. Geological Survey, 1675 West Anklam Road, Tucson, AZ 85745 
jebowers @usgs.gov 


ABSTRACT 


Mammillaria grahamii is an outcrossing species that can flower as many as four times a year. The 
number of flowers produced by an individual plant depends largely on plant volume. Fruit set is high, 
about 77%. Flowers normally live one day but can open a second day when pollination is inadequate. 
Flower production within a population varies interannually and increases with rain in the week before 
flowering. Even heavy rains do not induce all potentially fertile tubercules to flower during a single event. 
Because some tubercules are withheld, plants can flower several times a year, given appropriate conditions. 
Mammillaria grahamii plants risk poor fruit set if they happen to flower when pollinator populations are 
low or when pollinators preferentially visit other species. Withholding of tubercules might allow plants 
to spread this risk across the entire flowering season, thus increasing the probability that flowering will 


coincide with pollinator activity. 


Key Words: Cactaceae, floral biology, floral longevity, Mammillaria grahamii. 


When and how often plants bloom has profound 
consequences for seed set, seed dispersal, and seed- 
ling recruitment, thus making the study of flower- 
ing phenology an important aspect of plant popu- 
lation biology (Primack 1985; Rathcke and Lacey 
1985; Newstrom et al. 1994). Because flowering 
patterns affect the behavior and reproductive suc- 
cess of pollinators and granivores, phenology un- 
derlies the study of evolutionary and community 
ecology as well (Brody 1997). Knowledge of flow- 
ering patterns is, moreover, crucial to conservation 
and management of rare species (Newstrom et al. 
1994). For example, Coryphantha_ robustispina 
(Schott ex Engelmann) Britton & Rose subsp. 
scheeri (Muehlenpfordt) N. P. Taylor, a small, en- 
dangered cactus, can bloom several times per year, 
but because the proportion of sexually mature 
plants that flower during any single event varies 
considerably (Roller 1996), within-year flowering 
patterns must be considered when estimating repro- 
ductive effort. 

In general, flowering patterns and reproductive 
biology of small cacti in the arid southwestern 
United States are poorly known (Johnson 1992). 
The few species that have been studied are for the 
most part endangered or threatened, giving us a 
poor idea of what phenological and floral traits are 
associated with successful reproduction and estab- 
lishment. This paper examines the reproductive 
ecology of Mammillaria grahamii Engelmann, a 
small cactus that is common and widespread in and 
around the Sonoran Desert (Shreve and Wiggins 
1964; Aguilar et al. 2000). The species has been 
known as M. microcarpa Engelmann, not a validly 
published name (Aguilar et al. 2000). 

The reproductive ecology of M. grahamii has not 


been studied in any detail. Flowers are bowl-shaped 
and are relatively small for the family, about 2 to 
4 cm in diameter (Anderson 2001). They arise from 
areoles located in the axils of old tubercules, that 
is, tubercules formed the previous year or earlier 
(Shreve and Wiggins 1964). Tubercules are modi- 
fied leaf bases that appear as regularly spaced pro- 
jections on the plant body. Once an areole has flow- 
ered, it cannot flower again (Gibson and Nobel 
1986). Petals are deep pink or white with a pink 
midrib. Size, shape, and diurnal habit of the flowers 
indicate that they are bee-pollinated (Grant and 
Grant 1979). Stems are solitary or branching and 
grow to a height of 10 to 20 cm. Maximum lifespan 
is about 11 years (Goldberg and Turner 1986). 

Mammillaria grahamii can flower several times 
per year and in any month from March to Septem- 
ber (personal observation; M. Dimmitt personal 
communication). The ultimate goal of this study 
was to determine the benefits of multiple flowering 
events and the conditions under which they occur. 
Specific objectives were to determine: 1) relation 
between plant size and flower production, 2) effect 
of rain on flower production, 3) longevity of indi- 
vidual flowers, 4) minimum reproductive size, 5) 
breeding system, and 6) fruit set and seed produc- 
tion. 


METHODS 


Study area. The study site is located at 720 m 
above sea level about 11 km northwest of Tucson, 
Pima County, Arizona (32°16'N, 111°2’W). Terrain 
is a level to gently sloping alluvial terrace. Soils 
are derived from rhyolitic parent material, and the 
surface is gravelly to cobbly. Vegetation is char- 
acteristic of the Arizona Upland subdivision of the 


202 


Sonoran Desert (Shreve and Wiggins 1964). Dom- 
inant plants include Cercidium microphyllum 
(Torr.) Rose & Johnston, Ambrosia deltoidea (A. 
Gray) Payne, Acacia constricta Benth., Larrea tri- 
dentata (Moc. & Ses.) Cav., Krameria grayi Rose 
& Painter, Carnegiea gigantea (Engelmann) Britton 
& Rose, Opuntia engelmannii Salm-Dyck., Fero- 
cactus wislizeni (Engelmann) Britton & Rose and 
Opuntia leptocaulis DC. Annual precipitation (300 
mm) is seasonally distributed as a highly variable 
winter and early spring (November to March), an 
arid late spring (April to June), a predictable sum- 
mer monsoon (July to August), and a highly vari- 
able autumn (September to October). Maximum 
temperatures in summer often exceed 40°C. Mini- 
mum temperatures rarely drop below —6°C in win- 
ter. Although freezing nights can be frequent in 
winter, daytime temperatures always rise above 
O°C. 

Flower production and plant size. On July 24, 
1996, height and diameter of all stems on 60 M™. 
grahamii plants were measured to the nearest 0.5 
cm, and the number of flowers and flower buds on 
each stem was counted. Plants too small to flower 
were excluded from the sample. Stem volume was 
approximated using the formula for a cylinder, then 
individual volumes were summed to get volume of 
the entire plant. Number of flowers was used as the 
dependent variable in separate linear regressions 
against height of the tallest stem, diameter of the 
thickest stem, number of stems, and plant volume. 


Annual flowering patterns. For monitoring fre- 
quency, duration, and intensity of bloom, all M. 
grahami plants (n = 68) within an area approxi- 
mately 15 m by 20 m were marked on July 27, 
1996 by affixing numbered aluminum tags to the 
ground next to the plant. The sample included some 
but not all of the plants sampled previously and 
also included plants that were too small to flower. 
Height and diameter of the tallest stem of each 
plant were measured to the nearest 0.5 cm. The 
number of open flowers on each plant in the sample 
was counted daily from the first to the last day of 
bloom whenever the sample flowered in 1996 (with 
one exception, described below) and 1997. In ad- 
dition, open and spent flowers were counted on the 
last day of a single blooming event in 1999. Daily 
values in 1996 and 1997 were summed to deter- 
mine the total number of flowers produced per plant 
during each event. Flower production during the 
first blooming event of 1996 was determined on the 
last day of the event by counting number of open 
and spent flowers. At the first event in 1996, flow- 
ers were counted on 25 single-stemmed plants only. 
Thereafter, all marked plants were sampled at every 
event. 


Rain and flower production. The effect of rain 
on flower production was assessed for six flowering 
events in 1996, 1997, and 1999 using only those 
plants for which flowers had been counted at all six 


MADRONO 


[Vol. 49 


events (n = 15 plants). The mean number of flow- 
ers per plant at each event was calculated, then 
Spearman rank-order correlation was used to deter- 
mine the strength of association between flower 
production and rain (mm) in the weeks before flow- 
ering. 


Pollinator visitation. Visits by potential pollina- 
tors were monitored from August 3 to 6, 1996. Al- 
together, 25 flowers on 14 plants were watched for 
10 minutes each, during which time the number of 
bee visitors was recorded. No attempt was made to 
identify the bees. Beetles, which are not effective 
pollinators of cactus flowers (Grant and Connell 
1979), were not included in these surveys. Obser- 
vations were made between 0830 and 1130 hr, the 
period of greatest pollinator activity. 


Characteristics of fruits and seeds. Twenty-four 
ripe fruits were collected in September 1996 and 
individually weighed to the nearest 0.001 g. Length 
and width of each fruit were measured to the near- 
est 0.5 mm. Seeds were removed and air-dried, then 
the mass of seeds from each fruit was measured to 
the nearest 0.001 g. Seed set was determined by 
counting the number of seeds in each fruit. The 
weight of an individual seed was calculated by di- 
viding seed mass by number of seeds. 


Fruit set. Fruit set, defined as the proportion of 
flowers that produced fruits, was studied during two 
separate blooming events in August 1997. Thirty 
reproductively mature plants, none included in the 
previous samples, were numbered and tagged. Fif- 
teen were randomly selected and covered by wire 
mesh cages to prevent pollinators from getting ac- 
cess to the flowers. The number of set fruits was 
counted several weeks after flowering ended. Sep- 
arate Mann-Whitney tests were used to determine 
the effect of treatment (caged or open-pollinated) 
and blooming event (first or second) on the pro- 
portion of flowers that set fruit. 


Flower longevity. The lifespan of individual 
flowers was studied on the same sample of 30 
plants during the first summer blooming event in 
August 1997. All flowers opening for the first time 
were marked on a daily basis with glass-headed 
pins, using a different color of pin for each day. 
The number of freshly opened flowers and the num- 
ber that opened more than once were counted on 
each plant every day. 


Breeding system. Pollination and germination ex- 
periments in a greenhouse were used to determine 
the breeding system. Altogether, 10 self pollinations 
(two flowers from the same plant) and 6 cross pol- 
linations (two flowers from different plants) were 
made. For each cross, one of the flowers was 
tagged with a label indicating date and cross num- 
ber. Stamens were removed from both flowers, then 
the stamens of the untagged flower were used to 
pollinate the stigmas of the tagged flower. Tagged 


2002] 


iz 

® 

= 

2 1997 

TgeOut 1 
60 + =| 
40 - - 
20 | =| 
: L 

146 166 186 206 226 246 266 
Day of Year 
Fic. 1. Flowering curves for Mammillaria grahamii in 


1996 (top) and 1997 (bottom). Y axis represents percent 
of sample in flower; n = 68 except for first event of 1996, 
where n = 25. The first event in 1996 lasted five days but 
flowers were counted only on the final day. 


flowers were checked regularly for developing 
fruits. Mature fruits were collected and dried, and 
the number of seeds in each was counted. 

Seed germination was tested using fruits from 
cross and self pollinations. Twenty seeds from each 
fruit were planted in a four-inch-square plastic pot 
on a moistened, sterilized mixture of sand, turface, 
and milled peat moss. The pots were covered with 
clear plastic wrap to prevent desiccation while al- 
lowing exposure to light. After two months, the 
covers were removed and the pots watered weekly 
during the next three months, then every two weeks 
for the following seven months. The number of liv- 
ing and dead seedlings in each pot were counted at 
two months and twelve months. 


RESULTS 


Annual flowering patterns. As reported previous- 
ly, M. grahamii can flower three to four times in a 
year (Fig. 1). Flower dates in this study were as 
follows: July 21 to 25, 1996; August 2 to 11, 1996; 
May 26 to 30, 1997; August 14 to 20, 1997; August 
29 to September 3, 1997; and September 21 to 23, 
1997. In both years, the most intense blooming 
events occurred in summer and lasted more than a 
week (Fig. 1). Typically, 60 to 80% of the sample 
flowered during the first summer event. Later sum- 
mer events involved fewer plants and lasted only 
three to four days. No flowers were produced in 
spring of 1996. Spring flowering in 1997 involved 
about 40% of the population and lasted five days. 


Flower production and plant size. In the sample 
of reproductively mature plants (n = 60), volume 


BOWERS: FLOWERING PATTERNS OF MAMMILLARIA 203 


100 — re 
” 80 + = 
10) 
= ® e 
= 60} 
re 
ro) ® 
= 40 ¢ ® e as 
3 we ° y = 4.28 + 0.04x 

20 ope ia e r = 0.75 

o f= <wAte ane a 2) Se es Ee : 
0 500 1000 1500 2000 
Plant volume (cm*) 

Fic. 2. Flower production in a single flowering event as 


a function of plant volume (n = 60). 


explained 75% of the variation in flower number 
(P = 0.001) (Fig. 2). Number of stems and height 
of the tallest stem respectively accounted for only 
44% (P = 0.001) and 32% (P = 0.001) of the var- 
iation in number of flowers. Diameter of the thick- 
est stem was not significantly correlated with flower 
production (r7 = 0.06, P = 0.07). In the 68-plant 
sample, which included plants too small to flower, 
the smallest plants to bloom were 2.5 cm in height; 
virtually all plants > 4 cm in height produced flow- 
ers. 


Rain and flower production. Spearman correla- 
tion analysis showed that rain in the week before 
flowering was strongly correlated with mean num- 
ber of flowers produced by 15 plants during six 
flowering events (r, = 0.94, P < 0.05) (Fig. 3). 


14 - 
= 42 - : 
oO 
a 
O 
Qa 
2 [- T = 
(00) 
= LL 
ne) ~ 
= [ : 
oO 
) 

E 

2 

= 

iS 

| | 

3 = 5 
Event 

Fic. 3. Correlation between flower production and rain- 


fall. Top: mean number of flowers/plant (+ 1 SE) at six 
flowering events. Bottom: rain (mm) in the week before 
flowering. First day of flowering as follows: event 1, Au- 
gust 2, 1996; event 2, August 21, 1996; event 3, May 27, 
1997; event 4, August 15, 1997; event 5, August 29, 1997; 
event 6, July 14, 1999. 


204 


TABLE 1. FRUIT AND SEED CHARACTERISTICS, MAMMILLARIA 
GRAHAMII; PEARSON CORRELATION COEFFICIENTS (N = 24 
Fruits). * = Significant at P < 0.001. 


Fruit Fruit Fruit Seed 
mass length width mass 
Fruit length 0.54 
Fruit width OFS 0.13 
Seed mass 0.40 =().37 0.75% 
Seed number 0.45 =()),25 OVBe 0.89* 


Mean flower production was also correlated with 
rain in the two weeks before flowering (r, = 0.90, 
P < 0.05) but not with rain in the three weeks be- 
fore flowering (r, = 0.77, P > 0.05). 


Pollinator visitation. Visitation varied consider- 
ably over four days: visits/flower/hr (+SD) on the 
first through fourth days of observation were 0.0, 
5.3 = 3.6, 5.0 = 2.4, and 41.1 + 25.3, respectively. 
Cloudy, humid weather on the first day of obser- 
vation may have limited pollinator activity; the re- 
maining days were clear and hot. On the second 
and third days of observation, all flower visitors 
were small bees, whereas on the fourth day, most 
visitors were honeybees. 


Characteristics of fruits and seeds. In the wild 
population, the number of seeds per fruit averaged 
149 = 45 (SD). Fruit length and width (+SD) were, 
respectively, 26.0 mm + 5.2 mm and 7.2 mm + 
0.9 mm. Mass of all seeds in a single fruit averaged 
0.038 g + 0.016 g (SD). Calculated mass of an 
individual seed was 0.0003 g. Seed mass constitut- 
ed on average 9.1% of fresh fruit mass. Fruit width 
was the best predictor of number of seeds, seed 
mass, and fruit mass (Table 1). In addition, seed 
mass and number of seeds were highly correlated 
(Table 1). 


Fruit set. Fruit set of caged and open-pollinated 
plants averaged, respectively, 1.4% and 68.0% in 
the first blooming event, 1.1% and 85.0% in the 
second. The difference between treatments was 
highly significant (Mann-Whitney U = 0.0, P = 
0.001), but the difference between blooming events 
was not (Mann-Whitney U = 350.0, P = 0.70). 


Flower longevity. Most flowers, whether caged 
or not, opened only a single day. Nevertheless, the 
proportion of flowers that opened a second day was 
almost 14 times greater for caged (63 of 284 flow- 
ers) than uncaged plants (3 of 188 flowers). It ap- 
pears that flowers typically live a single day if the 
level of pollination is adequate and that they can 
sometimes open a second day if it is not. 


Breeding system. Fruit set was 100% for crossed 
flowers (6 of 6) but only 10% for selfed flowers (1 
of 10). The single fruit produced from the self-pol- 
linated flowers contained 281 seeds, but none ger- 
minated. Thus, in the rare instance when self pol- 
lination yielded fruit, the seeds apparently were not 


MADRONO 


[Vol. 49 


viable. The average number of seeds in fruits pro- 
duced by cross pollination was 208 + 45 (SD). This 
was considerably higher than in the wild population 
and probably reflects a difference in pollen loads. 
Seeds from cross-pollinated fruits were highly ger- 
minable. Two months after planting, germination 
averaged 11.7%. After twelve months, average ger- 
mination was 78.3%. 


DISCUSSION 


In some respects, the floral biology of M. gra- 
hamii is similar to that of other small cacti in the 
arid southwestern United States (Table 2). As for 
Echinomastus erectocentrus (J. M. Coulter) Britton 
& Rose (Johnson 1992) and Escobaria robbinso- 
rum (W. H. Earle) D. R. Hunt (Schmalzel et al. 
1995), plants reach reproductive maturity at a small 
size, and flower production increases as plants 
grow. As with Sclerocactus polyancistrus (Engel- 
mann & Bigelow) Britton & Rose (May 1994), var- 
iability in flower production from year to year or 
event to event is strongly correlated with rain. 

In other respects, the reproductive strategy of M. 
grahamii, a common and relatively widespread spe- 
cies, differs from that of small cacti that are rare or 
narrowly distributed. The number of flowers per 
plant and seeds per fruit is considerably higher (Ta- 
ble 2). The potential flowering season is longer, as 
well, and there can be two to four times as many 
flowering events per year (Table 2), a pattern that 
Newstrom et al. (1994) describe as “‘subannual.”’ 
The abundance and distribution of any species nec- 
essarily arises from multiple intrinsic, environmen- 
tal, and historical factors and cannot be ascribed to 
reproductive biology alone. Nevertheless, the com- 
bination of subannual blooming, large number of 
flowers per plant, high fruit set, and high seed pro- 
duction makes M. grahamii substantially more fe- 
cund than other small cacti (Table 2) and doubtless 
contributes to its success. 

Small cacti are at high risk of illicit collection 
(Bennett et al. 1986). During one study, for exam- 
ple, collectors illegally removed 31% of 324 M. 
grahamii and 44% of 9 M. thornberi Orcutt (Ben- 
nett et al. 1986). The latter species is considered 
vulnerable (Nabhan et al. 1989). Although the per- 
centages are roughly equivalent, the number of 
plants remaining—23 versus 5—is not. Especially 
for small cacti that have low fecundity, such dep- 
redations can represent a substantial portion of the 
reproductive capacity of the population. 

It remains to be seen whether M. grahamii, like 
other small cacti in the region, is obligately out- 
crossing. In greenhouse experiments, only 1 of 10 
self-pollinated flowers set fruit. Because these flow- 
ers received ample pollen, low fruit set was likely 
a consequence of self incompatibility rather than 
inadequate transfer of pollen from stamens to stig- 
mas. None of the seeds from the selfed fruit ger- 
minated, in contrast to seeds from cross-pollinated 


2002] 


BOWERS: FLOWERING PATTERNS OF MAMMILLARIA 


205 


TABLE 2. COMPARATIVE FLORAL BIOLOGY OF SELECTED SMALL CACTI. Species abbreviations and data sources as follows: 
CORY, Coryphantha robustispina subsp. scheeri (Roller 1996); ECHI, Echinomastus erectocentrus (Johnson 1992): 
ESCO, Escobaria robbinsorum (Schmalzel et al. 1995); MAMM, Mammillaria grahamii; SCLE, Sclerocactus polyan- 
cistrus (May 1994). Flower and stem sizes are from Anderson (2001). Other abbreviations: SS, single-stemmed: MS, 
multiple-stemmed; n.d., no data. Information on status is from http://arizonaes.fws.gov and http://www.cnps.org. 


CORY ECHI ESCO MAMM SCLE 
Flower diameter 5-7 4-5 1-2 2-4 5 
Stem height (cm) 5-15 10—37 2-6 7-20 10—40 
Stem diameter (cm) 5-9 7-12 2-6 7-11 5-9 
Flowers/plant/yr (range) 1—23 1-16 1-7 1-136 1-6 
Flowers/plant/yr (mean) iL n.d. 2 23 4 
Height at first lowering (mm) 30 24 13 DS 30 
Breeding system Outcrossing Outcrossing Outcrossing Outcrossing Outcrossing 
Fruit set (percent) 71 94 93 a n.d. 
Seeds/fruit 89 oF 20 149-209 120 
Germination (percent) 89 n.d. n.d. 12-78 “low” 
Flower longevity (days) ] n.d. n.d. 1 (2) a7 
Flowering season May-—Jul Mar—Apr Mar—Apr Mar—Sep Apr—May 
Flowering events/yr 1-3 1 1 1-4 | 
Duration of events (days) 1 Dag | n.d. 3-11 n.d. 
Lifespan (yr) <30 n.d. ilg/ 11 17 
Plant morphology SS, MS SS SS SS, MS SS 
Status Endangered Candidate Threatened Not listed “Watch 
fruits, which germinated at a high rate. Taken to- ACKNOWLEDGMENTS 


gether, the pollination and germination experiments 
suggest that M. grahamii is obligately outcrossing; 
additional experimental work is needed to deter- 
mine whether this is indeed the case. 

Because M. grahamii produces only a single 
flower in each axil, flower production in one year 
is ultimately limited by the number of tubercules 
produced in previous years and by the proportion 
of old tubercules that have already flowered. An- 
nual tubercule production is in turn determined by 
plant volume and, probably, rainfall. The intensity 
of bloom during any single event is also a function 
of rain, at least in part. When rains are minimally 
adequate, the proportion of tubercules that flower 
is small; when rains are relatively large, many tub- 
ercules bloom (Fig. 3). On the other hand, even 
heavy rains do not induce all potentially fertile tub- 
ercules to flower during a single event. Because 
some tubercules are withheld, plants can flower 
several times a year, given appropriate conditions. 

Fruit set of M. grahamii and certain other small 
cacti is relatively high (Table 2). In the case of 
Echinomastus, fruit set apparently is not affected 
by pollinator abundance or effectiveness (Johnson 
1992). This might not be true for M. grahamii. The 
results of the caging experiment suggest that plants 
risk poor fruit set if they happen to flower when 
pollinator populations are low or when pollinators 
preferentially visit other species. This does happen 
at least occasionally, as in 1996 when visitation on 
one day was essentially nil. Withholding of tuber- 
cules might allow plants to spread the risk across 
the entire flowering season, thus increasing the 
probability that flowering will coincide with polli- 
nator activity. 


Thanks to S. P. McLaughlin for reading the manuscript 
and for providing data from his greenhouse studies. 


LITERATURE CITED 


AGUILAR, R. P., T. R. VAN DEVENDER, AND R. S. FELGER. 
2000. Cactaceas de Sonora, México: su diversidad, 
uso y conservacion. Arizona-Sonora Desert Museum 
Press, Tucson, AZ. 

ANDERSON, E. FE 2001. The cactus family. Timber Press, 
Portland, OR. 

BENNETT, P., R. R. JOHNSON, AND M. R. KUNZMANN. 1986. 
Cactus collection factors of interest to resource man- 
agers. Pp. 215-223 in T. S. Elias (ed.), Conservation 
and management of rare and endangered plants. Cal- 
ifornia Native Plant Society, Sacramento, CA. 

Bropy, A. K. 1997. Effects of pollinators, herbivores, and 
seed predators on flowering phenology. Ecology 78: 
1624-1631. 

GiBsON, A. C. AND P. S. NOBEL. 1986. The cactus primer. 
Harvard University Press, Cambridge, MA. 

GOLDBERG, D. E. AND R. M. TURNER. 1986. Vegetation 
change and plant demography in permanent plots in 
the Sonoran Desert. Ecology 67:695-—712. 

GRANT, V. AND W. A. CONNELL. 1979. The association 
between Carpophilus beetles and cactus flowers. 
Plant Systematics and Evolution 133:99—102. 

GRANT, V. AND K. A. GRANT. 1979. The pollination spec- 
trum in the southwestern American cactus flora. Plant 
Systematics and Evolution 133:29-—37. 

JOHNSON, R. A. 1992. Pollination and reproductive ecol- 
ogy of acuna cactus, Echinomastus erectocentrus vat. 
acunensis (Cactaceae). International Journal of Plant 
Science 153:400—408. 

May, R. W. 1994. The ecology of Sclerocactus polyan- 
cistrus (Cactaceae) in California and Nevada. Desert 
Plants 11:6—22. 

NABHAN, G. P., E. SAUCEDO M., P. OLWELL, P. WARREN, 


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W. Hopcson, C. GALINDO, R. BITTMAN, AND S. AN- 
DERSON. 1989. Plants at risk in the Sonoran Desert: 
an international concern. Agave 3:14—15. 

NewstTroM, L. E., G. W. FRANKIE, H. G. BAKER, AND R. 
K. COLWELL. 1994. Diversity of long-term flowering 
patterns. Pp. 142-160 in L. A. McDade (ed.), La Sel- 
va: ecology and natural history of a neotropical rain 
forest. University of Chicago Press, Chicago, IL. 

PRIMACK, R. B. 1985. Patterns of flowering phenology in 
communities, populations, individuals, and single 
flowers. Pp. 571-593 in J. White (ed.), The popula- 
tion structure of vegetation. Dr. W. Junk Publishers, 
Dordrecht, The Netherlands. 


MADRONO 


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RATHCKE, B. AND E. P. LAcEy. 1985. Phenological patterns 
of terrestrial plants. Annual Review of Ecology and 
Systematics 16:179—214. 

ROLLER, P. S. 1996. Distribution, growth, and reproduction 
of Pima Pineapple Cactus (Coryphantha  scheeri 
Kuntz var. robustispina Schott). M.S. thesis. Univer- 
sity of Arizona, Tucson, AZ. 

SCHMALZEL, R. J., E W. REICHENBACHER, AND S. RUTMAN. 
1995. Demographic study of the rare Coryphantha 
robbinsorum (Cactaceae) in southeastern Arizona. 
Madrono 42:332—348. 

SHREVE, FE AND I. L. WiGGINs. 1964. Vegetation and flora 
of the Sonoran Desert. Stanford University Press, 
Palo Alto, CA. 


Maprono, Vol. 49, No. 4, pp. 207-227, 2002 


THE EFFECT OF FIRE AND COLD TREATMENTS ON SEED 
GERMINATION OF ANNUAL AND PERENNIAL POPULATIONS OF 
ESCHSCHOLZIA CALIFORNICA (PAPAVERACEAE) IN 
SOUTHERN CALIFORNIA 


ARLEE M. MOoNnrtTALVO!, LAURA J. FEIST-ALVEY’, AND CATHERINE E. KOEHLER? 
Department of Botany and Plant Sciences and Center for Conservation Biology, 
University of California, Riverside, CA 92521 


ABSTRACT 


Throughout its native range, the California poppy, Eschscholzia californica, exhibits substantial mor- 
phological and life-history variation, including variation in seed dormancy and ability to perennate. Pop- 
ulations from xeric southern California habitats have high seed dormancy over a range of habitats that 
span from coast to desert and across vegetation types of varying fire frequency. Understanding variation 
in the cues that break dormancy in this species is especially important to managing natural populations 
with prescribed fire, and in production and use of local ecotypes for restoration, erosion control, and 
ecological landscaping. We explored the influence of sequential treatments of low temperature and com- 
ponents of fire (heat, dry smoke, smoke water, a commercially concentrated smoke water we call “‘liquid 
smoke’’, charrate, and nitrate) on seed germination for both annual and perennial populations and com- 
pared results to those of widely used domesticated seeds. We also examined the effect of light and seed 
age. Domesticated seeds had no seed dormancy and, except for heat-treated seeds, germination was close 
to 100% across treatments, including water controls. In contrast, seeds of all wild southern California 
populations showed some dormancy, germination was highly conditional on test factors, and light inhib- 
ited germination. We found differences in dormancy rates among wild populations and years since seed 
collection, with annuals having higher dormancy than perennials in the first year following collection but 
not after aging > two years. Of the fire treatments, heat (85°C for 10 min), or heat plus smoke, resulted 
in significantly reduced germination and viability of all populations tested, including domesticated seeds. 
All smoke treatments significantly improved germination of dormant-seeded populations over water con- 
trols, but neither nitrate, water soaking, charrate, nor cold treatment alone broke dormancy. In the absence 
of cold treatment, both liquid and dry smoke yielded higher germination than controls in seeds aged 8— 
10 months (65-95% for liquid smoke, 21—60% for dry smoke, vs. 14-59% for controls). Moist cold 
treatment (3—9°C) by itself did not usually break seed dormancy but it did act synergistically to increase 
germination of smoke-treated seeds and did not harm controls. In contrast, for most populations colder 
pretreatment (~2°C) resulted in a small decrease in germination of water controls but not in seeds smoke- 
treated before cold treatment. Seed age affected germination of controls and the ability of smoke to break 
dormancy. Germination of controls and smoke-treated seeds increased between 2 and 4 mo of aging in 
the lab, with no further increase at 8 mo. Dormancy of controls was substantially higher in seeds aged 
in the lab > 27 months from collection relative to seeds aged 8—10 months (n = 7 and 5 populations, 
means = 92% and 63% dormant, respectively). Smoke succeeded in breaking dormancy of older seeds 
to half the extent as in younger seeds, suggesting either a decline in germinability as seeds degrade, 
induction of a deep secondary dormancy, or both. Given the large differences between domesticated and 
wild populations in dormancy and germination requirements, and that seed dormancy is probably heritable 
and adaptive, non-dormant domesticated seeds are not appropriate for restoration, especially in xeric 
environments that naturally support plants with dormant seeds. 


Key Words: Eschscholzia, fire, germination treatments, life-history variation, restoration, seed dormancy, 
smoke, stratification. 


As the use of native plants for revegetation and 
landscaping gains popularity, and as restoration 
projects become more prevalent and species-inclu- 
sive, understanding seed germination biology be- 


' Author for correspondence, e-mail: montalvo @ citrus. 
ucr.edu 

* Current address: Remediation Division, Montana De- 
partment of Environmental Quality, PO. Box 200901, He- 
lena, MT 59620-0901. 

* Current address: University of California Davis— 
McLaughlin Reserve, 26775 Morgan Valley Rd., Lower 
Lake, CA 95457-9411. 


comes essential from both basic and applied eco- 
logical perspectives. In warm and dry climates, na- 
tive seeds can be difficult to germinate because of 
dormancy, and they often require very specific ger- 
mination cues. As is the case with many traits, both 
dormancy and response to dormancy-breaking sig- 
nals can be locally adaptive and can differ dramat- 
ically within species, especially when they exist in 
a wide variety of habitats (Cruden 1974; Capon et 
al. 1978; Keeley 1986; Meyer et al. 1990; Meyer 
and Monson 1992). If cues that break seed dor- 
mancy are identified, then practitioners can use the 
information to maximize germination of out-plant- 


208 


ed seeds and of seeds used for agricultural increase, 
thus limiting selection against genes conferring dor- 
mancy. Seed increase encourages use of local seed 
sources for restoration, landscaping, and roadside 
erosion control, thereby minimizing mismatching 
of important adaptive traits to planting location. 
Mismatches can decrease project success and intro- 
duce maladapted genes into wild populations (Mon- 
talvo et al. 1997; Keller et al. 2000; Montalvo and 
Ellstrand 2000, 2001). 

The effects of fire on germination response are 
so common that it is worthwhile to include fire 
treatments in studies of seed dormancy, especially 
for species with close relatives known to be fire 
followers. Seeds of many species in Mediterranean 
climates, where fire is common, germinate in re- 
sponse to different fire-related cues (Keeley and 
Keeley 1987; Keeley 1991; Roche et al. 1997a, b, 
1998). Also, as prescribed burning becomes an in- 
creasingly important management tool, it is impor- 
tant to examine the effects of fire on seeds, and to 
determine if populations from different environ- 
ments have different requirements and vulnerabili- 
ties. 

We chose California poppy (Eschscholzia cali- 
fornica Cham.) as a model species for studying var- 
iation in seed dormancy and response to different 
germination cues. California poppy occupies many 
habitats throughout its native range from Baja Cal- 
ifornia to southern Washington state, and is native 
to a diverse range of climates, edaphic environ- 
ments, and plant communities that vary in fire fre- 
quency, from coastal sand dunes to inland deserts, 
including fire-prone coastal sage scrub, chaparral, 
and grasslands. Populations exhibit a wide spec- 
trum of morphologies and life histories, varying in 
flower color, size, seed dormancy, and other attri- 
butes, and can be annual (semelparous), long-lived 
perennial (iteroparous), or a facultative annual 
(Cook 1962). In addition, this species has promi- 
nent economic importance as the California state 
flower and is frequently used in restoration and re- 
vegetation. Information about variation in its ger- 
mination response to the effects of fire and other 
factors is essential for long-term success of projects 
that utilize prescribed burning for restoration, for 
commercial seed production, and for conservation 
practice. 

In southern California, native poppies (both an- 
nuals and perennials) germinate during the rainy 
season in mid winter; plants flower in late winter 
to spring, and produce seeds in the spring (ate 
April to early June), although some perennials can 
defer flowering to the second spring and produce 
seeds over a longer season. Seeds appear to remain 
dormant (or conditionally dormant) until the next 
winter rains, which prevents early germination after 
infrequent summer storms and subsequent death 
from desiccation in the hot, dry summer environ- 
ment. Thus plants appear to fit the ““winter annual 
and perennial’”’ syndrome (Baskin and Baskin 1998, 


MADRONO 


[Vol. 49 


p. 54-56), but it is unknown whether the seeds fol- 
low an annual cycle of summer dormancy followed 
by non-dormancy or conditional dormancy during 
the winter. Based on the small linear embryo of 
California poppy seeds, seeds have been assumed 
to have morphophysiological dormancy in which 
embryos complete their maturation after seed dis- 
persal and an environmental cue is required to 
break physiological dormancy (Martin 1946; Bas- 
kin and Baskin 1998). It is unknown if embryo 
growth and dormancy break require the same or 
different conditions. 

In interior shrub- and grassland habitats of south- 
ern California, prolific displays of poppies in the 
absence of fire are periodic and tend to be separated 
by many years. In some years, flowering of annual 
populations is absent or scant and can be followed 
by boom years, demonstrating the existence of a 
substantial seed bank. This suggests that seeds may 
cycle through dormancy and conditional dormancy 
depending on environmental conditions. In addi- 
tion, the conspicuous presence of poppies on recent 
burns when blooming is poor elsewhere (A. Mon- 
talvo personal observation, G. Hund, R. Noll, J. 
Crossman personal communication) suggests that, 
at least in these habitats, seeds survive fire and 
some component of fire aids in breaking seed dor- 
mancy. 

In California, fire and fire components have been 
shown to break seed dormancy in many “‘fire-fol- 
lowing” species, especially in sage scrub and chap- 
arral communities (Keeley 1991). For example, seed 
dormancy of Emmenanthe penduliflora Benth., Sal- 
via columbariae Benth., S. mellifera Greene, Pha- 
celia grandiflora (Benth.) Gray, P. minor (Harv.) 
Thell., and Lotus scoparius (Nutt.) Ottley is broken 
by heat, smoke, or other components of fire (Keeley 
1991; Keeley and Fotheringham 1998a, b). Each of 
these species occurs in a number of different plant 
communities, frequently concomitant with California 
poppy (Munz and Keck 1968). In studies of intra- 
specific germination response to heat, very different 
patterns were found among populations of perennial 
S. mellifera (Keeley 1986) and annual S. columbar- 
tae (Capon et al. 1978) from desert habitats vs. chap- 
arral and coastal sage scrub. Given that the distri- 
bution of California poppy is even more diverse than 
for these Salvia species, its germination response to 
fire is likely at least as complex. The response of 
poppy seed germination to components of fire has 
not been studied, although prescribed burning is be- 
ing used to manage portions of its native habitat, 
including the Antelope Valley California Poppy Re- 
serve (AVCPR, www.calparksmojave.com/poppy; 
www.parks.ca.gov/parkindex, J. Crossman, personal 
communication). 

Data on the general germination biology of Cal- 
ifornia poppy are scant, and few authors have iden- 
tified use of wild rather than domestic seeds in their 
work (e.g., Cook 1961, 1962; Fox et al. 1995). 
Cook (1961, 1962) documented geographic varia- 


2002] 


TABLE 1. 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 


209 


SEED POPULATIONS USED IN SEED GERMINATION EXPERIMENTS. Abbreviations: Pop = population; A = annual; 


P = perennial. Source, locality, and life-history strategy are listed for each study population. 


Source of seeds and collection date 


S&S Seeds (Carpinteria, CA) Lot #T8979, 
~May 1998 

L. Feist, May 25, 2000 

Stover Seed Co. (Los Angeles, CA), harvested 
for 2000 market 

J. Crossman, May 1995 


G. Fox, ~May 1996 

L. Feist, May 17, 2000 

A. Montalvo and L. Feist, May 27, 2001 

S&S Seeds (Carpinteria, CA), Lot #87383, 
~May 1997 

R. Noll, S&S Seeds (Carpinteria, CA), Lot 
#T1008, Apr. 29—May 11, 1998 

L. Feist, May 11—23, 2000 

A. Montalvo, L. Feist, May 8-13, 2001 

G. Fox, ~May 1996 

L. Feist, May 25 and June 6, 2000 

A. Montalvo and L. Feist, May 27, 2001 

R. Noll, S&S Seeds (Carpinteria, CA), Lot 
#T1022, May 7-9, 1998 

R. Noll, S&S Seeds (Carpinteria, CA), Lot 
#V 1022, May 8-11, 2000 

A. Montalvo and J. Skillman, May 19, 2001 


Pop Life 

Code history Collection locality 

LC98 P Antelope Valley area, Lancaster, CA 

AVO0O Pp Antelope Valley area 2, CA 

Dom P Cultivar domesticated in Salem, OR 

25-30 generations (origin s. California) 

PR95 A California State Parks, Antelope Valley Cali- 
fornia Poppy Reserve, CA (AVCPR) 

PR96 A as in PR95 

PROO A as in PR95 

PROI A as in PR95 

CB97 A Carlsbad, CA 

NH98 A North Hills of Western Riverside County 
Multi-Species Reserve, CA 

NHO0OO A as in NH98 

NHO1 A as in NH98 

FM96 E Fairmont Butte (s. of AVCPR) 

FMO0O Pp as in FM96 

FMO1 Pp as in FM96 

EM98 A Vicinity of Estelle Mt., ne Lake Elsinore, 
Riverside Co,. CA 

EMO00 A as in EM98 

SR101 lf Serpentine site, Sedgwick Ranch Reserve, 
Santa Barbara Co., CA 

RVO1 P Riverside, Riverside Co., CA, domesticated 


population 


tion in seed dormancy that was loosely correlated 
with habitat and longevity; annual and perennial 
populations from xeric sites had mostly dormant 
seeds, whereas perennial populations from mesic 
sites had little to no dormancy. He found that dor- 
mancy could sometimes be broken with gibberellic 
acid (GA3), but he did not identify the natural cues 
that break dormancy. 

Here we assessed the germination response of E. 
californica seeds from several southern California 
populations to various components of fire and other 
potential dormancy breaking conditions. We includ- 
ed seeds from both annual and perennial wild pop- 
ulations, neighboring and distant populations, and 
a variety of climate regions including coastal, in- 
land, and desert. We examined whether the popu- 
lations varied in seed dormancy, effect of seed ag- 
ing, and in germination response to a variety of fire 
treatments, including heat, smoke, charrate, and ni- 
trogen. Poppies do not usually germinate until well 
into the cold season (December—February). Be- 
cause many GA3 responsive seeds also respond to 
cold treatment, we also explored the effect on ger- 
mination of exposure to a short period of cold tem- 
peratures (e.g., Roche et al. 1998), using varying 
temperatures, length of exposure to cold, and mois- 
ture levels during exposure to cold. 


A. Montalvo May 27, 2001 


METHODS 


Study populations. All wild populations were 
from southern California and represented both an- 
nual and perennial life-histories (Table 1). Seed col- 
lection codes designate site of collection followed 
by year of collection (1.e., 97 for 1997; 00 for 2000 
and so on). Seeds were collected usually in May 
just before capsules exploded. Habitats of wild 
source populations varied as to vegetation type, 
precipitation, edaphic features, and other factors. 
Four study sites (LC, PR, FM, and AV) were grass- 
land mixed with forbs in the Antelope Valley of the 
upper Mojave Desert at or near AVCPR, and two 
sites (SR) were shrub-grassland ecotones in the 
foothills of Santa Barbara Co. Of these, only site 
PR (from AVCPR) supported annuals. The remain- 
ing annual populations were from sage scrub hab- 
itat in Riverside (NH, EM), and coastal San Diego 
Counties (CB). Mean temperatures of winter 
months vary among sites (Table 2), and were used 
to guide choice of chilling and incubation temper- 
atures. The domesticated population (Dom) has 
been repeatedly planted and harvested for com- 
mercial seed production in Oregon for at least 20 
years. The original source population for Dom is 
unknown, but thought to have been from southern 
California. The garden population RVO1 was plant- 


210 


MADRONO 


[Vol. 49 


TABLE 2. MEAN DarILy MINIMUM, MAXIMUM, AND MEDIAN TEMPERATURES FOR SOUTHERN CALIFORNIA WEATHER STA- 


TIONS NEAR STUDY POPULATIONS. 


Closest weather Study 
station location population 
Lake Cachuma (1952-2000) SR 
Lancaster (1945-1960) AV, FM, LC, PR 
San Diego (1940-1960) CB 
San Jacinto (1948—2000) NH 


Sun City (1973-2000) EM 


ed before 1990 from an unknown commercial seed 
source. 


Seed germination experiments. Six experiments 
were conducted, with each building on the infor- 
mation gained from previous experiments. General 
procedures are described below. Details of popu- 
lations and experiments are in Table 3, including 
seed age at time of incubation. 


General protocol. After we collected or re- 
ceived seeds, we stored them in the lab at ~22 + 
3°C. Before we obtained seeds from other sources, 
they had been stored under variable conditions: 
S&S Seeds stored seeds at ambient conditions in 
warehouse in coastal, Santa Barbara Co.; Fox and 
Crossman stored seeds in offices; domesticated 
seeds were stored under unknown conditions before 
purchase in June 2000. All wild site collections, 
including those from S&S Seeds, were document- 
ed. 

For all experiments, we chose visibly undam- 
aged, plump seeds of uniform shape, and which had 
obtained a brown to blackish brown color. Seeds 
were always randomly assigned to treatments (de- 
scribed below). For incubation, one filter paper 
(Whatman No. 1, 90 mm) was used per petri-dish 
(Fisher, polystyrene, 90 X 15 mm). Immediately 
before incubation, for treatments Con, S, H, SH, 
and Q (below), we placed seeds on filter paper 
moistened with 2 ml of pH7.5 filter sterilized tap 
water (hereafter ““water’’). For treatments L, W, and 
N, we placed seeds on filter paper moistened with 
2 ml of the treatment solution. Unless otherwise 
indicated, incubation then occurred under a diurnal 
cycle of 12 hr/22°C and 12 hr/12°C (henceforth 22/ 
12°C). We incubated seeds in winter (excepting 
Exp. | & 2) when they would be germinating in 
nature to control for any seasonal cycling of dor- 
mancy under shelf conditions (see Baskin and Bas- 
kin 1998). We aged seeds at least 8 mo in the lab 
except when testing for after-ripening (Exp. 6), 
considered here as a period of embryo maturation 
following seed harvest that occurs before seeds can 
germinate (Nikolaeva 1969). 

For cold treatment, seeds were placed in dishes 
between two filter papers and either left dry or 
moistened (see below). Before cold treatment or in- 
cubation, dishes were individually wrapped in alu- 


Mean min/max temperature °C 
(median temperature) 


December 


3.6/19.3 (11.5) 
—1.8/14.9 (6.5) 
9.5/19.6 (14.5) 
1.1/19.4 (10.3) 
1.4/19.8 (10.6) 


January 


3.7/18.5 (11.1) 
~1.9/13.8 (5.9) 
8.1/18.3 (13.2) 
1.1/18.4 (9.7) 
2.4/18.9 (10.7) 


minum foil to exclude light and placed in sealed 
plastic bags to prevent moisture loss or gas ex- 
change between treatments. Cold treatments and in- 
cubation took place in the dark because light was 
reported to inhibit germination of California poppy 
(Goldthwaite et al. 1971). 

We confirmed that light inhibits germination in 
wild-collected seeds but not in domesticated seeds. 
We tested the effect of light on germination by in- 
cubating 100 moistened seeds from each of four 
wild and one domesticated population in light (no 
foil wrap) vs. dark (foil wrapped) under alternating 
11 hr days and 13 hr nights (temperature ranged 
10—20°C). After 19 days, mean germination under 
light was 11.7%, 3.6%, 3.2%, 5.7%, and 99%, 
while germination in the dark was 29.3%, 16.5%, 
25.8%, 100%, and 100% for populations EMOO, 
PROI, PalaOl, RMO1, and Dom, respectively 
(RMO1 = perennial from coastal San Luis Obispo 
County; PalaOl = annual from inland San Diego 
County). 

At harvest, we scored radicle emergence of at 
least | mm as successful germination. Any remain- 
ing, ungerminated seeds were counted and assayed 
for viability with 2,3,5-triphenyl tetrazolium chlo- 
ride (TTC) (Fischer), a metabolic indicator dye. 
Seeds were pierced with a probe, stained by soak- 
ing overnight in a 1.0% TTC solution (Kitchen and 
Meyer 1992), then dissected under a dissecting mi- 
croscope. A seed was scored as viable if more than 
two thirds of its embryo stained dark pink or red. 
Live, ungerminated seeds were considered dor- 
mant. The proportion of viable seeds in each dish 
was ((#germinated seeds + #viable ungerminated 
seeds)/total # seeds) and proportion germinated was 
(#germinated seeds/#viable seeds), thereby stan- 
dardizing germination to the response of viable 
seeds. 


Fire treatments. Dry fire treatments (S, H, SH) 
were given before any temperature treatments 
whereas wet fire treatments (N, L, W, Q) were giv- 
en after temperature treatments in the first two ex- 
periments. Thereafter, wet fire treatments were giv- 
en before temperature treatments. 

S (dry smoke): Smoke stimulates germination in 
many species and can be applied by various meth- 
ods. Here, we applied cool dry smoke directly to 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY PAI 


2002] 


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212 


dry seeds as per Keeley and Fotheringham (1998b). 
We placed dry seeds in small plastic dishes in a 
large glass chamber. A 500 ml airtight metal can 
was filled 2/3 full with equal portions of air-dried 
Avena fatua L., Bromus madritensis L., B. diandrus 
Roth, Lessingia filaginifolia (Hook. & Arn.) M. A. 
Lane, and Nassella pulchra (A. Hitchc.) Barkworth. 
The vegetation was ignited, and the smoke was 
transferred into the glass chamber with forced air 
through a long tube. The chamber was filled with 
the cooled smoke for 30 seconds, then sealed, and 
seeds were smoked for five minutes. Our prelimi- 
nary experiments with domesticated poppy seeds 
showed that smoking up to 15 minutes does not 
affect seed viability. 

H (heat): A brief heat shock has been shown to 
enhance germination of numerous species (Keeley 
1991; Baskin and Baskin 1998), while killing or 
inhibiting germination of heat-sensitive seeds. To 
explore E. californica’s response to temperatures 
similar to those that may be experienced by buried 
seeds during a fire, we heated dry seeds in open 
Pyrex beakers in a forced air oven at 85°C for 10 
minutes. Preliminary trials using a temperature as- 
say range of 70—115°C demonstrated that viability 
of both wild and domesticated seeds decreased at 
115°C, but not all seed populations lost viability at 
85°C/10 min (Feist and Montalvo, unpublished 
data). 

SH (smoke + heat): A random subset of dry- 
smoked seeds was heated as in H. 

N (nitrogen): Levels of available N as nitrate 
have been shown to increase after fire (Sweeney 
1956; Franco-Vizcaino and Sosa-Ramirez 1997), a 
chemical shift which enhances germination in some 
species (Thanos and Rundel 1995; Baskin and Bas- 
kin 1998). Also, E. californica tends to respond 
positively to disturbance, which in turn is positively 
correlated with nitrogen availability. We used a so- 
lution of 10 mM KNO, (Fischer) which has been 
used successfully to stimulate germination in some 
fire-following species (Thanos and Rundel 1995; 
Keeley and Fotheringham, 1998b) to moisten seeds. 

L (leached charrate): A leachate of the charred 
remains of plant material enhances germination in 
some species (Keeley 1991), and simulates water 
transporting charred plant particles to the seed. We 
prepared charrate for adding to seeds by burning 
equal portions of air-dried vegetation (same as in 
S) until blackened but not ashed. This material was 
ground and added to water (5 g per 100 ml), stirred 
overnight, then filtered through several layers of 
cheesecloth (modified from Keeley 1991). 

W (smoke water): In this smoke treatment, wa- 
ter picks up smoke particles/chemicals and transfers 
them to the seed (Keeley and Fotheringham 1998b). 
We filled an airtight 500 ml can with one of the 
vegetation types (see S above), set the material on 
fire, and forced the smoke to bubble though 1 liter 
of water via a tube. Dried material of each species 


MADRONO 


[Vol. 49 


(S above) was burned in series until all the material 
was completely blackened (1—5 minutes). 

Q (liquid smoke): Commercially produced “‘liq- 
uid smoke’”’ products enhance germination in many 
plant species, and one such product has been used 
successfully in Australia to increase germination on 
mine reclamation sites (Roche et al. 1997b). We 
obtained commercial liquid smoke (Regen 2000 
Smokemaster® Seed Germination Solution, Regen, 
Glasgow, KY) made from passing smoke through 
water and concentrating it. 

In Exp. 1-2, we soaked seeds after their cold 
treatments (or after dry control) in a dilution of 1: 
50 (Regen 2000:water) for 20 hours. This treatment 
was modified for subsequent experiments. In the 
first modification (mQ-1), seeds were soaked in a 
dilution of 1:50 for 12 hours, then air dried at room 
temperature. Seeds were then subjected to appro- 
priate cold treatments after smoke treatment (Exp. 
3—4). The second modification (mQ-2) explored the 
effects of liquid smoke concentration (Exp. 5, be- 
low). Although the manufacturer recommends us- 
ing a 1:10 dilution, a 1:50 dilution had been used 
in order to decrease the risk of harming seeds with 
high liquid smoke concentrations. To assess the ef- 
fects of dilution strength on germination, we soaked 
seeds 12 hr in either a 1:50, 1:25. or 1:10 dilution, 
then transferred wet seeds to the appropriate cold- 
shock treatment (see below). 

Con (control for fire treatments): No fire treat- 
ments were given, and seeds were subjected to the 
appropriate temperature and moisture treatments 
(below). 


Temperature and moisture treatments. 

Cold: Seeds in petri dishes between two filter pa- 
pers were subjected to prolonged cold temperatures 
in a cold chamber (ranging from two to eight 
weeks) while either moist or dry. Seeds were moist- 
ened with 2 ml of fluid (moist-cold), 3 ml of fluid 
(wet-cold), or left dry (dry-cold). Wet-cold treat- 
ment provided free water for rapid imbibing, moist- 
cold provided for slower imbibing and high relative 
humidity, while the dry method tested whether pop- 
py seeds can be affected by cold treatment when 
dry. In addition, Exp. 5 tested the effects of a brief 
cold-shock (48 hours at 2°C) instead of prolonged 
cold treatment. 

Cold treatment was intended to occur near or be- 
low 4°C. Problems with one cold room resulted in 
higher cold-treatment temperatures during Exp. 3 
(see below), averaging near 7°C (range 4—9°C), 
which resulted in a positive germination response 
and provided the impetus to test variation in incu- 
bation temperature in Exp. 5. 

No-cold (control for cold and moisture treat- 
ments): For dry, no-cold conditions, henceforth 
‘“‘no-cold’’, we stored seeds dry at room tempera- 
ture until moistened and incubated. 


Experiments. All experiments included a control 
treatment, for which seeds received no fire or cold 


2002] 


treatments prior to incubation. At the time of in- 
cubation, those seeds collected in spring 2001 were 
between 2—7.5 months old (henceforth ‘“‘fresh 
seeds’’), seeds collected in 2000 were between 8— 
10 months old (henceforth “‘young seed’’), and all 
other wild seeds were > two years old (henceforth 
‘“‘aged seed’’). 

Experiment | and Exp. 2 were run concurrently 
and were overlapping subsets of one experiment. In 
Exp. 1, we examined germination response of three 
‘“‘aged seed”’ populations and domesticated seeds to 
cold treatment at three different moisture levels 
(wet, moist, and dry) in combination with five 
‘“‘wet’’ fire-effects treatments (Table 3). The wet fire 
treatments were applied after seeds had been cold- 
treated. 

In addition, for Exp. 2 we added three “‘dry”’ fire 
treatments (S, H, and SH) under three cold treat- 
ments (moist-cold, dry-cold, no-cold treatment) and 
on seeds of two additional wild populations (PR95 
and FM96, for a total of six populations). The ap- 
plication of dry fire treatments before seeds were 
chilled, simulated the natural sequence of events in 
the wild. The chilling temperature ranged from 2-— 
4°C; most often close to 2°. For both Exp. 1 and 
Exp. 2, incubating seeds were checked using dim 
green lights after 3, 8, 13, and 20 d of incubation. 
No increase in germination was seen after 13 days. 

In Exp. 3, we investigated five aged and one 
young seed population, increased the number of 
replicates from three to five petri-dishes to increase 
statistical power, and examined the response of wet 
and dry smoke (mQ-1 and S) to a longer 8 wk cold 
treatment and a more natural sequence of smoke 
and cold events (Table 3). Both the S and mQ treat- 
ments were applied before seeds were subjected to 
moist cold treatments. The cold room cycled be- 
tween 4—9°C, and was mostly just below 7°C. After 
cold treatment, any germination was tallied, and 
ungerminated seeds were transferred into new dish- 
es with fresh filter paper, moistened with 2 ml wa- 
ter, and incubated for 20 days before scoring ger- 
mination and viability. 

Experiment 4 assessed germination of young 
seeds collected in the same year to investigate their 
response to smoke and cold treatments (Table 3). 
Data from Exp. 3 suggested that young seeds (<1 
yr) may be less dormant or have dormancy more 
easily broken than aged seeds (2+ yr) which could 
complicate experiments with mixed age seeds. Ex- 
tended shelf storage can trigger secondary dorman- 
cy in some species while decreasing dormancy in 
others (Roche et al. 1997a; Baskin and Baskin 
1998). 

In Exp. 5 we tested different concentrations of 
liquid smoke (mQ-2), different incubation temper- 
atures, and the effect of a brief cold-shock on ger- 
mination and viability of two populations, one with 
young seeds and one with aged seeds (Table 3). 
Different incubation temperatures (22/12, 18/6, and 
7.5°C constant) were tested to determine if lower 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 


2B, 


incubation temperatures improved germination, 
given that germination was possible and sometimes 
higher at quite low temperatures (Exp. 3). In ad- 
dition, we subjected half the seeds to a 48 hr cold- 
shock at 2°C instead of prolonged cold because 
short cold treatments are sometimes sufficient for 
either breaking or inducing dormancy and are eco- 
logically realistic within E. californica’s range. Fi- 
nally, to discern between the possible effects of 
soaking which may leach germination inhibitory 
chemicals from seeds, and the effects of wet fire 
treatments, we added a wet-control treatment (12 hr 
soak in water). 

We used annual population EMOO for the full 
Exp. 5. In addition, seeds from the deeply dormant 
NH98 population were run with only a wet-control 
and with 18/6°C as the incubation temperature 
(based on incubation temperature with highest ger- 
mination in Exp. 4) to see if germination could be 
improved with higher or lower concentration of lig- 
uid smoke. 

We ran Exp. 6 to test if recently collected 
“fresh”? wild and domesticated California poppy 
seeds germinate at different rates with and without 
smoke treatment over the course of several months 
after fruit dehiscence. If seeds require lengthy after- 
ripening or if in the lab they experience an annual 
cycle of dormancy followed by conditional dor- 
mancy or loss of dormancy as expected under nat- 
ural conditions, then these behaviors could affect 
the outcome of germination experiments run at dif- 
ferent seed ages or times of the year. We collected 
seeds into paper envelopes in May 2001 from four 
wild populations (perennial FM and SR; annual NH 
and PR, Table 3), and from garden plants in Riv- 
erside (RV—started from commercial seed source 
> 10 years earlier, Table 3). Six weeks after col- 
lection and shelf storage, we randomly sorted seeds 
into microcentrifuge tubes, and then tested at 2, 4, 
and 7.5 months (herein “‘8’’ mo) following seed 
harvest. We incubated controls and liquid smoke- 
treated seeds (mQ-1) in the dark at alternating 
18°C/6°C for 11/13 hours for 15 days and scored 
germination and viability. Subsamples of smoked 
seeds aged 4 mo and 8 mo were given moist-cold 
treatments (4 wk at 3.5°C and 2 wk at 3°C, respec- 
tively) before incubation. 


Data analysis. Data were analyzed with ANOVA 
using Proc GLM of SAS (Release 6.12). In all mod- 
els, source POPULATION was a random effect 
while COLD (e.g., cold vs no-cold), FIRE (e.g., 
smoke, heat, nitrate, water, etc.), or AGE treatments 
were fixed effects. Response variables included 
proportion of viable seeds germinated/dish, and the 
proportion of viable seeds out of total seeds/dish. 
Dishes were the replicates. Before analysis, all data 
were angularly transformed (arcsin(proportion)'”) 
to enhance the normality of the residuals. We used 
either Duncan’s Multiple Range test or Tukey’s test 
for posteriori comparisons among means. In mod- 


214 


els where we found significant interactions among 
main effects, we ran separate analyses on each pop- 
ulation or treatment depending on structure of the 
interaction and the particular question. Given the 
mixed model ANOVAs, we used the RANDOM 
statement in SAS to calculate the denominator 
mean squares for F-tests using the Satterwaite mod- 
el. 


RESULTS 


General patterns. Over experiments, seed dor- 
mancy (Table 4) and the effect of potential dor- 
mancy breaking treatments varied substantially 
among years of collection and among populations 
(Tables 5, 6; Fig. 1-5). The domesticated commer- 
cial (Dom) and garden (RV) populations were the 
only ones with no seed dormancy (Table 4). Other 
populations ranged from 41% to 100% dormant un- 
der control conditions. In general, young seeds col- 
lected in 2000 had higher germination in controls 
and higher germination following treatment with 
liquid smoke than aged seeds (mean 65% vs. 92%; 
Table 6). 

There were differences in seed dormancy among 
young (year 2000) collections from the Antelope 
Valley, with much lower dormancy of the perennial 
AVOO0O and FMOO than the annual PROO. Aged seeds 
had uniformly high dormancy (range 93—100%). 

Seed viability also varied among wild collections 
and ranged from 54—100% following treatments 
(Table 4). Populations NH98 and FM96 showed 
different levels of viability between experiments, 
possibly because of differences among technicians 
in sorting bulk seeds to be used in experiments. The 
range in viability shows the importance of basing 
% germination on live seeds rather than total seeds. 

There were highly significant effects of fire and 
cold treatments and significant population effects. 
In addition, the many significant two and three way 
interactions between FIRE, COLD, AGE, and POP- 
ULATION main effects (Table 5) indicate variation 
among populations in response to at least some 
treatments. Because of this complexity, we present 
results of each experiment separately and break up 
analyses to examine interactions and effects of fire, 
cold, and age treatments. 

Given their lack of seed dormancy, we did not 
statistically compare Dom and RV with wild pop- 
ulations. The domesticated seeds had nearly 100% 
germination and viability in all treatments except 
those involving heat. In addition, unlike the dor- 
mant-seeded populations, Dom seeds germinated 
during cold treatment at 2—4°C. Separate analysis 
of population Dom seeds under the no-cold regime 
showed that there was no significant difference 
among the Con, S, N, W, L, or Q treatments. How- 
ever the H and SH treatments reduced germination 
significantly suggesting that heat treatment inhibit- 
ed germination and even killed some seeds. Via- 
bility of seeds from the H and SH treatments ranged 
from 64—100%, compared to 100% in controls. 


MADRONO 


[Vol. 49 


Experiment 1—Effect of wet-fire and cold treat- 
ments (moist-cold, wet-cold, dry-cold, and no-cold). 
Analysis of germination from the three study pop- 
ulations of different regions (annual NH98, peren- 
nial LC98, and annual CB97) revealed significant 
main effects and interactions except POPULATION 
and FIRE x POPULATION (Table 5). When ger- 
mination was analyzed for each population sepa- 
rately, FIRE, COLD and FIRE X COLD were still 
significant for each population (all P values = 
0.0113). This significant interaction shows that 
each population responded to the set of treatments 
differently, obscuring whether any particular cold 
treatment or fire treatment resulted in the highest 
germination. However, for all populations, the no- 
cold regime resulted in the highest mean germina- 
tion (Fig. 1), suggesting that the 2°C cold treatment 
was cold enough to slightly inhibit germination in 
the three populations examined. 

We also analyzed germination response separate- 
ly by level of cold treatment and found no signifi- 
cant FIRE < POPULATION interactions, and in 
the wet-cold analysis we found no significant FIRE 
or POPULATION effects (Fig. 1). In the moist-cold 
treatment, annual NH98 had significantly higher 
germination than perennial LC98 and annual CB97 
seed populations, while in the no-cold regime CB97 
seeds had significantly higher germination than 
LC98 and NH98 (Fig. 1). Inland NH98 seeds ap- 
pear to germinate under cold conditions more read- 
ily than coastal CB97 seeds. There were significant 
effects of FIRE treatment only in the dry-cold and 
no-cold regimes, potentially because liquid fire 
treatments were applied after chilling in moist-cold 
and wet-cold. In no-cold and dry-cold, treatment Q 
resulted in the highest germination, although Q and 
W were not statistically different under the no-cold 
control conditions (Fig. 1). 

With ANOVA of Experiment 1, we examined the 
effect of treatments on seed viability to reveal if 
some treatments harmed seeds. We found no sig- 
nificant main effects, but the COLD X FIRE xX 
POPULATION interaction was significant (F534 129 
= 1.67; P < 0.04), suggesting that seed viability of 
different populations was affected differently by the 
cold and fire treatments. After running viability 
data separately by population, no significant effects 
of COLD or FIRE treatments were found for CB97. 
Population LC98 showed a significant effect of 
FIRE treatment (Fy 3, = 3.07; P = 0.027) and a 
FIRE X COLD interaction (Fy, 4 = 2.26; P = 
0.028). COLD had a significant effect on NH98 
(F’; 49 = 6.76; P < 0.001), with the no-cold regime 
more viable than all other cold treatments. No other 
effects were significant. Overall, the wet-cold treat- 
ment resulted in the lowest germination in NH98 
(Sigal): 

When viability was analyzed for each cold re- 
gime separately, the only significant effect of POP- 
ULATION was in the no-cold treatment (Fy , = 
4.73; P = 0.044), with NH98 having greater via- 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 25 


2002] 


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216 MADRONO [Vol. 49 


TABLE 5. ANOVA TABLES FOR GERMINATION RESPONSE IN EXPERIMENTS 1—6. For Experiments 1—4 and 6, full models 
are shown; COLD refers to cold treatments, and FIRE refers to fire treatments (see Table 3). For exp. 5, only results 
for population EMO0 are presented; SHOCK, INCUBATION, and FIRE refer to those treatments as noted in Table 3 
under Cold, Incubation, and Fire treatments, respectively. Num and Den df are numerator and denominator degrees of 
freedom, respectively. Exp. 5 is a fixed effects model so Den df is that of error term. 


Source Num df Den df lia IP < 
EXPERIMENT 1 
Cold 3 6 49.89 0.001 
Fire 4 8 17.41 0.001 
Population 2 Do 1.40 0.325 
Cold X Fire 12 24 6.76 0.001 
Cold < Population 6 24 4.94 0.002 
Fire <X Population 8 24 OFT: 0.635 
Cold X Fire X Population 24 120 1.67 0.038 
Error 120 
EXPERIMENT 2 
Cold 1 4 12.47 0.024 
Fire 7 28 16.46 0.001 
Population 4 4.6 1.67 0.302 
Cold X Fire 7 28 9:32 0.001 
Cold X Population 4 28 5.48 0.002 
Fire X Population 28 28 1.41 0.184 
Cold X Fire X Population 28 160 DS 0.001 
Error 160 
EXPERIMENT 3 
Cold 2 10 alg) 0.001 
Fire 2 10 203.07 0.001 
Population 5 V2 2B 3D) 0.001 
Cold X Fire 4 20 2.00 0.096 
Cold < Population 10 20 1.34 . 0.276 
Fire X Population 10 20 1.41 0.244 
Cold X Fire X Population 20 216 Jiepoi] 0.002 
Error 216 
EXPERIMENT 4 
Cold 2 6 9).35) 0.001 
Fire 2 6 231.81 0.001 
Population 3 6.8 5.26 0.034 
Cold X Fire 4 12 6.18 0.001 
Cold < Population 6 12 DES 0.079 
Fire X Population 6 12 20.09 0.001 
Cold X Fire X Population 12 144 0.59 0.847 
Error 144 
EXPERIMENT 5 
Shock 1 4.32 0.040 
Incubation 2 7.01 0.001 
Fire 4 220.20 0.001 
Shock X Incubation 2 ele 0.328 
Shock X Fire 4 0.45 0.775 
Incubation X Fire 8 0.24 0.983 
Shock X Fire X Incubation 8 0.96 0.468 
Error 120 
EXPERIMENT 6 
Age group 8.02 8.05 0.012 
Fire (Con vs. mQ) 4 23.08 0.009 
Population 7.74 2S 0.144 
Age X Fire 8.08 9.04 0.009 


Age X Population 

Fire < Population 

Age X Fire X Population 
Error 


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2002] 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 


PN] 


TABLE 6. SUMMARY OF MEAN % GERMINATION OF YOUNG VS. AGED SEEDS COMPARING RESPONSE TO DRY SMOKE (S), 
LIQUID SMOKE (MQ), AND WATER CONTROLS ACROSS EXPERIMENTS. Population abbreviations as in Table 1. No-cold = 
germination under no-cold (and no cold-shock) treatments. Moist-cold = moist-cold treatments as in Table 3. Values 
in bold represent higher germination under moist-cold compared to no-cold for the same smoke treatment. * = liquid 
smoke concentrations of 1:10 (all others were 1:50); ** = Exp. 2 had liquid smoke applied after cold treatment. Dashes 


represent absence of treatment. See Table 3 for seed age. 


Moist-cold 

Pop Liquid 

code Exp Control smoke 
LC98 2 LS —_ 
AVOO 3 54.5 86.8 
AVOO 4 45.8 88.2 
PR95 2 0 —_* 
PR96 3 0 Las 
PROO 4 14.0 70.8 
NH98 2 24.4 —_ 
NH98 3 23.8 47.5 
NH98 5 — — 
NHOO = 3335 97.2 
FM96 2 0 rE 
FM96 3 32 31.9 
FMO0O - 44.5 77.8 
EM98 3 28.2 59.4 
EMO00 5 — — 


bility than LC98 and CB97 (Tukey’s test, alpha = 
0.01). Thus, the significant COLD FIRE x POP- 
ULATION effect of the full model analysis is due 
in part to population NH98 having much improved 
viability in the no-cold treatment. This small neg- 
ative effect of 2°C on viability for NH98, suggests 
that any ungerminable seeds in the process of via- 
bility degradation may have died more readily than 
such marginal seeds of other populations, thus in- 
fluencing the germination rate for NH. Otherwise, 
there was little effect of cold or other treatments on 
viability. 


Experiment 2: Effect of all fire and cold treat- 
ments (moist-cold vs. no-cold). ANOVA of Exper- 
iment 2 germination data showed that all effects 
were significant except POPULATION and FIRE X 
POPULATION (Table 5). We then broke the anal- 
ysis down by the two cold treatments because of 
the significant 3-way interaction of main effects. 

In both the moist-cold and no-cold regimes, 
FIRE, POPULATION and FIRE X POPULATION 
were all significant (P < 0.001), so the analysis was 
further broken down by population for each cold 
regime separately. Under moist-cold, for all popu- 
lations except NH98, we found a significant effect 
of FIRE treatment on germination (all F, ,, > 4.13: 
P < 0.009), with S producing significantly higher 
germination. For NH98, mean germination was 
highest for S, but non-significant (Fig. 2). A Ken- 
dall’s analysis of the rankings of fire treatment 
within each population showed a strong association 
between fire treatment and germination (0.01 < P 


Mean % germination 


No-cold 

Dry Liquid Dry 
smoke Control smoke smoke 
55.4 MB) 37.8 29.4 
70.6 52.6 83.3 60.2 
68.7 58.9 82.9 49.7 
aS 6.3 46.5 44.0 

0) 0 3.6 0 
48.7 ihe s7/ 65.5 215 
24.1 11.9 41.9 19.3 
25.6 11.8 33.8 2S 
= Fh GEN 34.0* — 
132 29.9 94.9 oy es 
24.8 3.8 10.5 10.7 

3.6 4.8 25.6 6.6 
735 53.9 69.7 59.4 
36.6 9.3 65.2 [5:2 
— Sil 94.6 = 


< 0.001). Treatment S had the highest overall rank- 
ing, HS and H had the lowest overall rankings (in- 
dicating heat reduces germination), while all other 
treatment rankings were very similar. 

Under the no-cold regime, germination of all 
populations except FM96 was significantly affected 
by FIRE treatment (all F,,, => 6.2;.P.= 0.001)..A 
Kendall’s analysis of the rankings of treatments 
within each population showed a strong association 
(P < 0.001) between fire treatment and germina- 
tion. For all populations, Q produced the highest 
mean germination, while S and W tied for second 
place (Fig. 2). Averaged over the five populations, 
the no-cold controls germinated only 25% as well 
as seeds treated with liquid smoke (Q). The H and 
HS treatments were ranked lowest overall. Again, 
the heat treatments reduced germination. Germi- 
nation of H treated seeds for the five populations 
averaged 38% of control seed germination. Q per- 
formed best in the no-cold regime, while S_ pro- 
duced the highest germination in the cold regime, 
again suggesting the effect of Q would improve if 
applied before chilling. This hypothesis was veri- 
fied in Exp. 3, below. 

ANOVA of Exp. 2 viability data showed that 
POPULATION, FIRE, and COLD were all signifi- 
cant effects (POP F, ,; = 5.76; P = 0.049; FIRE 
Fa 93, = 4.53; P = 0.002; COLD F, , = 12.4; P = 
0.024). There were no significant interactions (all 
P >0.11). In the full model, viability was unaffect- 
ed by all treatments except H and SH, which sig- 
nificantly decreased viability (Duncan’s Multiple 


218 MADRONO [Vol. 49 
= 100 Moist-cold 
o 80 
£ 60 
E 40 
0) 
© 20 
xs (0 
Con L N Q WwW Con L N Q WwW Con L N Q W 
LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 
No-cold a 
c 
no) 
—s 
© 
= 
= 
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Con L N Q WwW Con L N Q Ww Con L N Q WwW. 
LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 
Dry-cold 
S 100 
= 80 
£ 60 
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0) 
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Con L N Q WwW Con L N Q WwW Con L N Q WwW 
LC98 LC98 LC98 LC98 AV98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 
c 100 Wet-cold 
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= 
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Con L N Q Ww Con 
LC98 LC98 LC98 LC98 LC98 CB97 CB97 CB97 CB97 CB97 NH98 NH98 NH98 NH98 NH98 


Fic. 1. 


N Q WwW Con L N Q Ww 


Experiment 1 backtransformed germination data, analyzed for each population within each cold treatment. 


Lower and upper case letters represent significant differences between treatments and populations, respectively, based 
on Tukey’s tests (alpha = 0.05). Within each cold treatment, populations or treatments that share a letter are not 
significantly different. In moist-cold, NH98 had higher germination than CB97 and LC98, but there were no significant 
effects of fire treatment. Error bars = 1 SD (n = 3). Abbreviations: Con = control, L = leached charrate, N = nitrogen 


(KNO,;), Q = liquid smoke, and W = smoke water. 


Range Test, using alpha = 0.05). Viability of SH 
and H treated seeds for the five populations aver- 
aged 6% lower than viability of control seed, com- 
pounding the effect of reduced germination. Over- 
all, FM96 had significantly higher viability than all 
other populations, and CB97 had significantly low- 
er viability than all but population LC98 (Tukey’s 
test, alpha = 0.01) (Table 4). 

For remaining experiments, analysis of seed vi- 
ability will not be presented in detail. Additional 
treatments had little affect on seed viability. 


Experiment 3: Effect of dry smoke, liquid smoke 
and moist-cold, dry-cold, and no-cold (smoke be- 
fore cold). In Exp. 3, many seeds germinated un- 
expectedly during moist cold treatment, including 
some FIRE treatment controls. For populations that 
germinated during cold treatment, very few of the 
remaining seeds germinated after shifting to the 


warmer incubation period. Therefore, in ANOVA 
of total % germination, the main effect of COLD 
represents three treatment levels: moist-cold = in- 
cubation at 4—9°C; no-cold = incubation at 22/ 
12°C; and dry-cold = incubation at 22/12°C follow- 
ing 8 wk dry cold treatment. 

ANOVA of total % germination showed a sig- 
nificant COLD X FIRE X POPULATION interac- 
tion (Table 5). When data were broken up by pop- 
ulation, treatment mQ (modified liquid smoke) pro- 
duced significantly greater germination than the 
other treatments in all populations (averaging over 
fire treatments shown in Fig. 3), but there was a 
significant COLD X FIRE interaction within EM98 
and NH98. EM98 and NH98 were the only popu- 
lations significantly affected by COLD (P < 
0.0024), and in both cases moist-cold produced 
higher germination than dry-cold and no-cold (Fig. 


2002] MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 219 
= 
= 100 - MH no-cold (11 moist-cold 
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Fic. 2. Experiment 2 backtransformed germination data, analyzed separately for moist-cold and no-cold treatments 


within each population. Lower and upper case letters indicate significant differences (Tukey’s test, alpha = 0.01) 
occurring between treatments within no-cold and moist-cold regimes, respectively. FIRE treatments that share a letter 
are not significantly different. Error bars = 1 SD (n = 3). Abbreviations: as in Fig. 1 plus S = dry smoke, H = heat, 
and SH = smoke plus heat. 


Evidently, germination decreased as seed age in- 
creased. 


3). In a separate analysis by cold regime, there was 
a significant interaction of FIRE x POPULATION 
in no-cold and dry-cold, likely due to populations 


CB97 and EM98, for example, having much larger 
differences between mQ and control treatments 
than population AVOO. However, in all cold treat- 
ments, mQ produced the best germination, and was 
highest for AVOO, then EM98, and iowest for PR96. 


Experiment 4: Effect of dry and liquid smoke vs. 
cold treatments on ‘‘young”’ seeds. ANOVA of ger- 
mination data showed that all effects were signifi- 
cant except COLD X POPULATION and COLD xX 
TREATMENT X POPULATION (Table 5). Even 


220 MADRONO [Vol. 49 
8 © 100 

= E 60 

8 8 40 

xe sx 20 

~ E b s i<e) 0 

oO (ep) 

O Ne Me = 

m m m 


OS 


Cc 
Con Con Con Q Con Con Con Ss S S mQ mQ mQ 


NH98 % Germination 
aS 
(@) 


Cc NC MC DC NC MC DC NC MC DC 
Q Con Con Con Ss S S mQ mQ mQ 


AV00 % Germination 


Con Con Con Ss S Ss 


2 E 

ie 8 

E € 

(i) = 

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2 g 

fi NC mc be NC me De NC me OC f NC MC DC NC MC DC NC MC DC 
Con Con Con S&S S Ss mQ mQ mQ Con Con Con S&S Ss Ss mQ mQ mQ 

Fic. 3. Experiment 3 backtransformed germination data, analyzed for each population. Upper case and lower case 


letters indicate significant differences between COLD or FIRE treatments, respectively (Tukey’s test, alpha = 0.05). 
COLD or FIRE treatments that share a letter do not have significantly different germination. For populations EM98 
and NH98, no Tukey’s tests were run due to significant COLD X FIRE interactions. Error bars = 1 SD (n = 5). 
Treatment abbreviations: Con = control, NC = no-cold, MC = moist cold, DC = dry cold, S = smoke, mQ = modified 


liquid smoke. 


though the strength of different TREATMENTS 
varied by population and cold treatment, the 
TREATMENT controls had the lowest mean in 11 
of 12 comparisons of population/cold combinations 
(Fig. 4). The significant interaction of TREAT- 
MENT X POPULATION appears to be due to the 
annual populations (NHOO and PROO) having high- 
er seed dormancy and larger differences among 
treatment means than the perennials (AVOO and 
FMOO0). 

Because of the significant interactions, we used 
ANOVA to analyze germination separately for each 
population. For AVOO, there was a significant in- 
teraction of COLD X TREATMENT (F;, 3, = 3.91; 
P = 0.009), however TREATMENT was highly 
significant (Ff, 3, = 48.70; P < 0.001). Germination 
of AVOO in moist-cold and dry-cold was lowest in 
fire controls, higher with S, and highest with mQ, 
but position of no-fire Con and S were switched 
under no-cold (Fig. 4). For FMOO, S and mQ were 
not significantly different from each other, but both 
smoke treatments were significantly higher than 
controls. The trend seen in AVOO was even stronger 
in the two annual populations (PROO and NHOO; 
Fig. 4). In all three cold treatments, mQ resulted in 
significantly higher germination than S, and S re- 
sulted in higher germination than Con. Cold treat- 
ments were significant only for annuals (PROO 
F, 3 = 5.73; P = 0.007; and NHOO F, ;, = 4.80; 
P = 0.014) with moist-cold producing significantly 
higher germination than dry-cold in both cases, and 


also higher than no-cold for PROO (Fig. 4), sug- 
gesting a small synergistic effect of cold and 
smoke. 


Experiment 5: Effect of different liquid smoke 
concentrations (mQ), incubation temperatures, and 
cold-shock. ANOVA was run on the two popula- 
tions separately because NH98 underwent only a 
subset of treatments. The full model analysis of 
EMO0 showed significant effects of SHOCK, IN- 
CUBATION TEMPERATURE, and TREATMENT, 
with no significant interactions (Table 5). The 3 mQ 
concentrations yielded much higher germination 
than the no-fire controls, and all concentrations per- 
formed equally well (Fig. 5). In addition, the two 
colder incubation temperatures (7.5°C and 18/6°C) 
resulted in higher germination than warmer incu- 
bation (22/12°C; Tukey’s test, alpha = 0.05). Seeds 
readily imbibed fluids and there was no difference 
between wet and dry controls, indicating leaching 
of inhibitors is not a factor in success of liquid pre- 
germination treatments. Finally, cold-shock de- 
creased germination of EMO0 relative to no-shock 
(Fig. 5) but the difference was slight. 

ANOVA analysis of the NH98 germination re- 
vealed a significant effect of TREATMENT only 
(F; 35 = 11.43; P = 0.001). Unlike in EMOO, ger- 
mination of NH98 seeds was significantly higher as 
smoke concentration increased (Fig. 5; Tukey’s test, 
alpha = 0.05), and cold-shock had no effect on ger- 
mination (F 3, = 0.01; P = 0.934). 


2002] MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 29) || 
S 5 400 C Cc Cc B B B 
© € 380 
€ E 60 
© 5 
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x = 
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Con Con Con S S S mQ mQ mQ Con Con Con S S S mQ mQ mQ 
NC DC MC NC DC MC NC DC MC NC DC MC NC DC MC NC DC MC 
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Con Con Con S S S mQ mQ mQ Con Con Con S Ss S mQ mQ mQ 
NG DG SMG =NG. IDG yaMG TING | DG” IMC NC DC MC NC DC MC NC DC WMC 


Fic. 4. Experiment 4 backtransformed germination data, analyzed for each population. Upper case and lower case 
letters indicate significant differences (Tukey’s test, alpha = 0.05) among smoke or cold treatments, respectively. Smoke 
or cold treatments that share a letter do not have significantly different germination. For populations AVOO no Tukey’s 
test was run due to significant COLD X smoke TREATMENT interaction. Error bars = 1 SD (n = 5). Abbreviations 


as in Fig. 3. 


Experiment 6: Effect of aging 2—8 months from 
collection and of liquid smoke (mQ-1) on germi- 
nation: The garden RVO1 seeds germinated to 99% 
under both control and liquid smoke conditions at 
2 mo of age so we did not continue to examine 
them at 4 mo and 8 mo, nor was RV included in 
Statistical analysis. We ran ANOVA separately on 
each wild population because of significant 2-way 
interactions between population, smoke treatment, 
and seed age (Table 5). The interactions indicate 
that populations respond differently to both treat- 
ment and aging, either in degree or direction of 
response. Controls of FMO1, NHO1, and PROI had 
significantly lower germination at all ages than did 
mQ-1 treated seeds (Fig. 6). Except for FMO1, 2- 
mo old seeds had significantly lower germination 
than older seeds, but there was usually no differ- 
ence between germination of 4 mo and 8 mo-old 
seeds. The strongest evidence for after-ripening was 
for perennial populations from Santa Barbara 
County (SR10O1 and SR601) where there was no 
increase in germination of mQ-1 treated seeds at 
age 2 mo, but thereafter mQ-1 treatment resulted in 
Significant increases. This result, together with 
higher germination of both controls and smoked 
seeds at 4 and 8 mo age suggests that ~60% of 
seeds required an after-ripening period of > 2 mo 
before they were capable of dormancy brake and 
germination (Fig. 6). We also looked at effects of 
mQ-1 and a cold treatment at 3—3.5°C on 4 and 8 
mo-old seeds (Fig. 6). A separate ANOVA of treat- 
ment (cold + mQ-1 vs. mQ-1) and age (4 vs. 8 mo) 
on germination, showed no significant effect of 
cold treatment or age in PROI or FMO1. However, 


cold significantly increased germination in NHO1 
and SRIO1 seeds and decreased germination in 
SR601. Germination was significantly higher for 4 
mo-old seeds in these later three populations. Thus 
cold treatment had an inconsistent effect on ger- 
mination relative to just smoke-treated seeds both 
among age groups and among populations. 


DISCUSSION 


Our study populations came from a set of con- 
trasting southern California environments. We used 
seeds of both annual and perennial life-histories 
and with different levels of seed dormancy. We ex- 
amined whether cold treatment, some specific com- 
ponents of fire, or some combination enhances ger- 
mination of dormant seeds, and found important di- 
rect effects of smoke. This is important because his- 
torically, fire has been naturally occurring and 
periodic in both shrub and grasslands of California, 
(Parsons 1981; Keeley 1991) where poppies occur. 
Additionally, for thousands of years prior to Euro- 
pean settlement, indigenous tribes of southern Cal- 
ifornia managed many areas with fire to encourage 
growth of animal forage as well as certain food and 
textile plants (Bean and Lawton 1973; Timbrook et 
al. 1982; Lewis 1993). With such repeated exposure 
to fire, we expect that many species that have not 
been considered fire followers may, nonetheless, re- 
spond to chemical constituents of fire. Even in de- 
sert areas that do not have a history of frequent 
wildfire, the incidence of fire is increasing due to 
fuel loading by invasive grasses coupled with in- 
creasing human activity (Brooks 1999), underscor- 


DDD MADRONO 


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80 B 5 
60 4 2 a b b 


% Germination 
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Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 
NCS NCS NCS NCS NCS CS CS CS CS _ CS 


EMO00 @ 18/46 C A A A A A A 
a a a b b b 
100 B B 
80 B B 


a a 


% Germination 
£& OD 
oo 
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oOo 


Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 
NCS NCS NCS NCS NCS CS CS CS CS CS 


EM00@ 75 C ‘ mA ES ude: 


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a a 


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oO 


% Germination 
bh 
(o) 


Cn-D Cn-W 1:50 1:25 1:10 Cn-D Cn-W 1:50 1:25 1:10 
NCS NCS NCS NCS NCS CS CS CS CS _ GCS 


NH98 @ 18/6 C 


100 ie B AB A a a a a 
80 a a a a 


% Germination 
L 
oO 


Cn-W 1:50 1:25 1:10  Cn-W_ 1:50 1:25 1:10 
NCS NCS NCS NCS CS CS CS CS 


Fic. 5. Experiment 5 backtransformed germination data, 
analyzed for each population and incubation temperature 
separately. Upper case letters indicate significant differ- 
ences (Tukey’s test, alpha = 0.05) among the three liquid 
smoke (mQ) concentrations. Lower case letters designate 
ANOVA result for differences between cold-shocked (CS) 
and no-shock control (NCS) seeds. Error bars = 1 SD (n 
= 5). Solid bars are NCS and open bars are CS treatments. 
Cn-D = dry control; Cn-W = wet control. 


120 
AcQ 


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SR101 —————_» 


[Vol. 49 


ing the need to understand the effects of fire on 
plant reproduction. 


Fire treatments. Smoke applied dry or wet was 
successful in substantially increasing germination 
of all dormant seeded populations, sometimes by as 
much as three to four fold over controls. Dry smoke 
(S), smoke water (W), and liquid smoke (Q, Qm- 
1, Qm-2) all improved germination substantially 
over controls showing that smoke can break dor- 
mancy when delivered to seeds in various ways. 
Because seeds readily imbibed water, the mecha- 
nism for the effect of smoke on germination in this 
species may not involve changing the structure of 
the seed coat cuticle as has been detected for Em- 
menanthe penduliflora (Egerton-Warburton 1998; 
Keeley and Fotheringham 1998a). It is possible that 
ethylene or other components of smoke influence 
seed germination in California poppy, but this 
needs further study. Both dry smoke and smoke 
water contain ethylene (Sutcliffe and Whitehead 
1995), which is known to promote germination by 
several different mechanisms and can sometimes 
Overcome secondary dormancy or particular tem- 
perature requirements for germination (Corbineau 
and Come 1995; Baskin and Baskin 1989). 

Commercially produced liquid smoke gave the 
best results and outperformed dry smoke except 
when applied after chilling. In each case, the seeds 
responded best to the most natural sequence of 
events. Any “‘smoke”’ event would likely occur in 
the dry season before the winter rains or as a result 
of rain carrying smoke particles from the soil sur- 
face to the seedbank in late fall or early winter. 
Results were more erratic for dry smoke than for 
liquid smoke treatments, possibly because adsorp- 
tion of dry smoke among experiments is more dif- 
ficult to control. 

Other fire treatments were less effective or even 
inhibited germination. We found no significant ben- 
efit of charred plant leachate (L), or nitrate (KNO,) 
on germination. It is possible that charate and NO, 


I 
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Experiment 6 backtransformed germination data. Data are arranged by population, treatment, and seed age (in 


months) along the x-axis. Seeds aged 7.5 weeks are noted as “‘8”’. Data for liquid smoke followed by cold treatment 
(cQ) were compared with mQ-1 treatment in a separate analysis. Week 4 and week 8 cQ seeds were exposed to 4 


weeks at 3.5°C and 2 weeks at 3°C, respectively. Error bars 


— I SDi@) = 5): 


2002] 


could be more effective if applied before a cold 
treatment, but this was not tested. Exposure to 85°C 
for 10 minutes (H and SH) generally reduced ger- 
mination of viable seeds, decreasing it to about a 
third of control values. Viability of heat-treated 
seeds also decreased by an average of 6%, stressing 
a need for further studies on heat sensitivity of this 
species. 

We expect that seeds near the soil surface where 
temperatures are higher will be largely killed or in- 
hibited by the heat of fire. In low intensity burns in 
chaparral, temperatures ranged from about 60°C at 
a depth of 7-8 cm, about 82°C at 2—3 cm, about 
100°C at 1-2 cm deep, to over 180°C at the surface. 
Also, as fire intensity increased with increase in 
fuel load, temperatures deep into the seed bank be- 
came higher (Moreno and Oechel 1991). Higher 
fire intensities are known to deplete seed banks of 
other species (Odion 2000). Presumably, prescribed 
burning of grassland and relatively open California 
poppy habitats, with less above ground biomass 
than chaparral, will result in lower fire intensity and 
heat < 85°C at depths below 2 cm. Our testing at 
85°C may adequately represent conditions of a low 
intensity fire in the upper 2 cm of dry soil where a 
high density of seeds is expected to occur. 

Moist soil has higher thermoconductivity which 
can result in much higher temperatures to greater 
depths (DeBano et al. 1998). This has implications 
for the survival of seed banks of temperature sen- 
sitive species following spring burns over moist 
soil. In addition, Roche et al. (1998) found much 
higher germination and survival from smoke treat- 
ed seedbanks when treated in the dry autumn be- 
fore winter rains than when treated in winter or 
spring. Further studies are needed to determine the 
proportion of seeds killed at different burial depths 
over a range of fire soil temperatures and moisture 
levels. 


Cold treatment and incubation. Many species re- 
quire cold temperatures to break dormancy or for 
optimal germination of conditionally dormant seeds 
(Bewley and Black 1994; Baskin and Baskin 1998), 
and intra-species differences in response to chilling 
have been detected in other geographically variable 
species. For example, in Artemisia tridentata Nutt., 
which also germinates in winter, among-population 
differences were found in germination response to 
different temperatures, with the responses correlat- 
ing to mean January temperature (Meyer et al. 
1990; Meyer and Monsen 1992). In E. californica, 
populations differed in response to chilling, but it 
is too early to tell if response correlates with hab- 
itat. 

We examined chilling under different moisture 
levels (dry, moist, wet) because cold winter tem- 
peratures in California (mid December—February) 
occur partly while the soil is still dry, and partly 
after the onset of winter rains. Furthermore, some 
species have higher seed germination after dry-cold 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 


223 


storage (Padgett et al. 1999). However, for Califor- 
nia poppy, compared to no-cold controls, we found 
a slightly negative effect of wet-cold, no significant 
benefit of dry-cold, and only a weak effect of 
moist-cold treatment. Moist-cold resulted in higher 
germination than no-cold treatment in only six of 
13 trials (Table 6). Two reviews of seed dormancy 
report dormant embryos generally need to be im- 
bibed before they respond to chilling (Nikolaeva 
1969; Baskin and Baskin 1998). It is unlikely that 
a chilling period longer than 4—8 wk would im- 
prove germination. In southern California, poppies 
germinate primarily in January and February, so 
seeds frequently receive a relatively short period of 
cold moist exposure, perhaps 2—8 wk, depending 
on location and onset of rains. In winter 2000, for 
example, native soils were dry into January in Riv- 
erside County, yet there was a spectacular bloom 
that spring. 

The response of seeds to the various cold treat- 
ments was, in part, consistent with the dormancy 
behavior of a winter annual/perennial strategy. 
Many species in lowland Mediterranean climates 
germinate and grow in the cool winter rainy season 
between late fall and late winter. In some of these, 
prolonged warm summer temperatures or shelf stor- 
age can break dormancy while many have the abil- 
ity to germinate at temperatures above about 5°C 
(Baskin and Baskin 1998). Instead of cold temper- 
atures necessarily breaking dormancy, very cold 
temperatures can sometimes send seeds of winter 
annuals back into dormancy (Baskin and Baskin 
1998) or increase degradation of viability (Priestley 
1986). In our first two experiments with the lowest 
temperature cold treatment (2—4°C), dry cold did 
not cause a decrease in germination, but moist- and 
wet-cold treatments appeared to either increase dor- 
mancy of some wild seeds or increase degradation 
of viable seeds to the extent they lost the ability to 
germinate. This cold had no effect on domesticated 
seeds which germinated to nearly 100%. Dry 
smoke treatment applied before cold had an ame- 
liorating effect on the lowering of germination 
(Exp. 2), but smoke treatments applied after cold 
did not. Cold-shock at 2°C actually decreased ger- 
mination slightly or had no effect (Exp. 5), a result 
consistent with a “‘typical’’ winter annual strategy. 
Treatment at 3—3.5°C did not obviously depress or 
enhance germination (Exp. 6). Seeds exposed to 
cold treatments above 4°C began germination in the 
cold chamber and continued to germinate after 
moving to warmer incubation chambers (Exp. 3, 4). 
There was no consistent trend indicating which of 
the cold temperature treatments improved germi- 
nation most. Treatment at 4—9°C (mostly 7°C) did 
not break dormancy in fire controls but did increase 
germination of smoke-treated seeds. Across exper- 
iments, germination appeared to improve under 
cooler incubation temperatures, especially once 
dormancy was broken with smoke. In addition, the 
two lower incubation temperatures (7.5 and 18/6°C 


224 


vs. 22/12°C) resulted in small significant increases 
in germination (Exp. 5). Lower temperatures are 
consistent with seed lab testing of California poppy 
at 15°C (Association of Official Seed Analysts 
1981). The temperature range (2—4°C—22/12°C) un- 
der which perennial, domesticated poppies germi- 
nated to 100% was quite large, suggesting broad 
tolerances in the original source populations or se- 
lection for broad tolerances and loss of conditional 
dormancy under domestication. Overall, these re- 
sults show that the effect of cold is subtle as well 
as population-specific. The results merit further in- 
vestigation into optimal germination temperatures, 
how warm storage temperatures (higher than for 
shelf storage) affect dormancy break, and how ef- 
fects vary among populations of different life-his- 
tories (annual vs. perennial) and from different cli- 
mates. 

Although California poppy has a small linear 
embryo, it does not appear to have the morpho- 
physiological dormancy expected by Martin (1946). 
If there is such dormancy in California poppy, it is 
not general to all populations. Even in wild popu- 
lations with dormant seeds, some seeds appeared to 
lack physiological dormancy. Extraordinarily high 
germination of domesticated seeds from commer- 
cial sources and naturalized garden plants even 
within two weeks of collection without pretreat- 
ment, may be due to lost genetic components 1m- 
portant to germination cycles in natural popula- 
tions. Most of the fresh seeds collected in 2001 
became less dormant as they aged from 2 to 4 and 
~8 months. There was some germination by two 
months suggesting seeds underwent some after-rip- 
ening by 2 months. Increased germination at 4 mo- 
old suggests seeds continued to mature in lab stor- 
age. For all but the Santa Barbara seeds, smoke 
treatment promoted higher seed germination than 
controls even at 2 mo, but was even more effective 
after 4 mo of aging for all populations. Further 
studies are required to reveal if embryos grow dur- 
ing dry storage or if their growth follows hydration 
with or without dormancy breaking smoke treat- 
ment. The ability of smoke to break dormancy in 
seeds less than 4 mo of age suggests that smoke 
from early summer fires may break dormancy pre- 
maturely, possibly making seeds vulnerable to ger- 
minating after summer rain if germination could 
occur at warm temperatures. 

Because species with morphophysiological dor- 
mancy sometimes respond to moist-warm temper- 
ature followed by cold or by GA3 (Nikolaeva 1969; 
Hidayati et al. 2000), we exposed dormant, aged 
seeds of FM96 and NH98 to one week of warm- 
moist stratification at 28°C (A.M.M. and L.F un- 
published). We chose one week because in southern 
California’s hot dry summer environment, soil dries 
out at most within a week of summer storms. Warm 
stratification did not improve germination over con- 
- trols even when followed by GA3 (SO0ppm). GA3 
treatment improved germination much less than liq- 


MADRONO 


[Vol. 49 


uid smoke. We plan to test additional combinations 
of warm stratification followed by cold incubation 
treatments. 


Synergistic effects of smoke and cold. We did 
find a weak synergistic effect of cold treatment in 
combination with smoke treatments for six popu- 
lations in an overview of experiments in which 
smoke treatment was applied before cold treatment 
(Table 6). Under no-cold, there was consistently 
higher germination for liquid smoke than no-smoke 
controls in 15 of 15 trials; under moist-cold, liquid 
smoke resulted in even higher germination in seven 
of nine trials (Table 6, Exp. 2—5). Similarly, under 
no-cold, dry smoke treatment resulted in higher 
germination than no-smoke controls in 10 of 13 tri- 
als; under moist-cold, dry-smoke resulted in even 
higher germination in 10 of 13 trials. This syner- 
gism suggests smoke is triggering a growth se- 
quence that must occur before cold temperatures 
can succeed in promoting germination and may, in 
part, be substituting for other environmental cues 
that would normally occur before cold treatment 
(e.g., warm summer temperatures or exogenous 
chemicals in the soil). 


Seed age. Interestingly, young seeds had lower 
seed dormancy than aged seeds from the same pop- 
ulations, and dormancy was easier to break than in 
aged seeds. In young seeds, dry smoke increased 
germination over the controls, and liquid smoke 
produced the highest germination of any treatment 
(62—95% in young seeds vs. O—70% in aged seeds). 
Viability dropped no more than 6% between young 
and aged collections from the same sites (viability 
confirmed with TZ and checked with fluoroscien 
diacetate methods, unpubl. after Windholm 1972), 
indicating that older seeds entered a secondary dor- 
mancy during prolonged shelf storage, or that seeds 
experienced deterioration in germinability that can- 
not be detected with chemical viability tests. Ad- 
ditional cues may be required to break any second- 
ary dormancy. The ability to enter secondary dor- 
mancy is important to seasonal cycling of dorman- 
cy, the building of a seed bank, and presence of a 
bet-hedging strategy in unpredictable environ- 
ments. This may be especially important to seeds 
that germinate in dark so that the seed bank is not 
exhausted in any particular year. 


Variation within and among populations. Even 
though smoke alone or with cold succeeded in in- 
creasing germination across populations, popula- 
tions differed substantially in response to those 
treatments and in baseline dormancy of untreated, 
shelf-stored seeds. Populations also differed in re- 
sponse to wet or dry smoke treatments, often in 
association with annuality or perenniality (Exp. 4). 
Furthermore, in all wild populations and treatments 
a fraction of the viable seeds did not germinate. 
This is important for several reasons: 1) we have 
not identified a general natural cue that promotes 
germination of young seeds enmasse in the absence 


2002] 


of fire, or of aged seeds enmasse with or without 
fire; 2) there are differences among seeds within 
populations for dormancy and dormancy breaking 
requirements; and 3) if the variation in response is 
heritable it can be selected and thus the observed 
variation in response among populations may be 
the result of adaptation to local conditions and may 
be linked to different life-history strategies (Cook 
1962; Young and Augspurger 1991). Variation 
within populations, including changes with seed 
age, may provide a good bet-hedging strategy. 

Clearly, the domesticated (Dom and RIV) pop- 
ulations of California poppy with their non-dormant 
seeds were very different from all wild populations 
in our study. Interestingly, some perennial popula- 
tions from coastal northern California also have no 
seed dormancy (Cook 1962; Montalvo personal ob- 
servation, e.g., RMO1). Large differences among 
wild-collected populations, and between wild and 
the domesticated populations, show the importance 
of noting seed source and population traits when 
doing research or restoration with this species. Our 
results indicate that results of seed dormancy ex- 
periments from one or two populations cannot be 
generalized correctly to the species level. Research 
on a range of wild populations which have different 
levels of seed dormancy, including perennial pop- 
ulations with non-dormant seeds needs to be done 
before any generalizations can be made about light 
inhibition of germination and requirements for ger- 
mination in this species. 


In pursuit of unknown cues. We have not discov- 
ered how to break dormancy without smoking 
seeds. Given that many California poppy seeds ger- 
minate in nature in the absence of fire, future work 
should explore the combined effects of seed burial 
and seasonal changes in temperature on seed ger- 
mination, with and without smoke. In several Aus- 
tralian species, prolonged seed burial was found to 
affect the seed coat by increasing permeability to 
water and seed coat breakage in ways that facili- 
tated germination (Tieu and Egerton-Warbuton 
2000). In addition, Roche et al. (1997a) found a 
synergistic effect of lengthy seed burial followed 
by smoke addition on germination of 60% of over 
100 Australian species tested. Smoke treatment by 
itself approximately doubled germination of seeds, 
but smoke treatment combined with soil storage 
quadrupled germination. Seed burial was also 
found to be important to germination of Dendro- 
mecon rigida Benth., a fire following species in the 
Papaveraceae (Keeley and Fotheringham 1998b) 
that is closely related to the genus Eschscholzia. It 
is possible that some dormant poppy seeds require 
a combination of seed burial and changing temper- 
atures before they can successfully germinate. Al- 
though exposure to warm summer temperatures 
breaks dormancy in many winter annuals (Baskin 
and Baskin 1998), in a recent seed burial study 
where the soil was dry, as is natural in this region, 


MONTALVO ET AL.: SEED GERMINATION IN CALIFORNIA POPPY 225 


we found that exposure to natural summer and early 
fall temperatures (July—mid November) did not no- 
tably improve germination relative to shelf-stored 
controls (A. Montalvo and C. Koehler unpublished 
data). 


Implications for conservation and_ restoration. 
Given the beneficial effect of smoke treatments on 
California poppy germination, prescribed fire might 
prove useful for poppy management as long as heat 
does not penetrate into the seed bank too deeply. 
Liquid smoke treatment of the seed bank may sim- 
ulate the beneficial effects of fire when prescribed 
burning is not feasible or when heat might unduly 
affect viability of poppy or other native species. On 
a cautionary note, evaluation of the differential ef- 
fects of smoke on other species, both native and 
exotic, is warranted before using smoke and/or fire 
aS a management tool. Seeds of exotic and native 
species may differ in response to smoke treatment 
and changes in seed coat chemistry may be irre- 
versible (Roche et al. 1998). If smoke facilitates a 
non-reversible breaking of seed dormancy for a 
large fraction of the seed bank, then effects of fre- 
quent or even occasional burning could last more 
than one season and could disrupt the selective val- 
ue of dormancy. The seedbank could be depleted if 
seeds germinate under conditions detrimental to 
successful establishment. It is important to weigh 
the consequences of such negative direct effects of 
repeated burning on the seed bank, against the ben- 
eficial effects of killing competitors, or perceived 
benefit of managing for large flowering displays. 

Our results can help to develop regionally adapt- 
ed populations of California poppy for restoration 
and revegetation projects in southern California, 
which have in the past met with mixed success. In 
xeric, non-irrigated locations where native poppies 
tend to have largely dormant seeds, plantings of 
domesticated seeds can die out within a few years 
(A. Montalvo personal observation). Most com- 
mercially available poppy seeds are non-dormant, 
perennial, and have resulted from many generations 
of seed increase, a practice that selects against seed 
dormancy. The seed industry has avoided dealing 
in native, dormant seeded populations, in part be- 
cause it is difficult to break their seed dormancy 
(Victor Schaff, S&S Seeds personal communica- 
tion). In the future, the seed industry could smoke- 
treat dormant native poppy seeds before planting to 
avoid selecting against seed dormancy. Given the 
evidence for locally adaptive differences and home 
site advantage in many other plant species (for re- 
view see Langlet 1971; Montalvo and Ellstrand 
2000), and the sometimes adverse effects of hy- 
bridization among genetically differentiated popu- 
lations (Millar and Libby 1989; Knapp and Rice 
1994: Montalvo et al. 1997, Montalvo and Ellstrand 
2001), the use of local poppy seeds for restoration 
and reseeding would likely increase the long-term 
success of planted populations. 


226 


ACKNOWLEDGMENTS 


We thank John Crossman, Gordon Fox, Robert Noll of 
Noll seeds and Victor Schaff of S&S Seeds for generous 
contributions of native seed collections, two anonymous 
reviewers for suggestions to improve the manuscript, Hi- 
lary Wall for unpublished information about effects of liq- 
uid smoke on other local native plants, L. Egerton-War- 
burton for discussions and information about seed ger- 
mination biology and seed viability testing, undergraduate 
students M. Sripracha, J. Terrell, and M. Blatt who helped 
with seed processing and scoring, the Antelope Valley 
California Poppy Reserve and Western Riverside Multi- 
Species Reserve for permission to collect seeds from re- 
serve populations, Regen of Glasgow Kentucky and Gray- 
son Australia of Bayswater, Victoria for information and 
samples of Regen 2000 liquid smoke, and California De- 
partment of Parks and Recreation (Agreement No. 67834) 
and Metropolitan Water District of Southern California 
(Agreement No. 4602) for financial support. 


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MADRONO, Vol. 49, No. 4, pp. 228-236, 2002 


TEMPERATURE LIMITATIONS FOR CULTIVATION OF EDIBLE 
CACTI IN CALIFORNIA 


PARK S. NOBEL!, ERICK DE LA BARRERA, DAVID W. BEILMAN, 
JENNIFER H. DOHERTY, AND BRIAN R. ZUTTA 
Department of Organismic Biology, Ecology, and Evolution, University of 
California, Los Angeles, CA 90095-1606 


ABSTRACT 


Hylocereus undatus (a hemiepiphyte) and Opuntia ficus-indica (“prickly pear’’) are cultivated world- 
wide as specialty fruit and vegetable crops, so the role of temperature in determining regions suitable for 
commercial growth of these cacti was investigated for California, the leading agricultural state in the 
United States. Air temperatures below —2.5°C and above 45°C are lethal for H. undatus compared with 
below —10°C and above 65°C for O. ficus-indica, demonstrating the latter’s greater tolerance of extreme 
temperatures. Mean nighttime air temperatures influence net CO, uptake for these Crassulacean acid 
metabolism species, optimal uptake occurring at 20°C for H. undatus and at 14°C for O. ficus-indica. 
Extreme air temperatures over a 30-year period for 326 weather stations and mean nighttime temperatures 
for 259 stations were mapped to identify where these species could be cultivated. Only 2% of the state’s 
total area avoided temperatures lethal to H. undatus, mostly along the southern coast. In contrast, 36% 
of the state’s area was possible for O. ficus-indica, exclusion occurring in mountainous regions. A Tem- 
perature Index (net CO, uptake over 24-hour periods at a particular temperature divided by uptake at the 
optimal temperature) was also utilized to evaluate a region’s suitability for growing these cacti. The 
Temperature Index was below 0.7 for 59% of the weather stations for H. undatus but for only 16% for 
O. ficus-indica. In the regions where lethal extreme temperatures did not occur, the Temperature Index 
averaged more than 0.8 for both species. Use of a Temperature Index based on net CO, uptake together 
with extreme temperature events can help evaluate regions for cultivating cacti with edible fruits or other 
new crops. 


Key Words: CO, uptake, Crassulacean acid metabolism, fruit, Hylocereus undatus, Opuntia ficus-indica. 


As a result of its geology, topography, and cli- 
mate, California is the most biodiverse state in the 
United States, having nearly 6000 native species of 
vascular plants (Hickman 1993). Such variety also 
permits the production of about 350 agricultural 
commodities that generate $30 billion in revenue 
per year, more than for any other state, such com- 
modities being responsible for 10% of the jobs in 
California (California Department of Food and Ag- 
riculture 2001). Much of the agricultural economy 
derives from intensively managed specialty crops— 
e.g., minor crops such as grapes, tomatoes, straw- 
berries, lettuce, and flowers generate about 20% of 
the agricultural sector’s annual revenue (California 
Department of Food and Agriculture 2001). In this 
regard, considerable interest exists among both 
government officials and also producers to develop 
new specialty crops, such as the cacti with edible 
fruits considered here. 

Given their potentially high productivity and tol- 
erance of high temperatures (Nobel 1988), cacti 
have become important crops in arid and semiarid 
regions worldwide (Barbera 1995). The prickly 
pear cactus Opuntia ficus-indica (L.) Miller is cul- 
tivated on over one million hectares in more than 
30 countries primarily for fodder but also as a fruit 


' Author for correspondence, e-mail: psnobel@biology. 
ucla.edu 


crop (“‘cactus pears’’) and on a limited scale as a 
vegetable crop (“‘nopalitos’’; Nobel 2000). Second 
in importance among edible cacti are hemiepiphy- 
tes in the genera Hylocereus and Selenicereus, 
whose fruits are commonly referred to as “‘pitahay- 
as,’ which are cultivated in 20 countries, with par- 
ticularly noteworthy success in Southeast Asia 
(Nerd et al. 2002; Nobel and De la Barrera 2002). 
Cacti have been cultivated in California since the 
eighteenth century, when O. ficus-indica was plant- 
ed around the Spanish missions along the coast for 
its fruit and its mucilage, which was utilized as a 
binding material for adobe bricks (McLeod 1975). 
In the early 1900s, Luther Burbank, who pioneered 
several specialty crops, developed a “‘spineless”’ 
variety of O. ficus-indica (Savio 1989). The 
D’ Arrigo Brothers have plantations of spineless O. 
ficus-indica in the Santa Clara Valley (also known 
as the Salinas Valley) near Gilroy that were estab- 
lished in the 1920s for fruit (Curtis 1977), and var- 
ious companies in southern California are currently 
exploring the pitahaya market (Savio 1989; Valdi- 
via 2000; P.S. Nobel personal observation). 

Most gas exchange for cacti occurs at night when 
air temperatures are lower than during the daytime, 
a characteristic of the Crassulacean acid metabo- 
lism (CAM) photosynthetic pathway. This physio- 
logical strategy, which has evolved for species in 
over 30 vascular plant families, reduces transpira- 


2002] 


tional water loss (Nobel 1999; Taiz and Zeiger 
2002). Optimal nocturnal air temperatures for net 
CO, uptake by CAM plants are generally from 10 
to 20°C (Nobel 1988). Extreme temperatures limit 
CO, uptake and also can damage cacti, with epi- 
sodic freezing temperatures generally being more 
limiting to plant distribution than high tempera- 
tures. For instance, based on the uptake of a vital 
stain into the central vacuoles, the photosynthetic 
cells (chlorenchyma) of O. ficus-indica have 50% 
mortality (LT;,) at the extremely high temperature 
of 62.4°C for plants kept at day/night air tempera- 
tures of 40/30°C and at 66.6°C for plants kept at 
50/40°C, indicating that O. ficus-indica has a high- 
temperature acclimation (hardening) of 4.2°C per 
10°C increase in air temperature (Nobel 1988). The 
LT;, for low-temperature tolerance of O. ficus- 
indica is —7.7°C for plants kept at day/night air 
temperatures of 20/10°C, decreasing to —8.8°C for 
plants kept at 10/0°C, indicating a low-temperature 
acclimation of 1.1°C for a temperature decrease of 
10°C (Nobel 1988). Hylocereus undatus (Haworth) 
Britton & Rose, which is native to neotropical for- 
ests (Britton and Rose 1963; Backeberg 1966; 
Barthlott and Hunt 1993) where temperatures tend 
to be warm and vary little over the course of a year 
(Croat 1978; Liittge 1997), shows a high-tempera- 
ture acclimation of only 1.4°C per 10°C increment 
in air temperature, as LT;, occurs at 54.0°C for 
plants kept at day/night air temperatures of 25/15°C 
and at 55.4°C for plants at 35/25°C; furthermore, 
plants kept at 40/30°C develop stem tissue necrosis, 
leading to death after 19 weeks (Nobel and De la 
Barrera 2002). Because the responses of H. undatus 
to freezing temperatures have not been reported, 
one of the objectives of the present research was to 
determine its low-temperature tolerance and its 
low-temperature acclimation. 

An Environmental Productivity Index (EPI), 
which can help to evaluate the feasibility for ex- 
panding the area of cultivation of crops such as 
cacti, indicates the primary influence of water, tem- 
perature, and light on net CO, uptake and hence 
biomass productivity of plants (Nobel 1988, 1999). 
EPI is defined as the Water Index X the Tempera- 
ture Index X the PPF Index (PPF refers to the pho- 
tosynthetic photon flux, composed of wavelengths 
of light from 400 to 700 nm), where each compo- 
nent index ranges from 0.00, when that environ- 
mental factor eliminates net CO, uptake, to 1.00, 
when that factor is optimal for net CO, uptake (No- 
bel 1988). EPI ignores secondary interactions, such 
as the different response to PPF when temperature 
is limiting, but in any case net CO, uptake is gen- 
erally low under such conditions. The individual 
indices are determined in the laboratory under con- 
trolled conditions over 24-hour periods by varying 
the environmental parameter to be studied, while 
keeping the other factors constant at optimal values. 
EPI can then be calculated under field environmen- 
tal conditions, as has been done to predict quanti- 


NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 


229 


— Hylocereus undatus 
—— Opuntia ficus-indica 


Daily net CO, uptake per unit stem area 
(fraction of maximum) 


eee L Spee l 
5 10 15 20 25 30 35 


Mean nighttime air temperature (°C) 


Fic. 1. Responses of total daily net CO, uptake by Hy- 
locereus undatus and Opuntia ficus-indica to mean night- 
time air temperatures. Plants were maintained for 7 to 14 
days at a particular temperature before measurement. Data 
for H. undatus are from Raveh et al. (1995) and Nobel 
and De la Barrera (2002) and for O. ficus-indica are from 
Nobel (1988) and Nobel and Bobich (2002). 


tatively the growth of Agave deserti along an ele- 
vational gradient (Nobel 1984) and to establish new 
plantations of O. ficus-indica in Chile (P. S. Nobel 
personal observation). For agricultural purposes, 
the Water Index can be increased to 1.0 by irriga- 
tion, and the PPF Index can be manipulated by 
varying the spacing between plants. However, it is 
difficult to control temperatures in the field. Deter- 
mining the Temperature Index may therefore help 
producers decide the suitability of a particular re- 
gion for growing a certain crop. In this regard, net 
CO, uptake is optimal at a mean nighttime temper- 
ature of 20°C for H. undatus and 14°C for O. ficus- 
indica (Fig. 1). Also, H. undatus has a narrower 
temperature range than does O. ficus-indica within 
which the Temperature Index is above 0.5 (9 to 
26°C versus 2 to 26°C) and above 0.8 (14 to 23°C 
versus 6 to 20°C). The present research uses the 
previously measured Temperature Indices for H. 
undatus and O. ficus-indica (Fig. 1), the low and 
the high temperature tolerances for both species, 
and climatic data to evaluate potential regions for 
their cultivation in California. This approach used 
for cacti here can also serve as a model for assess- 
ing the cultivation potential of other plant species. 


MATERIALS AND METHODS 


Temperature tolerances. The low-temperature 
tolerance has already been determined for Opuntia 


ficus-indica, as have the high-temperature toler- 


ance for it and Hylocereus undatus (Nobel 1982, 
1988; Nobel and De la Barrera 2002), using the 
vacuolar uptake of neutral red (3-amino-7-dimeth- 
ylamino-2-methylphenazine hydrochloride) fol- 
lowing exposure for 60 min to a particular extreme 
temperature (Onwueme 1979; Didden-Zopfy and 
Nobel 1982; Nobel et al. 1995). Hence, the low- 
temperature tolerance of H. undatus was similarly 


230 


determined here using the neutral red assay. For 
this purpose, five plants approximately 45 cm in 
shoot length were obtained from the Cactus Trad- 
ing Company (Jamul, CA). They were grown in 
each of two Conviron E-15 environmental cham- 
bers (Controlled Environments, Pembina, ND) 
with day/night air temperatures of 20/10°C or 30/ 
20°C for 4 weeks with weekly application of 0.2- 
strength Hoagland’s solution supplemented with 
micronutrients and a total daily PPF of 16 mol m 
* day-!, environmental conditions that are near the 
optimal for H. undatus (Raveh et al. 1995; Nobel 
and De la Barrera 2002). 

Low temperatures at 1 to 2°C intervals decreas- 
ing from 4°C were obtained in an ULT-80 ultra- 
low-temperature freezer (Rheem Manufacturing, 
West Columbia, SC). Approximately 1.5 g of stems 
were removed with a scalpel, placed in contact with 
a copper-constantan thermocouple 0.51 mm in di- 
ameter, and wrapped in aluminum foil to prevent 
desiccation; the samples were then cooled at 5°C 
hr~', similar to stem cooling rates observed in the 
field (Nobel 1988; Nobel et al. 1995). After expo- 
sure to a particular temperature for 60 min, the 
samples were sliced into sections approximately 
700 wm thick using razor blades and then placed 
in 0.2% (w/w) neutral red for 90 min for stain up- 
take, which occurs for the vacuoles of living cells 
only and indicates membrane integrity (Onwueme 
1979; Nobel et al. 1995). The tissue samples were 
then placed for 10 min in 0.25 M potassium phos- 
phate buffer (pH 7.8) at 25°C followed by 24 hours 
at 6°C in distilled water to help remove excess stain 
and hence to sharpen the images, after which ap- 
proximately 130 intact cells per sample were ex- 
amined at 100X using a BH-2 phase-contrast mi- 
croscope (Olympus, Lake Success, NY) to check 
for stained (living) versus unstained cells. The low 
temperature treatment that halved stain uptake from 
the maximum occurring at 4°C (LT.,), a reliable test 
for predicting eventual tissue necrosis (Didden- 
Zopfy and Nobel 1982; Smith et al. 1984; Nobel et 
al. 1995), was determined graphically under each 
condition. Low-temperature acclimation was ana- 
lyzed by comparing LT;, for plants at day/night air 
temperatures of 20/10°C versus 30/20°C using an 
unpaired Student t-test. 


Extreme temperature limitation. For cacti in na- 
ture as well as under cultivation, infrequent freez- 
ing episodes can be severely limiting (Steenbergh 
and Lowe 1976; Russell and Felker 1987; Nobel 
1988). Moreover, fruit production by H. undatus 
and O. ficus-indica can occur two years after the 
establishment of a plantation, but approximately 10 
years may be necessary to obtain optimal yields 
and an even longer period for appropriate return on 
the initial investment (Mizrahi and Nerd 1999; In- 
glese et al. 2000; Nerd et al. 2002). Also, the fruit- 
ing potential of O. ficus-indica tends to decrease 
after 25 to 30 years (Inglese et al. 2002). Thus, 


MADRONO 


[Vol. 49 


instead of using annual minimum temperatures, the 
lowest air temperatures recorded between 1961 and 
1990 (the most recent period summarized with cli- 
mate normals) at 326 California weather stations 
were obtained from the Climate Atlas of the Con- 
tiguous United States (National Climatic Data Cen- 
ter 1995). Similarly, the highest air temperatures 
were obtained for the 318 weather stations with 
suitable records for the same period. The tempera- 
tures were converted from Farenheit to Celsius and 
then ranked in 2.5°C intervals for low-temperature 
extremes or 5°C intervals for high-temperature ex- 
tremes. In addition, weather station data were in- 
terpolated, correcting for elevation using a lapse 
rate of 6°C per km (Nobel 1999), to identify the 
areas with record minimum temperatures below 
—10°C or above —2.5°C as well as record maxi- 
mum temperatures above 45°C over the 30-year pe- 
riod (1961-1990). The resulting low-temperature 
and high-temperature maps were created in Arc- 
View 3.1 (ESRI, Redlands, CA). 


Temperature Index. Daily minimum tempera- 
tures averaged over each month for the California 
weather stations from 1961 to 1990 and then ay- 
eraged over the 30 years were also obtained from 
the Climate Atlas of the Contiguous United States 
(National Climatic Data Center 1995). Nighttime 
mean air temperatures were estimated by adding 
3°C to the average minimum temperature (Nobel 
1988) recorded at each of the 259 weather stations 
with sufficient records. The Temperature Index for 
total daily net CO, uptake per unit stem area for H. 
undatus and O. ficus-indica was then determined 
for each month using the known temperature re- 
sponses for these two species (Fig. 1). The twelve 
monthly values were averaged to obtain an annual 
Temperature Index for each weather station and 
maps were created in ArcView 3.1. 


RESULTS 


Low-temperature tolerance for Hylocereus un- 
datus. Neutral red accumulation in chlorenchyma 
cells of Hylocereus undatus decreased as the treat- 
ment temperature was lowered below O0°C (Fig. 2). 
For H. undatus growing at day/night air tempera- 
tures of 30/20°C, the percentage of cells taking up 
the vital stain was halved (LT;,) at —1.31 + 0.04°C. 
The LT... for H. undatus acclimated to day/night air 
temperatures of 20/10°C was —1.55 + 0.07°C. This 
species thus displayed a small, yet significant, low- 
temperature acclimation (hardening) of 0.24 = 
0.08°C per 10°C decrease in temperature (t = 2.98, 
P < 0.01, df = 8). 


Extreme temperature limitations. During the 30- 
year period considered (1961-1990), 40% of the 
326 weather stations had temperatures below 
—10°C, corresponding to 64% of the area of Cali- 
fornia, mainly in regions at high elevations in the 
Sierra Nevada, Coast and Diablo ranges, and the 
San Bernardino Mountains, as well as in northern 


2002] 


oy 

oO 

oO 
T 


ies) 
oOo 
T 


=O SANE 7 
—A— 20/10°C 


(% of maximum) 


Chlorenchyma cells taking up stain 


3 2 -1 0 1 2 3 4 
Temperature (°C) 


Fic. 2. Influence of day/night air temperatures of 20/ 
10°C and 30/20°C on the low-temperature tolerance of H. 
undatus. Uptake of neutral red was determined for pieces 
of chlorenchyma incubated at a particular treatment tem- 
perature for 60 min. Data are means = SE (n = 5 plants). 


>0 
0 to-2.5 
-2.5 to -5 
-5 to-7.5 
-7.5 to -10 
< -10 


*ODQA0 70 ® 


Fic. 3. 


NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 254 


California (Fig. 3A). Regions with extreme mini- 
mum temperatures from —5 to —10°C (49% of the 
Stations) are concentrated along the coast, through 
the Central Valley, and in southern California. Re- 
gions that were never below —5°C (11% of the sta- 
tions) are largely restricted to thin coastal regions 
in the San Francisco Bay area, the Channel Islands, 
and in southern California from Ventura to San Di- 
ego counties, in addition to lower inland elevations 
in southern California. Regions with extreme min- 
imum temperatures above —2.5°C (7% of the sta- 
tions), representing only 2% of the state’s area, are 
located along the coast in Ventura, Los Angeles, 
Orange, and San Diego counties plus one station in 
the San Francisco Bay area (Fig. 3B). Only one 
station (at the University of California, Los Ange- 
les) remained above 0°C during the period consid- 
ered (1961-1990). 

During the same 30-year period, five weather sta- 
tions had maximum temperatures above 50°C 


20 to 25 
25 to 30 
30 to 35 
35 to 40 
40 to 45 
45 to 50 
> 50 


3 ee et ar ef (| 


Temperature extremes at California weather stations from 1961—1990: (A) record minimum temperatures and 


(B) record maximum temperatures. Letters indicate specific ranges of extreme temperatures, e.g., c corresponds to a 


record minimum temperature between —2.5°C and —5°C. 


232. 


0.0 to 0.1 
0.1 to 0.2 
0.2 to 0.3 
0.3 to 0.4 
0.4 to 0.5 
0.5 to 0.6 
0.6 to 0.7 
0.7 to 0.8 
0.8 to 0.9 
0.9 to 1.0 


0 
1 
2 
3 
4 
5 
6 
la 
8 
9 


Fic. 4. 


MADRONO 


[Vol. 49 


Annual Temperature Index values at California weather stations averaged over 30 years (1961—1990) for (A) 


H. undatus and (B) O. ficus-indica. Each number represents the influence of temperature on the fraction of maximal 
total daily net CO, uptake averaged over the year, as calculated from monthly mean nighttime air temperatures, for 
each weather station. Numbers indicate specific ranges of the annual Temperature Index, e.g., 7 corresponds to 0.7 to 


0.8. 


(1.6% of the 318 stations), with the hottest site 
(53°C) in Death Valley (Fig. 3B). Maximum tem- 
peratures from 45 to 50°C occurred at 26% of the 
weather stations, concentrated at lower elevations 
in the Mojave Desert and Death Valley, along the 
Central Valley, and in inland southern California. 
Approximately 40% of the state’s area had temper- 
atures above 45°C during the period considered. 
Regions with maximum temperatures from 40 to 
45°C for the 30-year period considered (53% of the 
stations) are situated at higher elevations in north- 
ern California, in the Central Valley, and along the 
coast from the San Francisco Bay area through San 
Diego County. Only 20% of the stations, distributed 
in coastal regions or at high mountain elevations, 
recorded maximum temperatures below 40°C (Fig. 
3B). 


Temperature Index. For H. undatus, the annual 
Temperature Index (TI) was below 0.5 for 23% of 
the 259 weather stations, mostly those at high ele- 
vations, especially in northern and inland California 
(Fig. 4A). An annual TI of 0.5 to 0.7 occurred for 
36% of the stations, most occurring in the Central 
and Imperial valleys. An annual TI from 0.7 to 0.8 
was restricted to Sacramento, the San Francisco 
Bay area, and the southern California coast (33% 
of the stations; Fig. 4A). A TI of 0.8 to 0.9 was 
found only in Los Angeles, Orange, and San Diego 
counties as well as in Death Valley (8% of the sta- 
tions). 

Compared with H. undatus, the annual TI tended 
to be higher for Opuntia ficus-indica, only 4% of 
the weather stations having annual values below 0.5 
(Fig. 4B). An annual TI of 0.5 to 0.7 occurred for 


2002] 


12% of the stations, mostly in inland regions, es- 
pecially for northeastern California. Moderately 
high TI, from 0.7 to 0.8 (18% of the stations) and 
from 0.8 to 0.9 (32% of the stations), were located 
throughout California, except at high elevations in 
the Sierra Nevada. Annual TI values above 0.9 for 
O. ficus-indica occurred along the coast from the 
Oregon border south to Ventura County as well as 
in various inland regions, such as in the San Fran- 
cisco Bay area and the Los Angeles basin (34% of 
the stations; Fig. 4B). 


DISCUSSION 


Twenty-three species of cacti have been exam- 
ined for tolerance to extreme temperatures (Nobel 
1982, 1988; Smith et al. 1984), which are important 
in determining natural distributions and potential 
regions for cultivation. The least freezing tolerant 
of these species, Opuntia ramosissima, is native to 
the deserts of the southwestern United States and 
northern Mexico and has an LT;, (temperature that 
kills half of the cells compared with the control) of 
—4.4°C when maintained at day/night air tempera- 
tures of 10/0°C (Nobel 1982). Hylocereus undatus 
was even less freezing tolerant, with an LT,, of 
—1.6°C when maintained at 20/10°C. Taking into 
consideration its relatively small low-temperature 
acclimation of 0.2°C per 10°C decrease in air tem- 
perature observed here, the LT;, for H. undatus is 
only —1.8°C at 10/0°C, indicating that it is extreme- 
ly sensitive to freezing temperatures. In addition, 
H. undatus is not as tolerant of high temperatures 
as are the other cactus species examined (Nobel 
1988) and also exhibits little high-temperature ac- 
climation (Nobel and De la Barrera 2002). Indeed, 
acclimation is the key to tolerating extreme tem- 
peratures, and only one cactus species (Ferocactus 
covillei) has less low-temperature acclimation and 
none has less high-temperature acclimation than 
does H. undatus (Nobel 1988). In this regard, H. 
undatus is native to neotropical forests with mod- 
erate and rather stable temperatures (Britton and 
Rose 1963; Backeberg 1966; Croat 1978; Barthlott 
and Hunt 1993; Liittge 1997) and apparently is not 
genetically or physiologically capable of apprecia- 
ble acclimation to low or to high temperatures, al- 
though further studies are necessary to understand 
its intraspecific variation. 

LT.) is used for its ease of measurement and be- 
cause it is often the temperature where stem dam- 
age becomes visible, although cacti generally do 
not die until the cellular uptake of a vital stain is 
reduced to zero. As assessed by neutral red stain- 
ing, stem death of most cacti occurs approximately 
4°C below the low-temperature LT,, and 4°C above 
the high-temperature LT., (Nobel et al. 1986; Nobel 
1988). Moreover, LT., refers to tissue temperatures 
when damage occurs, not air temperatures, which 
can differ significantly. On clear nights, tempera- 
tures of cactus stems can be a few degrees Celsius 


NOBEL ET AL.: TEMPERATURE LIMITATIONS AND CACTI 


i) 
Oo 
oS) 


below air temperature due to transpirational cooling 
and especially net heat loss by infrared (longwave) 
radiation (Nobel 1988, 1999). Indeed, radiation 
frosts, when the tissue achieves freezing tempera- 
tures with air temperatures above O°C, are a severe 
agricultural problem in California, especially for 
the citrus industry (Pehrson 1984), and affect the 
suitability of a site for cactus cultivation. Freezing 
temperatures can cause extracellular ice crystal for- 
mation in cacti, which draws water out of the cells 
and can lead to irreversible damage (Burke et al. 
1976; Nobel 1982, 1988). For regions that experi- 
ence infrequent damaging or even lethal low tem- 
peratures, freeze-protection methods, such as shade 
cloth, heaters, and overhead irrigation, can mitigate 
freezing damage to perennial plants such as H. un- 
datus and O. ficus-indica (Pehrson 1984; Perry 
1998). 

Differences between air and tissue temperatures 
can be even larger during the day than at night, 
depending on stem orientation relative to solar ir- 
radiation and stem massiveness (Nobel 1988). Tis- 
sue temperatures of the relatively thin stems of H. 
undatus are not expected to rise more than | to 2°C 
above air temperature, especially in its typically 
shaded habitat (Nobel and De la Barrera 2002). In 
contrast, stem temperatures for O. ficus-indica can 
be more than 15°C above air temperatures (Wallace 
and Clum 1938; Konis 1950). Extremely high tem- 
peratures can denature proteins, degrade cell mem- 
branes, and disrupt metabolism in general (Nobel 
1988; Srinivasan et al. 1996; Taiz and Zeiger 2002). 
For H. undatus, daytime temperatures of 45°C can 
reduce flower and hence fruit production (Mizrahi 
and Nerd 1999). Shade cloth has been used to ame- 
liorate the effects of high temperatures (and high 
PPP) on A. undatus growing in Israel (Raveh et al. 
1998). In contrast, stems of O. ficus-indica usually 
are not damaged until air temperatures exceed 65°C 
and it can even tolerate 60 min at 70°C (Nobel et 
al. 1986; Nobel 1988), so high temperatures should 
not be a limiting factor for cultivation of this cactus 
in California. 

Hylocereus undatus can be grown in regions 
with extreme temperatures above —2.5°C and be- 
low 45°C (Fig. 5A), which occur for only 2% of 
the state’s area. On the other hand, O. ficus-indica 
is excluded only from regions of California where 
the minimum temperature is below —10°C (Fig. 
5B) and can be grown in 36% of the state’s area. 
The climate of California, which renders most of 
the state too cold for maximal net CO, uptake by 
H. undatus, resulted in a lower annual Temperature 
Index (TI) for it, averaging 0.57 throughout the 
state compared to 0.82 for O. ficus-indica. The low- 
er average annual TI for H. undatus reflects both 
its relatively high optimal mean nighttime temper- 
ature for net CO, uptake and also the more rapid 
decrease in net CO, uptake above and below the 
optimal value than is the case for O. ficus-indica. 
In the regions where the cacti can be cultivated be- 


234 


MADRONO 


[Vol. 49 


Fic. 5. 


Annual Temperature Index values (from Fig. 4) for the weather stations with extreme temperatures within the 


tolerable ranges for (A) H. undatus (extreme temperatures between —2.5°C and 45°C) and (B) O. ficus-indica (minimum 


temperature of —10°C and no maximum temperature). 


cause of lack of lethal extreme temperatures (Fig. 
5), the annual TI averages 0.83 for H. undatus and 
0.90 for O. ficus-indica, both high values, indicat- 
ing that the nighttime temperatures for regions 
within the extreme temperature limits are condu- 
cive to substantial net CO, uptake by these two spe- 
cies. The similarly high annual TI estimated for O. 
ficus-indica in areas suitable for cultivation (and in 
the entire state) reflect the wide range of nighttime 
temperatures at which this species can be grown 
successfully. Nevertheless, the exclusion of 64% of 
California’s area indicates that even single extreme 
low-temperature events can greatly damage plan- 
tations of O. ficus-indica (Russell and Felker 1987; 
Nobel 1988). Frost damage can be avoided with 
appropriate agricultural practices (Pehrson 1984; 
Perry 1998) or by the utilization of cold-tolerant 
cultivars of Opuntia, a genus with considerable ge- 
netic diversity as well as a long history of agricul- 


tural selection (Russell and Felker 1987; Parish and 
Felker 1997; Casas and Barbera 2002). 

The Temperature Index is the least manageable 
of the three components of the Environmental Pro- 
ductivity Index (EPI) in an agricultural setting, 
which is the reason for its consideration in the pre- 
sent study. The relationship between EPI, a predic- 
tor of net CO, uptake, and fruit production has not 
been investigated for H. undatus or O. ficus-indica, 
but EPI closely predicts leaf production for Agave 
tequilana (Nobel and Valenzuela 1987) and cladode 
production for O. ficus-indica (Nobel 1988) under 
cultivation. Besides the present focus on fruit crops 
and the use of young cladodes of O. ficus-indica as 
a vegetable, most cultivation of cacti worldwide is 
dedicated to fodder production (Nobel 2000), due 
to ease of management without irrigation or fertil- 
izer application and an acceptable protein content 
of 5 to 8% on a dry mass basis (Nobel 1988; Pi- 


2002] 


mienta Barrios 1990). Such fodder could be used 
in California as an input to the state’s livestock and 
poultry sector, which is responsible for 10% of the 
state’s agricultural revenue (California Department 
of Food and Agriculture 2001). In any case, the 
market for cactus fruits, which had been restricted 
for cactus pears in Mexico and southern Italy as 
well as for pitahayas in southeastern Asia, has re- 
cently expanded globally. The establishment of new 
plantations of these and other edible cacti in Cali- 
fornia, for domestic consumption by ethnic groups 
who traditionally consume cacti plus others who are 
developing a taste for these exotic fruits plus ex- 
portation, could contribute to the diversification of 
revenue production for this leading agricultural 
State. 


ACKNOWLEDGMENTS 


We thank Kevin Coniff for providing the plants of Hy- 
locereus undatus and the UCLA-Ben Gurion University 
Program of Cooperation for financial support through the 
generous gift of Dr. Sol Leshin and the continuing dedi- 
cation to such projects by Professor Samuel Aroni. 


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MADRONO, Vol. 49, No. 4, pp. 237-255, 2002 


SIX NEW SPECIES AND TAXONOMIC REVISIONS IN MEXICAN 
GAUDICHAUDIA (MALPIGHIACEAE) 


STEVEN L. JESSUP 
Department of Biology, Southern Oregon University, Ashland, OR 97520 
Jessup @ sou.edu 


ABSTRACT 


Six new species in the genus Gaudichaudia are described and sectional taxonomy in the genus is 
revised in accordance with findings from molecular analysis of chloroplast and nuclear genomes. Gau- 
dichaudia cycloptera, G. chasei, and G. mcvaughii are removed from section Gaudichaudia and placed 
in section Cyclopterys. Gaudichaudia krusei and G. subverticillata are removed from section Gaudi- 
chaudia and placed in section Archaeopterys. Gaudichaudia hirtella, comb. nov. is placed in section 
Oligopterys to accommodate the finding that Aspicarpa, at least in part, is nested within Gaudichaudia. 
Three nothosections are named to accommodate new species described as amphiploids among sections. 
Gaudichaudia implexa is described as a new amphiploid species formed from lineages in section Trito- 
mopterys and section Gaudichaudia (nothosection Tritomochaudia). Gaudichaudia symplecta and Gau- 
dichaudia synoptera are described as two new amphiploid species formed from lineages in section Tri- 
tomopterys and section Cyclopterys (nothosection Cyclotomopterys). Gaudichaudia zygoptera and Gau- 
dichaudia intermixteca are described as two new amphiploid species formed from lineages in section 
Tritomopterys and section Zygopterys (nothosection Zygotomopterys). Gaudichaudia andersonii is de- 
scribed as a new amphiploid species formed from lineages within section Cycloptera. 


RESUMEN 


Se describen seis especies nuevas en el género Gaudichaudia y se revisa la taxonomia seccional en el 
género de acuerdo con los hallazgos del andlisis molecular del cloroplasto y de los genomas nucleares. 
Se guitan Gaudichaudia cycloptera, G. chasei y G. mcvaughii de la secci6n Gaudichaudia y se colocan 
en la secci6n Cyclopterys. Se quitan G. krusei y G. subverticillata de la secci6n Gaudichaudia y se 
colocan en la seccion Archaeopterys. Se pone Gaudichaudia hirtella, comb. nov. en la secci6n Oligopterys 
para acomodar el hallazgo de que Aspicarpa por lo menos en parte, se anida dentro de Gaudichaudia. 
Se nombran tres nothosecciones para acomodar la nuevas especies descrita como anfiploide entre sec- 
ciones. Se describe Gaudichaudia implexa como una nueva especie anfiploide formada de linajes en la 
seccion Tritomopterys y la secci6n Gaudichaudia (nothosecci6n Tritomochaudia). Se describen Gaudi- 
chaudia symplecta y Gaudichaudia synoptera como dos especies nuevas del anfiploide formada de linajes 
en la seccion Tritomopterys y la secci6n Cyclopterys (nothosecci6n Cyclotomopterys). Se describen Gau- 
dichaudia zygoptera y Gaudichaudia intermixteca como dos especies nuevas del anfiploide formada de 
linajes en la secci6n Tritomopterys y la secci6n Zygopterys (nothosecci6n Zygotomopterys). Se describe 
el Gaudichaudia andersonii como una especie nueva del anfiploide formado de linajes dentro de la seccién 
Cycloptera. 


Key Words: Amphiploid, Gaudichaudia, polyploid complex, Malpighiaceae, nothospecies, systematics. 


Gaudichaudia (Malpighiaceae) is a genus of 
woody vines, vining shrubs, and suffrutescent sub- 
shrubs inhabiting xeric to mesic habitats in Me- 
soamerica and northern parts of South America. 
Most of the diversity in Gaudichaudia is geograph- 
ically concentrated in central, western and southern 
Mexico with variable wide-ranging lineages reach- 
ing into northeastern and northwestern Mexico, and 
south into Central America and northern South 
America. Several regional and narrow endemics oc- 
cur in central Mexico south of the Tropic of Cancer 
and west of the Isthmus of Tehuantepec. Gaudi- 
chaudia was monographed by Franz Niedenzu in 
1928, but little progress was made in understanding 
the genus until the work of Anderson in the 1980’s. 
Niedenzu attempted to coordinate the prior works 
of de Candolle, Adrien de Jussieu, Sessé and Mo- 
cino, Kunth, Chodat, Rose, and others who had de- 


scribed taxa in the genus in the previous century. 
Niedenzu himself added numerous names, con- 
structing an elaborate taxonomic system that in- 
cluded two subgenera and three sections covering 
eleven species and twenty-three infraspecific taxa. 
Although Niedenzu proliferated names in the ge- 
nus, he also significantly reduced the nomenclatural 
superfluity in Gaudichaudia by listing twenty-nine 
names from five genera in synonomy. While Nie- 
denzu’s work was a significant step in clarifying the 
systematics of Gaudichaudia, the profusion of 
names he introduced had the opposite effect. Nie- 
denzu’s taxa were largely based on narrow taxo- 
nomic concepts that relied on characters that are 
now clearly seen as variable within lineages and 
sometimes plastic even within a single individual. 
For example, he used the number of glands on the 
caylx to diagnose forms within varieties within sub- 


238 


species in both G. cycloptera and G. cynanchoides. 
Furthermore, Niedenzu’s reliance on floral features 
in Gaudichaudia, which like other Malpighiaceae 
is distinctive in the ancient (Taylor and Crepet 
1987) and conservative features of its flowers (An- 
derson 1979), yielded a taxonomy that overlooked 
some distinctive species while emphasizing rela- 
tively minor variations in widespread taxa. To fur- 
ther confound taxonomic matters in Gaudichaudia, 
the herbarium specimens available to Niedenzu for 
his monograph were largely collected prior to 1910, 
and most of the collections he cited, even those 
designated as types, are lacking specific locality 
data. Some of the types are fragmentary or consist 
of immature stages, in a few cases without fruits or 
mature flowers. The names based on inadequate 
type material may never be clearly referable to a 
lineage and are therefore of little or no taxonomic 
value today. 

Taxonomy in Gaudichaudia has been revisited in 
recent years by Anderson (1987, 1993), who clar- 
ified some of the nomenclature and described three 
new species that were missed by Niedenzu and his 
predecessors. Anderson (1993) accepted Nieden- 
zu’s sectional taxonomy: section Tritomopterys, 
section Zygopterys, and section Gaudichaudia, all 
of which are distinguished on fruit wing morphol- 
ogy. Fruit wing morphology in section Tritomop- 
terys (G. albida Schlecht. & Cham., G. diandra 
(Nied.) Chodat, G. hexandra (Nied.) Chodat) is 
characterized by highly asymmetric lateral wings 
(Fig. 1), and in section Zygopterys (G. galeottiana 
(Nied.) Chodat) by distally rounded, free and sym- 
metric lateral wings, and a well developed posterior 
wing (Fig. 2). Section Gaudichaudia is character- 
ized by a rounded fruit wing with the apex some- 
times notched, but with lateral wings scarcely free 
at the apex and completely confluent at the base 
(Fig. 2). Anderson (1993) includes G. cynanchoides 
H. B. K., G. cycloptera (DC.) W. R. Anderson, G. 
subverticillata Rose, G. chasei W. R. Anderson, G. 
mcvaughii W. R. Anderson, and G. krusei W. R. 
Anderson in section Gaudichaudia (Fig. 2 in part, 
Figs. 3, 4). Anderson’s (1987, 1993) published 
work on Gaudichaudia recognizes ten species, in- 
cluding the three he described (G. mcvaughii, G. 
krusei, G. chasei). Anderson (1993) has shown that 
all ten of the recognized species are diploids with 
n = 40 meiotic chromosome pairs. Anderson (per- 
sonal communication) also recognizes several spe- 
cies in Gaudichaudia that have yet to be published. 
Three of those (species-A, “‘G. intermedia’; spe- 
cies-B, ““G hirsuta’; species-C, ““G. velutina’’) are 
clearly in section Tritomopterys and closely related 
to the G. albida—G. diandra—G. hexandra com- 
plex (Fig. 1). At least some of the undescribed spe- 
cies in section Tritomopterys are also known dip- 
loids as shown by Anderson’s (1993) chromosome 
counts published as G. albida sens. lat. Another 
undescribed species in Gaudichaudia is a very dis- 
tinctive close relative of G. cycloptera (species-D, 


MADRONO 


[Vol. 49 


““G. mexiae’’) that is narrowly restricted to lower 
elevations in remote areas on the west slopes of 
coastal Jalisco, Nayarit and Sinaloa (Fig. 3). Judg- 
ing from morphology of the samaras and the lim- 
ited chromosome counts available, all of these un- 
described elements of Gaudichaudia would be eas- 
ily accommodated within the sections established 
by Niedenzu as endorsed by Anderson. While the 
sectional taxonomy in Gaudichaudia based on fruit 
wing morphology is supported by overall morpho- 
logical similarity of the fruits, recent evidence from 
comparative analyses of the chloroplast and nuclear 
genomes (Jessup 1994, 2002), clearly shows that 
section Gaudichaudia as previously constructed is 
not monophyletic. In this paper I briefly summarize 
molecular evidence supporting sectional revisions 
in Gaudichaudia and propose a new combination 
resulting from the finding that Aspicarpa is, at least 
in part, nested within Gaudichaudia (Cameron et 
al. 2001; Davis et al. 2001). I propose three new 
sections and three new nothosections in Gaudi- 
chaudia and reclassify anomalous species. Six new 
species are described. 


MOLECULAR EVIDENCE 


Total genomic DNA was extracted from 118 
Gaudichaudia specimens collected across a wide 
geographic region of Mexico. Extraction protocol 
and restriction site analysis followed procedures 
outlined in Dowling et al. (1996). Accession data 
and detailed laboratory procedure is presented in 
Jessup (1994). Restriction sites found on the chlo- 
roplast genome are presented in Table 1. Twenty- 
one informative sites were produced with 10 of the 
20 enzymes screened. Thirty-four distinct chloro- 
plast haplotypes were discovered, each present in 
between 1 and 29 specimens. Restriction site data 
for representatives of each of the 34 haplotypes is 
given in Table 2. Phylogenetic analysis of the 
cpDNA data was performed with Hennig 86 v.1.5 
(Farris 1989). Autapomorphies were excluded from 
the analysis. The procedure “‘mhennig*’’ was used 
to search for the shortest possible trees. Branch 
swapping on the shortest trees found by multiple 
initial passes with different OTU addition sequenc- 
es produced a set of trees that was then used as the 
starting point for procedure “‘bb’’, an extended 
branch-swapping algorithm that searches all possi- 
ble trees. As a check on thoroughness of the heu- 
ristic algorithms, a separate run of the procedure 
‘ie; bb’’, an implicit enumeration algorithm fol- 
lowed by branch-swapping, was used to find all of 
the shortest trees, which were then used to compute 
the majority rule consensus tree using PAUP 
(Swofford 1990). 

The majority rule consensus tree of 26 equally 
parsimonious trees obtained from restriction site 
characters is presented in Fig. 5. Terminal branches 
in the cpDNA tree represent the 34 distinct ge- 
nomes discovered in the sample of 118 plants. Ro- 


2002] JESSUP: GAUDICHAUDIA 


Section Tritomopterys 
@ G.albida 

A G.diandra 
3+ G. hexandra 


(undescribed species) 
% G.sp. ined. “intermedia” 
©. G.sp. ined. “hirsuta” 

¥€ G.sp. ined. “velutina” 


Fic. 1. Geographic distribution of species and fruit morphology in representative collections from section Tritomop- 
terys. A-E, G. albida, A) Jessup 4041, Oaxaca, B) Jessup 4052, C) Anderson 13275, Mexico, D) Jessup 4067, Guerrero, 
E) Jessup 4056, Chiapas; F—J, G. diandra, F) Jessup 4088, Nayarit, G) Jessup 4066, Guerrero, H) Anderson 13309, 
Michoacan, I) Jessup 4032, Michoacan, J) Anderson 12937, Morelos; K—N, G. sp. ined. “‘intermedia’’, K) Anderson 
13225, Chiapas, L) Jessup 4051, Oaxaca, M) Anderson 13224, Chiapas, N) Jessup 4055, Chiapas; O, G. sp. ined. 


“hirsuta”, Jessup 4048, Oaxaca; P-R, G. sp. ined. “‘velutina”’, P) Jessup 4058, Chiapas, Q) Jessup 4060, Chiapas, R) 
Jessup 4054, Oaxaca. Samara illustrations are approximately | x. 


240 


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[Vol. 49 


Section Gaudichaudia 
@ G. cynanchoides 


{J 
OP - uty Zz 


Section Zygopterys 
Y G. galeottiana 


Fic. 2. 


Geographic distribution of species and fruit morphology in representative collections from section Gaudi- 


chaudia and section Zygopterys. A—L, section Gaudichaudia: A) Jessup 4078, Jalisco, B) Koch 82260, Mexico, C) 
Jessup 4077, Jalisco, D) Nicolas 5078, Puebla, E) Jessup 4075, Michoacan, F) Anderson 13296, Michoacan, G) Jessup 
4111, Hidalgo, H) Anderson & Laskowski 3884, Michoacan, I) Jessup 4112, Hidalgo, J) Jessup 4109, Hidalgo, K) 
McVaugh 13204, Michoacan, L) Anderson & Laskowski, Michoacan; M-—Q, section Zygopterys: M—N) Galeotti X. 
1844, Tutepeji, Puebla (type), O) Jessup 4038, Puebla, P) Anderson & Laskowski 4294, Puebla, Q) Anderson 13128, 


Oaxaca. Samara illustrations are approximately | x. 


bustness of the phylogenetic estimate is indicated 
by the fact that all but two branches are supported 
by 100% of the trees in the consensus calculation. 
Detailed geographic mapping of the cpDNA hap- 
lotypes and taxonomic assignment of plants carry- 
ing those chloroplast genomes is presented in Jes- 
sup (1994). The cpDNA phylogeny taken together 
with morphological features supports recognition of 
six sections in Gaudichaudia. Sections Tritomop- 
terys and Zygopterys as recognized by Anderson 
(1993) are well supported as monophyletic by the 
cpDNA analysis. Section Gaudichaudia is mono- 
phyletic with the removal of G. cycloptera, G. 
mevaughii, G. chasei, G. subverticillata, and G. 
krusei. The geographically widespread species G. 
cynanchoides is shown to be a genetically diverse 
assemblage of morphologically similar lineages. 
The methods used in this study were unable to re- 
solve the basal polytomy. The cpDNA phylogeny 
is illustrated as an unbranched network (Fig. 5) to 
emphasize the lack of phylogenetic resolution 
among the sections in Gaudichaudia. 

To accommodate the species removed from sec- 
tion Gaudichaudia two new sections are proposed 
here; section Cyclopterys and section Archaeopter- 


ys. Section Cyclopterys is well supported by the 
cpDNA analysis as monophyletic and includes G. 
cycloptera, G. mcvaughii, and G. chasei, as well as 
the undescribed elements discussed above. Section 
Archaeopterys is paraphyletic and, based on mor- 
phological evidence and evidence from analysis of 
nuclear DNA discussed below, comprises a cluster 
of phylogenetically basal lineages within the genus. 
There is not, however, sufficient evidence to root 
the cpDNA network with section Archaeopterys. 
Section Oligopterys is proposed here to accom- 
modate species of Aspicarpa that have been found 
to nest within Gaudichaudia, as reported by Cam- 
eron et al. (2001), Davis et al. (2001) and Jessup 
(1994). 


TAXONOMIC REVISIONS AND NEW SPECIES 


Revision of sectional taxonomy of the diploid 
species. 


Sectio Cyclopterys Jessup sect. nov. (Fig. 3)— 
TYPE: Mexico, Michoacan: Tuzantla, 110 km N 
Huetamo, rd to Zitacuaro, 670 m. 15 October 
1988, Jessup 4033 (Holotype MICH; isotypes 
MO, UC, IEB). 


2002] JESSUP: GAUDICHAUDIA 241 


— - 
a Section Cyclopterys 


66 G. cycloptera 
@ G. chasei 


OF G. mcevaughii 


G. sp. ined. 
“mexiae” 


G. mevaughii 


Fic. 3. Geographic distribution of species and fruit morphology in representative collections from section Cyclopterys. 
A-D, G. cycloptera, A) Jessup 4029, Guerrero, B) Jessup 4083, Jalisco, C) Jessup 4033, Michoacan, D) McVaugh & 
Koelz 534, Nayarit; E—G, G. chasei, E) Lyonnet 28, Morelos, F) Jessup 4009, Morelos, G) Bates 3426, Morelos: H) 
G. sp. ined. ““mexiae”, Croat 45249, Nayarit; I-J, G. mcvaughii, 1) Anderson 12699, Colima, J) Koch & Fryxell 82218, 
Guerrero. Samara illustrations are approximately 1X. 


Section Archaeopterys 
@ G subverticillata 


+ G. krusei 


Fic. 4. Geographic distribution of species and fruit morphology in representative collections from section Archaeop- 
terys. A) G. subverticillata, Jessup 4087; B) G. krusei, Jessup 4069. Samara illustrations are approximately 1X. 


242 


TABLE 1. INFORMATIVE CPDNA RESTRICITON SITES FOUND 
IN 118 SPECIMENS. Each restriction site is represented by 
a letter code and referred to in the data matrix in Table 2. 


Enzyme Informative site(s) Site code 
Ban II 96> 82+ 1.4 Q 
Bcl I 4457+? H 
3.6 > 2.1 + 1.5 O 
Se) => 20) SPL) T 
Cla I 19.0 > 8.2 + 10.8 E 
2.8 > 1.2 + 1.6 F 
2.95 1.3 + 1.6 G 
Dra I 6.15 > 5.48 + .67 K 
5.48 > 4.2 + 1.28 L 
31 Ss Io aP IS M 
Eco RI 3.5) => 340 =P OLS A 
Eco RV 16.5 > 9.5 + 7.0 D 
Hae III ALS <> 1.3 se 2? I 
Lab > 12 se OD J 
16757?+? R 
Mgd) = EY S 
Hine II S19 => SD =P Dell P 
A232) => 2.3 a7 2O UW 
Hpa II 5.1 > 3.6 + 1.5 N 
Xba 3.8 > 2.0 + 1.8 B 
30 => Zo) ar ILS (C 


To promote nomenclatural clarity, and to bring 
evidence from DNA analyses to bear on taxonomic 
circumscription of this section, under provisions of 
the ICBN Saint Louis Code, Article 22.6 (Greuter 
et al. 2000), I am designating a different holotype 
for section Cyclopterys than the type for the name 
on which the section is based, which is Gaudi- 
chaudia cycloptera (DC.) Anderson. The earliest 
name for the species according to Anderson (1987) 
is Hiraea? cycloptera DC. Prodr. 1: 586. 1824, and 
the type for that name is an unnumbered illustration 
among de Candolle’s collection of plates, referred 
to simply as “fl. mex. ic. ined.” (de Candolle 
1824), a plate presumably copied from Sessé and 
Mocino’s original painting in the Jcones Florae 
Mexicanae, now at HU. The original watercolor il- 
lustration, the presumed virtual type, was titled, 
“Triopteris oblongifolia’ (HU accession number 
6331.0888). Many of the 279 names based on types 
that are plates or copies of plates from the Sessé 
and Mocino expedition were never vouchered with 
a collection (McVaugh 1980). This is most likely 
the case for G. cycloptera. So far as I am aware 
there is no known specimen corresponding to the 
plate, though the possibility remains that a speci- 
men will be found in MA or among the collections 
belonging to a recipient of specimens collected on 
the Sessé and Mocino expedition. Establishment of 
section Cyclopterys is warranted on the evidence 
from DNA analyses together with morphological 
comparisons. I am designating a type for the sec- 
tion that can be subjected to further DNA analysis. 
The specimen designated as holotype for the sec- 
tion carries cpDNA haplotype Z, one of five hap- 
lotypes found in a well supported monophyletic G. 


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[Vol. 49 


cycloptera (Fig. 5). G. cycloptera bearing cpDNA 
haplotype Z has not been discovered as a compo- 
nent in any amphiploid formed among sections in 
Gaudichaudia, although the closely related cpDNA 
haplotype AB is shared between some lineages 
within section Cyclopterys, in particular between G. 
cycloptera and G. mcvaughii, (Jessup 1994). 

Herbae fruticesve scandentes e basi fruticosa, 
caulibus longissimis volubilibus in vegetationem 
sustinentem extendentibus 4—12 m. Folia decussata 
et sensim decrescentia a basi ad ramos distales flo- 
rescentes, late elliptica ad ovata, subsessilia ad dis- 
tincte petiolata, basi hastata, cordata, truncata, vel 
cuneata. Indumentum partes maturas vegetativas to- 
tae plantae tegens, dense sericeum vel sparse hir- 
sutum ad grosse hispidum, e trichomatibus ramosis 
constans, trabeculis laevibus ad sinuosis, adpressis 
ad erecto-patentibus vel subaristatis. Flores omnes 
chasmogami, ad anthesin 1.5—3 cm lati, in umbellis 
geminatis 4-floris in axillis foliorum reductorum vel 
in dichasiis foliosis axillaribus dispositi, interdum 
in thyrsum elongatum distaliter fasciculati. Samarae 
orbiculares ad cordatae, 1—2 cm in diametro, alis 
lateralibus symmetricis, ad carpophorum confluen- 
tibus et rotundatis ad acutis vel late retusis, lobis 
apicalibus alarum obtusis ad acutis incisuram sinu 
lato ad angusto facientibus, vel alis lateralibus con- 
fluentibus ad apicem, margine distali itaque integra. 
Superficies dorsalis fructus alam distinctam inter- 
dum prominentem vel reticulum congestum pen- 
nularum supra nucem ferens. 

Vines and vining shrubs from a shrubby base, 
the longest twining stems reaching 4—12 m into the 
supporting vegetation. Leaves decussate and grad- 
ually reduced from the base to the distal flowering 
branches, broadly elliptic to ovate, subsessile to 
distinctly petiolate, leaf bases hastate, cordate, trun- 
cate or cuneate. Vesture thinly or densely sericeous 
or sparsely hirsute to coarsely hispid, comprising 
branched trichomes, the trabeculae smooth to sin- 
uose and appressed to erect spreading or subaris- 
tate, the indumentum covering mature vegetative 
parts of entire plant. Flowers all chasmogamous, 
1.5—3 cm in diameter at anthesis, on paired four- 
flowered umbels in the axils of reduced leaves, or 
in axillary leafy dichasia, sometimes clustered dis- 
tally to form an elongate thyrse. Samaras orbicular 
to cordate, 1—2 cm in diameter, the lateral wings 
symmetric, confluent and rounded to acute or 
broadly retuse at the carpophore, the apical wing 
lobes obtuse to acute, forming a notch with a broad 
to narrow sinus, or the lateral wings apically con- 
fluent, the distal margin thus entire. The dorsal sur- 
face of the fruit bearing a distinct, sometimes prom- 
inent wing or a congested nexus of winglets over 
the nut. 

Phylogenetic analysis of restriction sites in the 
chloroplast genome of Gaudichaudia (Fig. 5) clear- 
ly supports Niedenzu’s section Tritomopterys con- 
taining lineages related to G. albida. The phylo- 
genetic analysis also supports section Gaudichau- 


2002] JESSUP: GAUDICHAUDIA 243 


TABLE 2. CPDNA RESTRICTION SITE MATRIX FOR HAPLOTYPES. Restriction site codes refer to Table |. Each haplotype 


is designated by 1-2 letters. 


Haplotype Specimen Estado 
A J4047 Oaxaca 
B J4059 Chiapas 
(C J4055 Chiapas 
D J4015 Morelos 
E J4036 Michoacan 
F J4056 Chiapas 
G J4103 Zacatecas 
H J4064 Guerrero 
I J4087 Nayarit 
JJ J4069 Guerrero 
K A13216 Oaxaca 
J J4027 Guanajuato 
M A3707 Jalisco 
N J4102 Sinaloa 
O J4008 Morelos 
P J4035 Michoacan 
Q J4037 Michoacan 
S J4081 Jalisco 
ie J4018 Morelos 
R J4007 Guanajuato 
U J4112 Hidalgo 
V J4042 Oaxaca 
W J4049 Oaxaca 
xX A12990 Oaxaca 
a’ J4029 Guerrero 
7, J4033 Michoacan 
AA J4030 Michoacan 
AB J4024 Mexico 
AC A12699 Colima 
AD J4038 Puebla 
AE J4009 Morelos 
AF A4510 Guerrero 
AG J4039 Puebla 

Putative 

outgroup Apsicarpa 


Sites 
(refer to Table 1) 


ABCDEFGHIJKLMNOPORSTU 


000101000000110000000 
000111000000110000001 
010111000000110000001 
000111000100110000001 
000111000100100000000 
000110000100110000001 
000111000000001010010 
001111000000100000000 
101111000000100000000 
011111000000100000000 
010111101000110100000 
000111101000110100000 
000111101000010100000 
000110101000110100000 
000111101000110000000 
100111000000101011011 


OTA ARG OOOO CAs AO mea Os A 
AO) IE OONONCNOMONONGMENO)AL SE O}AEO) 
TL) LNAI ITO O)ONCNONON ONC) ALTOHAL 110) 4116) 


LOLTEALOOCCOPOOLOLLOT0 
101111000000001011010 
001111010010100000100 
101111010010100000100 
101111000010100000100 
101001000000100000000 
101001000000100000000 
001001000000100000000 
101011000000100000000 
101010000000100000000 
101111000001100000101 
101111000000100000000 
101111000000000000000 
001111000001100000101 


101111000000100000000 


dia containing lineages related to G. cynanchoides, 
as well as section Zygopterys containing the single 
species, G. galeottiana. In the phylogenetic analy- 
sis section Zygopterys and section Tritomopterys 
were each found to be monophyletic. Based on the 
molecular data, however, a monophyletic section 
Gaudichaudia cannot include G. cycloptera, G. 
mevaughii, G. subverticillata, G. krusei, or G. chas- 
ei as proposed by Anderson (1993). Those species 
are therefore moved to new sections as detailed be- 
low. Section Cyclopterys is strongly supported as 
monophyletic by the chloroplast DNA phylogeny, 
comprising the distinctive and geographically wide- 
spread type species, G. cycloptera and the narrow 
endemic, G. mcvaughii, known only from scattered 
and narrowly delimited low elevation localities on 
the Pacific coast of Colima, Guerrero, and Oaxaca. 
In addition to sharing, in some populations, a dis- 
tinct chloroplast haplotype with G. cycloptera, G. 
mcvaughii also shares morphological features of the 
fruit and habitat attributes with G. cycloptera. The 


elaboration of a dorsal wing in the fruits of both 
species, and the size and shape of the fruits provide 
further evidence that these taxa are closely related. 
Both G. cycloptera and G. mcvaughii are plants of 
mesic understory or forest edge thickets. Both are 
known diploids without cleistogamous flowers. 
Although the branch supporting the single col- 
lection of G. chasei included in the analysis (Fig. 
5) is unresolved with respect to branches support- 
ing sections, G. chasei has several morphological 
features in common with members of section Cy- 
clopterys, notably the expanded dorsal keel on the 
samara, and lateral fruit wings that are basally con- 
fluent. Gaudichaudia chasei, which Anderson 
(1987) argues is morphologically close to G. 
mevaughii, is known from only a small area in Mo- 
relos. The habitat where it grows is more similar to 
the mesic understory habitat of G. cycloptera from 
adjacent Edo. Mexico than the habitat of the typical 
species of section Gaudichaudia (G. cynanchoi- 
des), which grows in the xeric matorral of the Al- 


244 MADRONO [Vol. 49 


aie 13 ay) 
; mexia 


ARCHAEOPTERYS 


cycloptera mevaughii 


subverticillata Oo) @) @ AB) CO chasei 


CYCLOPTERYS 


OLIGOPTERYS (AF) 


cynanchoides 
sens. lat. 


hirtella 


galeottiana 


Majority rule consensus phylogeny of cpDNA haplotypes in Gaudichaudia. 


Fic. 5. Majority rule consensus phylogeny of cpDNA haplotypes showing the relationship of sections in an unrooted 
network. Each terminal branch represents a distinct cp DNA haplotype defined by a unique configuration of restriction 
sites. Letters in circles designate haplotypes referred to in Table 2. Representative fruits of species of non-hybrid origin 
illustrate the range of fruit morphology in each section. Position of the fruits relative to branches of the network is 


approximate. 


toplano Mexicano (Fig. 2). Gaudichaudia chasei, 
like G. cycloptera and G. mvaughii, lacks the cleis- 
togamous flowers that are a prevalent feature in G. 
cynanchoides. Though perhaps a divergent member 
of section Cyclopterys, G. chasei clearly fits better 
here than in the other sections and is not otherwise 
sufficiently distinct to justify a separate section. 
The molecular evidence, absence of cleistogamy 
and other morphological evidence mentioned by 
Anderson (1987), the similarity of geographic 
range and habitats, and the evidence from Ander- 
son’s (1993) study of chromosomes, indicating that 
all are diploids with n = 40 meiotic chromosome 
pairs, taken together strongly supports recognition 
of section Cyclopterys, containing G. cycloptera, G. 
mevaughii, G. chasei, and the undescribed lineage 


from Nayarit and Jalisco (G. sp. ined. “‘mexiae’’) 
as a discernable monophyletic clade within Gau- 
dichaudia. 

Section Gaudichaudia sensu Anderson (1993) 
included G. subverticillata and G. krusei, two spe- 
cies that he argued are closely related (Fig. 4). Both 
are subshrubs with rounded fruit wings resembling 
in outline the fruits of G. cynanchoides. The phy- 
logeny based on chloroplast genomes does not, 
however, support inclusion of those species in a 
monophyletic section Gaudichaudia. The fruits are 
generally similar in size and shape to those of sec- 
tion Gaudichaudia, and are quite distinct from 
fruits of species included in section Tritomopterys. 
Nevertheless, G. subverticillata and G. krusei are 
unambiguously placed as outgroups to the G. al- 


2002] 


bida complex (section Tritomopterys) in the parsi- 
mony analysis of the chloroplast genomes (Fig. 5). 
Taken together, G. subverticillata, G. krusei and 
section Tritomopterys are monophyletic. In view of 
the substantial divergence in gross morphology be- 
tween G. subverticillata and G. krusei on the one 
hand and members of section Tritomopterys on the 
other, however, their assignment to section Trito- 
mopterys is untenable. Gaudichaudia subverticil- 
lata and G. krusei are certainly similar in mor- 
phology: both are suffrutescent shrublets lacking 
cleistogamy, and both are known diploids. They oc- 
cupy similar habitats, and are both narrow endem- 
ics in a region of southwestern Mexico rich in nar- 
row endemics from many groups of plants, a fact 
that suggests the area may have served as an an- 
cient refugium where phylogenetically basal line- 
ages might be expected to persist. Furthermore, the 
comparative analysis of nuclear genomes based on 
randomly amplified DNA (Jessup 2002) places G. 
subverticillata and G. krusei as sister taxa close to 
members of section Cyclopterys, but on the periph- 
ery of the minimum spanning tree of Jaccard dis- 
tances. The evidence, taken together, corroborates 
the hypothesis that G. subverticillata and G. krusei 
are basal within Gaudichaudia. They are apparently 
closely related to each other, and more closely re- 
lated to section Tritomopterys and section Cyclop- 
terys than they are to section Gaudichaudia. To ac- 
commodate sectional placement of G. subverticil- 
lata and G. krusei the following new section is es- 
tablished. 


Sectio Archaeopterys S. L. Jessup sect. nov. 


Fruticuli suffrutescentes raro ramosi, ramis max- 
imis plerumque 0.5—1.0 m altis e basi lignosa. Folia 
brevipetiolata ad subsessilia, binata vel verticillata 
terna. Indumentum partes maturas vegetativas totae 
plantae tegens, sparse vel dense sericeum ad velu- 
tinum, e trichomatibus ramosis constans, trabeculis 
laevibus ad sinuosis et adpressis ad erecto-patenti- 
bus. Flores omnes chasmogami, in umbellis 4-floris 
verticillatis e nodis distalibus caulium primorum 
vel in dichasiis brevibus axillaribus a foliis caulinis 
superioribus subtentis dispositi. Samarae orbicular- 
es ad cordatae, alis lateralibus symmetricis, proxi- 
maliter acutis vel ad carpophorum infirme retusis, 
lobis distalibus alarum obtusis ad acutis, incisura 
apicali sinum obtusum ad acutum facienti, superfi- 
cies dorsalis fructus alam rudimentariam ad prom- 
inentem vel reticulum humile pennularum supra nu- 
cem ferens. 

Suffrutescent seldom branching shublets, the 
largest branches mostly 0.5—1.0 meter high from a 
woody base. Leaves short petiolate to subsessile, 
paired or in whorls of three. Vesture thinly or 
densely sericeous to coarsely velutinous, compris- 
ing branched trichomes, the trabeculae smooth to 
sinuose and appressed to erect spreading covering 
mature vegetative parts of entire plant. Flowers all 


JESSUP: GAUDICHAUDIA 


245 


chasmogamous, in verticillate four-flowered umbels 
from distal nodes of main stems, or on short axil- 
lary dichasia subtended by the upper stem leaves. 
Samaras orbicular to cordate, the lateral wings sym- 
metric, proximally acute to weakly retuse at the car- 
pophore, the distal wing lobes obtuse to acute, the 
apical notch forming an obtuse to acute sinus, the 
dorsal surface of the fruit bearing a rudimentary to 
prominent wing or low nexus of winglets over the 
nut. 


Type: Gaudichaudia subverticillata Rose 


The problem of long branch attraction makes in- 
terpretation of branching order among sections in 
Gaudichaudia problematic. Basal lineages with few 
close relatives in a genus of otherwise closely re- 
lated species complexes are particularly sensitive to 
inaccurate placement. Placement of section Ar- 
chaeopterys should therefore be tempered with 
skepticism until Tribe Gaudichaudieae can be ana- 
lyzed as a whole with sequence data chosen spe- 
cifically for this problem. 

With the foregoing sectional reassignments, sec- 
tion Gaudichaudia now comprises only vines with 
both chasmogamous and cleistogamous flowers 
producing rounded, essentially symmetric (cynan- 
choid) samaras. Section Gaudichaudia as revised 
includes only G. cynanchoides sens. lat., which is 
widespread in mesic to xeric ruderal habitats on the 
Altoplano Mexicano (Fig. 2). Although the only ac- 
cepted species remaining in section Gaudichaudia 
is G. cynanchoides, that name as now used encom- 
passes a diverse assemblage of microspecies prop- 
agating largely through abundant production of 
cleistogamous fruits. As evident in the series of dis- 
tinct chloroplast haplotypes and diverse morpho- 
types (Jessup 1994), G. cynanchoides is a species 
complex that should eventually be resolved into 
several closely related but geographically distinct 
species or subspecies. 

In addition to supporting the realignment of sec- 
tional taxonomy of recognized members of Gau- 
dichaudia, the results of molecular research clearly 
indicate that the genus Gaudichaudia itself, as con- 
ventionally delimited, is paraphyletic. My molecu- 
lar studies (Jessup 1994, 2002) included the genus 
Aspicarpa as the outgroup in several analyses, but 
in the maximum likelihood analysis (unpublished 
results) Aspicarpa was found to nest within Gau- 
dichaudia in a polytomy with section Cyclopterys, 
section Archaeopterys, and section Gaudichaudia. 
That result is corroborated by recent molecular 
phylogenies of the Malpighiaceae (Cameron et al. 
2001; Davis et al. 2001) showing that at least some 
species now included within Aspicarpa are nested 
within Gaudichaudia. To accommodate the evi- 
dence demonstrating that Aspicarpa is, at least in 
part, nested within Gaudichaudia, | here establish 
a new section in Gaudichaudia to include those el- 
ements of Aspicarpa that are properly considered 


246 


species within a monophyletic Gaudichaudia. The 
type of the new section is established by the fol- 
lowing new combination. 


Gaudichaudia hirtella (Rich.) Jessup, comb. nov. 


Aspicarpa hirtella Rich., Mem. Mus. Paris 2: 
BID, Nest). 

Aspicarpa urens Lagasca, Gen. Sp. Pl. Nov. 1. 
1816. 

Aspicarpa pruriens Desv., Desf. Cat. Hort. Paris. 
Gl, Bo ZIBB, M329). 

Gaudichaudia urens Chodat in Bull. Soc. Bot. 
Geneve Z Sew IDX, IOI, 


Section Oligopterys Jessup sect. nov. 


Fruticuli suffrutescentes caulibus paucis ad mul- 
tis erectis vel decumbentibus e basi lignosa. Folia 
opposita vel verticillata, basi rotundata ad cordata, 
sessilia ad subsessilia. Flores et chasmogami et 
cleistogami. Fructus sine ala laterali, nuculum 
oblique affixum sine carpophoro producentes, crista 
vel jugo dorsali instructi, sine ala dorsali. 


Type: Gaudichaudia hirtella (Rich.) Jessup, comb. 
nov. 


Suffrutescent shrublets with few to many erect to 
decumbent or trailing stems from a woody base. 
Leaves opposite or whorled, rounded to cordate at 
base, sessile to subsessile. Flowers both chasmo- 
gamous and cleistogamous. Fruits lacking a lateral 
wing, forming an obliquely attached nutlet lacking 
a carpophore, with a low dorsal crest or ridge, with- 
out a dorsal wing. 

Other species now placed in Aspicarpa will like- 
ly emerge as elements of Gaudichaudia. Only G. 
hirtella is included here in section Oligopterys 
since that was the species included in my study of 
chloroplast DNA phylogeny in Gaudichaudia. Al- 
though the chloroplast DNA parsimony analysis 
majority rule consensus tree does not resolve the 
branch supporting Aspicarpa with respect to 
branches supporting other sections, Aspicarpa is 
distinct enough in fruit morphology and plant habit 
to warrant a separate section. Circumstantial evi- 
dence from studies with labeled RAPD probes (Jes- 
sup 2002) support placement of section Oligopterys 
close to section Zygopterys. When amplified PCR 
products from G. galeottiana were probed against 
blots of RAPD gels in that study, lanes representing 
Aspicarpa hybridized the probe along with mem- 
bers of section Zygopterys and the intersectional 
amphiploids involving section Zygopterys. The 
probe also weakly hybridized the lane representing 
G. krusei, but did not hybridize lanes representing 
other lineages within Gaudichaudia. In the phenetic 
analysis of randomly amplified DNA (Jessup 1994) 
Aspicarpa clustered with members of section 77i- 
tomopterys on the minimum spanning tree of Jac- 
card distances. Recent DNA sequence analysis of 
generic phylogeny in Malpighiaceae (Cameron et 


MADRONO 


[Vol. 49 


al. 2001; Davis et al. 2001) found Aspicarpa nested 
with members of section Tritomopterys. Definitive 
resolution of the phylogentic placement of sections 
in Gaudichaudia must, however, be deferred until 
DNA sequences for a broader sample of represen- 
tative taxa in tribe Gaudichaudieae are available. 


Taxonomic revisions in the amphiploid complex- 
es. Anderson began the task of resolving reticulate 
ancestry in Gaudichaudia. In addition to his de- 
scriptions of new species and chromosome counts 
in Gaudichaudia, Anderson explored the relation- 
ship of Gaudichaudia to other genera in Tribe Gau- 
dichaudieae (Anderson 1985) and contributed to an 
understanding of the genus through studies of re- 
productive life history traits (Anderson 1980). In 
particular, cryptic self fertilization, which occurs in 
distinctive cleistogamous flowers, is common to all 
lineages examined so far that have n = 80 meiotic 
chromosome pairs. All of the plants examined thus 
far that bear fruits with morphologies intermediate 
between those of plants fitting neatly into the sec- 
tions of Gaudichaudia (as herein defined) are tet- 
raploids, and they all bear cleistogamous flowers. 
The evidence presented by Anderson strongly sup- 
ports his hypothesis that plants bearing fruits with 
intermediate morphologies are amphiploids or the 
products of amphiploids formed among lineages in 
different sections of Gaudichaudia. Results sup- 
porting that hypothesis have been corroborated by 
the results of molecular studies (Jessup 1994, 
2002). 

The amphiploids present a bewildering mélange 
of morphological variation that has heretofore been 
reticent to clean cut species delimitations. The com- 
bination of evidence now available from molecular 
studies and a morphological survey of a large num- 
ber of collections from across the geographic dis- 
tribution of Gaudichaudia in Mexico, however, re- 
veals several discrete elements among the tetra- 
ploids that can be clearly discerned and described 
as new species in Gaudichaudia. Five of the spe- 
cies described here are amphiploids formed among 
lineages from different sections, and one is appar- 
ently formed as an amphiploid among lineages 
within a section. In each case, the amphiploid ori- 
gin of the new species is supported by evidence 
from molecular studies. These are not simple F, 
hybrids, but rather wide-ranging lineages that prop- 
agate via selfing through cryptic self fertilization 
while maintaining outcrossing through chasmoga- 
mous flowers. Floral morphology is remarkably 
uniform across the Tribe Gaudichaudieae, and 
clearly fits the family-specific floral syndrome as- 
sociated with oil bee pollination (Buchmann 1987; 
Vogel 1990). Undescribed diploid lineages identi- 
fied by Anderson (Figs. 1, 3) are introduced and 
discussed in this paper in connection with their 
roles in formation of amphiploids among lineages 
in different sections of Gaudichaudia, and in delim- 
iting new taxa with which they might be confused. 


2002] 


Only amphiploid lineages with unambiguous mor- 
phological attributes and clear support from the 
molecular research are described in this paper. Sev- 
eral additional amphiploid species not treated here 
remain undescribed in Gaudichaudia. 


Nothosections and new amphiploid species. Pro- 
visions are made in the ICBN (Greuter et al. 2000) 
for naming nothotaxa, taxa of known hybrid origin, 
and those provisions are codified in the St. Louis 
Code, Appendix I. A diagnosis or description is not 
required for the naming of notho-subdivisions of 
genera, and such names do not have types, but the 
ICBN requires that names of the parental taxa are 
specified when the name of the nothotaxon is pub- 
lished (Article H.9). However, species that are 
known or suspected to be of hybrid origin need not 
be designated as nothospecies and may be desig- 
nated as species (Article H.3). Nothosections are 
proposed here to contain Gaudichaudia lineages 
that are clearly the products of intersectional hy- 
bridization. The combined evidence from morphol- 
ogy, chromosomal counts and molecular studies 
supports designation of nothosections with unam- 
biguous specification of the sectional source of pa- 
rental lines, even to the point of specifying section- 
al contributions of the pollen and ovules for indi- 
vidual members of species within the nothosec- 
tions. Sectional sources of pollen and ovule parents 
are indicated below for the types and paratypes 
where known. It has not, however, yet been feasible 
to unambiguously identify species-level lineages 
contributing to formation of the species in the no- 
thosections. The species named below can be clear- 
ly circumscribed and assigned to nothosections, but 
which of several possible combinations of species 
within the parental sections gave rise to them re- 
mains unknown or ambiguous. In some cases the 
member species in nothosections proposed here are 
likely the products of complex interbreeding among 
several independently evolving lineages from each 
of the contributing parental sections. With those 
reasons in mind the species proposed here are 
named as species rather than nothospecies. 


Nothosection Tritomochaudia S. L. Jessup notho- 
sect. nov. (Gaudichaudia sect. Gaudichaudia X 
Gaudichaudia sect. Tritomopterys), Fig. 6. Pres- 
ently I am recognizing only a single broadly de- 
limited species. The molecular data, the chro- 
mosome data, and the geographic distribution of 
morphological variation suggest that this taxon 
comprises a swarm of autogamously propagating 
amphiploid microspecies that retain viable chas- 
mogamy. 


Gaudichaudia implexa S. L. Jessup, sp. nov. (Fig. 
6)—TYPE: Mexico. Jalisco: south shore of Lago 
de Chapala, 6.4 km W of Jalisco/Michoacan state 
line, 1620 m, dry thorn scrub in hills above lake, 
3 Nov 1988, Jessup 4076 (Holotype MICH; iso- 
types CHAP, IEB, UC). 


JESSUP: GAUDICHAUDIA 


247 


Haec stirps variabilissima a Gaudichaudia cy- 
nanchoides secedit ala samarae lobos laterales an- 
ticos fere symmetricos ad valde asymmetricos acu- 
tos ad obtusos rotundatosve formanti, lobis sinu ad 
basin acuto ad obtuse angulato separatis, sinu in- 
terdum denticulo e margine antica alae lateralis vel 
in basin sinus ex apice nucis orienti, denticulo raro 
in lobum apicalem rotundatum inter lobos laterales 
alae crescenti. Samara autem Gaudichaudia cynan- 
choides ala laterali margine apicali fere integra gau- 
det, et quamquam ala incisuram apicalem parvam 
exhibet, samara ejus cynanchoidea est et lobis dis- 
cretis lateralibus sinu profundo separatis numquam 
instruit. Gaudichaudia albida, Gaudichaudia dian- 
dra et stirpes affines sectionis Tritomopterygos stat- 
im distinguendae sunt lobis lateralibus alae samarae 
valde asymmetricis, et cauda angusta postica alae 
Samarae vix vel haud confluenti cum ala laterali. 
Haec species interdum valde simulabit Gaudichau- 
dia galeottiana vel Gaudichaudia zygoptera aut alis 
lateralibus symmetricis rectis aut lobo lato postico 
constricto ad nucem aut utroque, sed clare distin- 
guenda est limbo texturae inter alas laterales al- 
amque posticam et alis lateralibus apicibus obtuse 
acutis pro alis plerumque latioribus rotundiorib- 
usque Gaudichaudia galeottiana et Gaudichaudia 
zygoptera; ubicumque autem admiscet speciebus il- 
lis, hae differentiae vix discernendae erit. Chro- 
mosomatum numerus, n = 80. 

This highly variable species is separable from G. 
cynanchoides by the samara wing forming two ap- 
proximately symmetric to strongly asymmetric 
acute to obtuse or rounded anterior lateral lobes 
apically separated by a sinus that varies from acute 
to obtusely angular at the base, the sinus sometimes 
with a small tooth originating from the antical mar- 
gin of the lateral wing or in the base of the sinus 
from the crown of the nut, the tooth rarely devel- 
oping as a rounded apical lobe between the lateral 
lobes of the wing. By contrast the samara in G. 
cynanchoides has the apical margin of the lateral 
wing largely entire, and though it occasionally has 
a small apical notch in the samara wing, the samara 
is cynanchoid and never has discrete lateral lobes 
separated by a deep sinus. Gaudichaudia albida, G. 
diandra and related species in section Tritomopter- 
ys can be immediately distinguished by the strongly 
asymmetric lateral lobes of the samara wing, and 
by the narrow posterior tail of the samara wing that 
is scarcely or not at all confluent with the lateral 
wing. In some cases this species closely resembles 
G. galeottiana or G. zygoptera in the development 
of symmetric upright lateral samara wings or a 
broad postical lobe constricted where it attaches to 
the nut. A few specimens bear both of those char- 
acteristics, but it is clearly distinguished by a flange 
of tissue between the lateral wings and the postical 
wing, and by the lateral wings with bluntly pointed 
apices rather than the generally broader more 
rounded wings of G. galeottiana and G. zygoptera. 
However, where it hybridizes with those species the 


248 


Fic. 6. 


MADRONO 


[Vol. 49 


Nothosection Tritomochaudia 


G. implexa 


Gaudichaudia implexa fruit morphology and geographic distribution of the holotype and paratypes. A—K, 


plants with 2 ancestor from section Gaudichaudia: A) Jessup 4076 (type), B) Rzedowski 32522, C) Jessup 4079, D) 
Jessup 4080, E) Jessup 4081, F) Jessup 4082, G) Jessup 4037, H) Jessup 4018, 1) Jessup 4100, J) Anderson 12624, 
K) Jessup 4105; L—X, plants with 2 ancestor from section Tritomopterys: L) Jessup 4006, M) Jessup 4108, N) Anderson 
& Laskowski 3707, O) Jessup 4115, P) Anderson & Laskowski 4293, Q) Anderson & Laskowski 4056, R) Jessup 4002, 
S) Anderson 13316, T) Jessup 4000, U) Jessup 4001, V) Jessup 4113, W) Jessup 4114, X) Jessup 4106. Samara 


illustrations are approximately 1X. 


distinctions will be difficult to discern. I collected 
flower buds of the type, prepared aceto-carmine 
squashes of meiotic pollen mother cells and count- 
ed the chromosomes. In three separate preparations 
I found n = 80 pairs. Anderson (1993) reported 
chromosome counts in several collections included 
here as paratypes. Anderson found meiotic chro- 
mosome number was n = 80 in Rzedowski 32522, 
Anderson & Laskowski 3707, Anderson & Las- 
kowski 4056, Anderson & Laskowski 4293, Ander- 
son 12624, and Anderson 13316. 


Paratypes. Mexico. Plants with 2 ancestor from 
section Gaudichaudia: Hidalgo: 7 km NE Mezqui- 
titlan, rd to Zacualtipan, 1800 m, 17 Nov 1974, 


Rzedowski 32522 (MICH, IEB). Jalisco: Cerro Vie- 
jo, trail S of Tlajomulco, 1800 m, 5 Nov 1988, 
Jessup 4079 (CHAP, IEB, MICH, UC); 10 mi NE 
of Cocula, between Guadalajara and Autlan, 1380 
m, 6 Nov 1988, Jessup 4080 (CHAP, IEB, MICH, 
UC); 10 mi NE of Cocula, between Guadalajara 
and Autlan, 1380 m, 6 Nov 1988, Jessup 4081 
(CHAP, IEB, MICH, UC); 11.3 km NE of Tecol- 
otlan, between Guadalajara and Autlan, 1480 m, 6 
Nov 1988, Jessup 4082 (CHAP, IEB, MICH, UC). 
Michoacan: Mpio. Benito Juarez, 17.5 km S of Zi- 
tacuaro, 0.5 km S of Guanoro, 500 m, 16 Oct 1988, 
Jessup 4037 (CHAP, IEB, MICH, UC). Morelos: 1 
km SE of Laureles village limit, Barranca Tezahu- 


2002] 


ate, ca. 3 km NW of Tlayacapan, 1800 m, 6 Oct 
1988, Jessup 4018 (CHAP, IEB, MICH, UC). So- 
nora: 18 km N of Yécora—Hermosillo hwy, on rd 
to Sahuaripa, 850 m, 12 Sep 1990, Jessup 4100 
(CHAP IEB. MICH, UC)...Zacatecas: 21 km_S. of 
Villanueva on rd to Jalpa, 1900 m, 12 Sep 1983, 
Anderson 12624 (MICH): 21 km S of Villa Nueva, 
1900 m, 16 Sep 1990, Jessup 4105 (CHAP, IEB, 
MICH, UC). Plants with 2 ancestor from section 
Tritomopterys: Guanajuato: 16 km NE of San Fi- 
lipe, 1890 m, 2 Oct 1988, Jessup 4006 (CHAP, IEB, 
MICH, UC); 19 km E of San Luis de la Paz on rd 
to Victoria, 2070 m, 17 Sep 1990, Jessup 4108 
(CHAP, IEB, MICH, UC). Jalisco: 9.7 km E of Vil- 
la Corona, above Lago Atotonilco, 1420 m, 24 Sep- 
tember 1966, Anderson & Laskowski 3707 (MICH). 
Nuevo Leon: 6.1 km S of Allende, between Linares 
and Monterrey, 10 Oct 1990, Jessup 4115 (CHAP, 
IEB, MICH, UC). Oaxaca: 7.4 km NE of Chazum- 
ba, 2150 m, 23 Nov 1966, Anderson & Laskowski 
4293 (MICH). San Luis Potosi: 32.9 km W of Cd 
Valles, 485 m, 18 Oct 1966, Anderson & Laskowski 
4056 (MICH); 3.2 km W of Cuidad Valles, 420 m, 
1 Oct 1988, Jessup 4002 (CHAP, IEB, MICH, UC); 
20 km E of Santa Catarina, 1200 m, 29 Oct 1983, 
Anderson 13316 (MICH). Tamaulipas: hwy 101, 
1.6 km S of bridge over Rio San Marcos, 14.5 km 
S of ject with hwy 85, 30 Sep 1988, Jessup 4000 
(CHAP, IEB, MICH, UC); 6.4 km N of jct hwy 85 
and hwy 40, N of Guayalejo, 750 m, 1 Oct 1988, 
Jessup 4001 (CHAP, IEB, MICH, UC); NW of 
Tampico, 4.8 km E of Gonzalez, 8 Oct 1990, Jessup 
4113 (CHAP, IEB, MICH, UC); 77 km N of Ciudad 
Victoria, 32 km N Rio Purificacion, 250 m, 8 Oct 
1990, Jessup 4114 (CHAP, IEB, MICH, UC). Za- 
catecas: Mpio. Jalpa, 5.1 km E of jct hwy 54 and 
hwy 70, rd to Aguascalientes, 16 Sep 1990, Jessup 
4106 (CHAP, IEB, MICH, UC). 

Gaudichaudia implexa is morphologically more 
variable and geographically more widespread than 
any other species in Gaudichaudia. The name, 
which means “‘entangled,”’ refers not only to the 
typical habit of the plant in relation to the surround- 
ing supportive, often spiny, prickly or thorny veg- 
etation, but to the fact that lineages within this spe- 
cies represent the entangled accretion of genetically 
intertwining amphiploid lineages formed among 
diploid species in the G. albida sens. lat. and the 
G. cynanchoides sens. lat. complexes. All of the 
lineages in this species proliferate via autogamous- 
ly produced samaras, exhibit samara wing mor- 
phology with some degree of shape intergradation 
among shapes typically found in the diploids, and 
on analysis of cpDNA exhibit a chloroplast haplo- 
type typical of either the G. albida species complex 
or the G. cynanchoides species complex (Jessup 
1994, 2002). Judging from the spectrum of samara 
wing morphology, the broad range of stem and leaf 
vesture, and the combination of cpDNA haplotypes 
found it seems likely that more than one of the 
species in section Tritomopterys (Fig. 1) is active 


JESSUP: GAUDICHAUDIA 


249 


in formation of the amphiploids. Ten cpDNA hap- 
lotypes were detected in samples representing sec- 
tion Tritomopterys, and twelve of the G. implexa 
collections (paratypes) shared one of those haplo- 
types with three species in section Tritomopterys 
(G. albida, G. diandra, Gaudichaudia sp. ined. *‘in- 
termedia’’). One additional haplotype closely relat- 
ed to other haplotypes carried by species in section 
Tritomopterys (Gaudichaudia sp. ined. “‘velutina,”’ 
Gaudichaudia sp. ined. “‘intermedia,”’ and G. dian- 
dra) was detected in a single collection of G. im- 
plexa. Of the eight cpDNA haplotypes detected in 
samples representing section Gaudichaudia, eight 
of the G. implexa collections shared one of those 
haplotypes with G. cynanchoides. Three additional 
haplotypes carried by specimens representing sec- 
tion Gaudichaudia were each detected in one col- 
lection of G. implexa. 

The plants are typically found in ruderal habitats 
and range in geographic distribution (Fig. 6) from 
southern Puebla westward through the Eje Volcan- 
ico Transversal to central Jalisco, northward on the 
Altiplano Mexicano to Nuevo Leon, in scattered 
locations in the Sierra Madre Occidental, and east- 
ward into the Sierra Madre Oriental in Tamaulipas, 
San Luis Potosi, Queretaro, and Hidalgo. Anoma- 
lous collections from Chihuahua, Coahuila and Du- 
rango probably represent rare isolated lineages of 
this species, similar to the northern disjunct popu- 
lation sampled from southeastern Sonora. Though 
formed as amphiploids among lineages in section 
Tritomopterys and section Gaudichaudia, the geo- 
graphic range and apparent ecological amplitude of 
G. implexa far exceeds that of either ancestral dip- 
loid lineage. 


Nothosection Zygotomopterys S. L. Jessup notho- 
sect. nov. (Gaudichaudia sect. Tritomopterys X 
Gaudichaudia sect. Zygopterys), Fig. 7. This no- 
thosection contains two amphiploid species from 
southern Puebla and west-central Oaxaca, with 
an isolated population of one species from cen- 
tral Guerrero. 


Gaudichaudia zygoptera S. L. Jessup, sp. nov. 
(Fig. 7)—TYPE: Mexico. Oaxaca: 10.1 km N of 
hwy 190 on rd to Guelatao, 1880 m, in thicket 
near stream, 22 Oct 1988, Jessup 4042 (Holotype 
MICH; isotypes CHAP, IEB, UC). 


A Gaudichaudia galeottiana facile distinguenda 
lobis lateralibus alae samarae valde asymmetricis, 
uno vel utroque lobo majori [quam eis Gaudichau- 
dia galeottiana], lobo postico angustiori, et nulla 
constrictione lobi postici ad nucem, vel aliquot his 
differentiis. A formis Gaudichaudia implexa lobum 
posticum alae samarae latum ferentibus distinguen- 
da apicibus loborum lateralium alae samarae late 
rotundatis pro apicibus obtuse acutis Gaudichaudia 
implexa. 

Readily distinguished from G. galeottiana by a 
marked asymmetry in the lateral lobes of the sa- 


250 


Fic. 7. 


MADRONO 


[Vol. 49 


Nothosection Zygotomopterys 
© G. intermixteca 


wy SB 
e® 


a | 


3+ G. zygoptera 


Nothosection Zygotomopterys fruit morphology and geographic distribution of the holotypes and paratypes. 


A-E G. intermixteca: A) Jessup 4040, B) Anderson 13031, C) Jessup 4043, D) Jessup 4047 (type), E) Jessup 4046, 
F) Jessup 4045; g—m, G. zygoptera: G) Anderson 12990, H) Jessup 4044, 1) Jessup 4038, J) Jessup 4072, K) Jessup 
4042 (type), L) Jessup 4049, M) Anderson 13138. Samara illustrations are approximately 1x. 


mara wing, by one or both lobes larger than those 
found in G. galeottiana, by a narrower postical lobe 
of the samara wing, and by the absence of a prom- 
inent constriction in the postical lobe where it at- 
taches to the nut, or by some combination of these 
features. Distinguished from phases of G. implexa 
that bear a broadened postical lobe of the samara 
wing by the broadly rounded apices on the lateral 
lobes of the samara wing, in contrast with the gen- 
erally bluntly acute apices typical of the lateral 
lobes in G. implexa. 


Paratypes. Mexico. Guerrero: 103 km N of Ac- 
apulco on rd to Chilpancingo, 8 km along rd to El 
Alquitran, 2000 m, 30 Oct 1988, Jessup 4072 
(CHAP, IEB, MICH, UC). Oaxaca: 22.5 km S of 
Huahuapan on rd to Oaxaca, 2100 m, 20 Oct 1988, 
Jessup 4049 (CHAP, IEB, MICH, UC); Mpio. Oa- 
xaca, vicinity of Monte Alban, 21 Oct 1988, Jessup 
4044 (CHAP, IEB, MICH, UC). Puebla: .8 km NW 
of Cacaloapan on rd between Puebla and Tehuacan, 
1970 m, 19 Oct 1988, Jessup 4038 (CHAP, IEB, 
MICH, UC); 89 km S of Teotitlan on rd to Oaxaca, 
1500 m, 10 Oct 1983, Anderson 12990 (MICH); 3 
km S of Ocotlan on rd from Oaxaca to Puerto An- 
gel, 1540 m, 15 Oct 1983, Anderson 13138 
(MICH). 

Plants of ruderal habitat ranging from central and 
southern Puebla south into central Oaxaca (Fig. 7). 


The name refers to the zygomorphy evident in the 
lobes of the lateral samara wing, which indicates 
the influence of ancestral hybridization with line- 
ages from section Tritomopterys. Though the evi- 
dence is weak for the branching order of sections, 
in the maximum likelihood analysis of cpDNA re- 
striction sites section Zygopterys and section Tri- 
tomopterys form a monophyletic group with section 
Tritomopterys haplotypes forming a crown cluster, 
indicating possible paraphyly of section Zygopter- 
ys. All but one of the specimens of G. zygopterys 
sampled for cpDNA haplotypes carry haplotypes 
closely related to that carried by G. galeottiana, 
indicating that most of the hybridization contribut- 
ing to the formation of this species involves pollen 
donors from section Tritomopterys. The paratype 
Anderson 12990 has n = 80 chromosome pairs at 
meiosis (Anderson 1993). 

Only one of the cpDNA haplotypes encompassed 
in section Zygopterys is unequivocally assigned to 
G. galeottiana. The other haplotypes were found 
exclusively in G. zygoptera. The possibility re- 
mains that variation seen in samara morphology in 
G. zygoptera is indicative of relictualism, repre- 
senting a tendency toward increased zygomorphy 
in the samara wing that persists in lineages that 
were ancestral to section Tritomopterys. It seems 
more probable, however, that cpDNA haplotype di- 


2002] 


versity in section Zygopterys is indicative of ances- 
tral diversity within G. galeottiana that has been 
swamped by introgression with adventive lineages 
from section Tritomopterys. One population of G. 
zygoptera, the western disjunct in central Guerrero, 
carries a cpDNA haplotype shared with three line- 
ages (G. albida, G. diandra, Gaudichaudia sp. 
ined. “‘intermedia’’) within section Tritomopterys. 
An anomalous plant collected in southern San Luis 
Potosi (Anderson & Laskowski 4043) that is mor- 
phologically well placed in G. implexa bears a 
cpDNA haplotype from section Zygopterys. That 
plant, growing 500 km north of the range of G. 
zygoptera, is probably a hybrid between G. zyg- 
optera and either G. cynanchoides or G. implexa. 
The nuclear DNA signature, based on randomly 
amplified DNA, places it close to G. implexa, nest- 
ed close to G. cynanchoides (Jessup 1994). 


Gaudichaudia intermixteca S. L. Jessup, sp. nov. 
(Fig. 7)—TYPE: Mexico. Oaxaca: Mpio. Oaxa- 
ca, vicinity of Monte Alban, 21 October 1988, 
Jessup 4047 (Holotype MICH; isotypes CHAP, 
IEB, UC). 


A stirpibus nothosectionis Cyclotomopterys, qui- 
buscum generatim similis est forma alae samarae, 
distinguenda est lobo postico angustiori, apice lobi 
postici latiori minusque acuto-acuminata, lobis la- 
teralibus vix ad valde asymmetricis, et samaris la- 
tioribus super centrum nucis, pro latioribus ad vel 
sub centrum nucis. A Gaudichaudia synoptera dif- 
fert pilis caulis minoribus adpressisque, non paten- 
tibus erectisve, brachiis trabeculae fere aequalibus 
in longitudine et numquam subaristatis ut frequent- 
er in Gaudichaudia synoptera. Gaudichaudia sym- 
plecta similis indumento caulis, sed pilis caulis mi- 
noribus saepeque pro ratione latioribus [quam eis 
G. symplecta|. Formae Oaxacae centralis alis sa- 
marae reductis a Gaudichaudia cynanchoides se- 
cedunt pilis parvis v-formibus omnino nullis. 

Distinguished from lineages in nothosection Cy- 
clotomopterys, with which it shares a general sim- 
ilarity in shape of the samara wing, by the narrower 
postical lobe of the samara wing, by the broader, 
less acute-acuminate apex of the postical lobe, by 
the slight to pronounced asymmetry in the lateral 
lobes of the samara wing, and in having the sa- 
maras typically widest above the center of the nut, 
rather than widest at or below the center of the nut. 
Differing from G. synoptera in having the stem 
hairs smaller and appressed rather than spreading 
or erect, with the limbs of the trabecula nearly 
equal in length and never subaristate as frequently 
found in G. synoptera. Somewhat similar in stem 
vesture to G. symplecta, but with smaller stem hairs 
that are frequently wider relative to their length 
than those in G. symplecta. In phases from central 
Oaxaca with reduced samara wings, separable from 
G. cynanchoides by the complete absence of small 
v-shaped hairs. 


JESSUP: GAUDICHAUDIA 251 


Paratypes. Mexico. Oaxaca: 15 km W of Oaxaca 
on hwy 190, 1710 m, 11 Oct 1983, Anderson 13031 
(MICH); Mpio. Oaxaca, vicinity of Monte Alban, 
21 October 1988, Jessup 4043 (CHAP, IEB, MICH, 
UC); Mpio. Oaxaca, vicinity of Monte Alban, 21 
October 1988, Jessup 4045 (CHAP, IEB, MICH, 
UC); Mpio. Oaxaca, vicinity of Monte Alban, 21 
October 1988, Jessup 4046 (CHAP, IEB, MICH, 
UC); (CHAP, IEB, MICH, UC). Puebla: 8 km S and 
W of Tehuacan on rd to Huahuapan de Leon, 1900 
m, Jessup 4040 (CHAP, IEB, MICH, UC). 

Gaudichaudia intermixteca means “‘among the 
Mixtec,’ reflecting the narrow distribution of this 
species in central Oaxaca, where it is especially 
abundant and diverse in the vicinity of Monte Al- 
ban, but the name also denotes the genetically ‘‘in- 
termixed”’ nature of this species. The species has 
clear affinities with lineages in section Tritomop- 
terys, as shown by phylogenetic placement of the 
cpDNA haplotype borne by all collections exam- 
ined. That cpDNA haplotype is, however, unique to 
G. intermixteca, suggesting an ancient hybridiza- 
tion that has persisted long enough to evolve a di- 
vergent genome. Nuclear DNA analysis places the 
group close to G. galeottiana and the G. albida 
complex (Jessup 2002). In addition to the concen- 
tration in the vicinity of Monte Alban, the species 
is found somewhat further north in Oaxaca and in 
extreme southern Puebla (Fig. 7). Anderson (1993) 
found n = 80 pairs of meiotic chromosomes in the 
paratype Anderson 13031. 


Nothosection Cyclotomopterys S. L. Jessup notho- 
sect. nov. (Gaudichaudia sect. Cyclopterys X 
Gaudichaudia sect. Tritomopterys), Fig. 8. This 
nothosection includes two new species with re- 
stricted geographic ranges in the central and 
western Eje Volcanico Transversal. 


Gaudichaudia synoptera S. L. Jessup, sp. nov. 
(Fig. 8)—TYPE: Mexico. Edo. Mexico, Mpio. 
Tepetilixpa, 3.2 km S of Tepetilixpa on rd be- 
tween Cuautla and Amecameca, 2160 m, 8 Oct 
1988, Jessup 4020 (Holotype MICH; isotypes 
CHAP. IEB} UC). 


A Gaudichaudia symplecta et Gaudichaudia in- 
termixteca secedit pilis caulis longis angustisque, 
erecto-patentibus ad subaristatis, numquam adpres- 
sis, caulibus hispidis, brachiis trabeculae inter se 
valde differentibus in longitudine. A G. andersonii 
differt samaris ovatis ad orbiculatis sine constric- 
tione postica alae lateralis prope basin nucis. 

Separable from G. symplecta and G. intermixteca 
by the relatively long and narrow stem hairs char- 
acterized by a marked difference in length of the 
trabecula limbs, the stem hairs erect-spreading to 
subaristate, never appressed, and the stems hispid. 
Distinct from G. andersonii in the ovate to orbic- 
ular samaras lacking a postical constriction in the 
lateral wing near the base of the nut. 


D5, 


Fic. 8. 


MADRONO 


[Vol. 49 


Nothosection Cyclotomopterys 


2 


3 G.synoptera 
. MS’ 
D 


Nothosection Cyclotomopterys fruit morphology and geographic distribution of the holotypes and paratypes. 


A-E, G. synoptera: A) Jessup 4022, B) Jessup 4023, C) Soto 4024, D) Jessup 4020 (type), E) Jessup 4015; F—H, G. 
symplecta: F) Jessup 4074 (type), G) Anderson 13291, H) Jessup 4025; 1, G. andersonii: Jessup 4026 (type). Samara 


illustrations are approximately 1X. 


Paratypes. Mexico. Guerrero: Soto 4024 
(MICH). Estado Mexico: 7 km W of Temascaltepec 
on rd to Real de Arriba, 2000 m, 10 Oct 1988, 
Jessup 4022 (CHAP, IEB, MICH, UC); 12 km SW 
of Temascaltepec on rd to Tejupilco, 1750 m, 11 
Oct 1988, Jessup 4023 (CHAP, IEB, MICH, UC). 
Morelos: ca. | km SE of Laureles village limit, Bar- 
ranca Tezahuate, ca 3 km NW of Tlayacapan, 1800 
m, 6 Oct 1988, Jessup 4015 (CHAP, IEB, MICH, 
UC). 


G. cycloptera 


= 


Gaudichaudia synoptera is one of two recogniz- 
able hybrid species formed between species in sec- 
tion Jritomopterys and section Cyclopterys. It is 
immediately separable from the other species in this 
nothosection by the distinct morphology of stem 
hairs (Fig. 9). The most probable origin for G. syn- 
optera is as an amphiploid between Gaudichaudia 
sp. ined. “hirsuta” and G. cycloptera. The evi- 
dence from morphology and DNA analyses sup- 
ports that conclusion. The stem vesture is similar 


1.0mm C i D 


Comparison of stem hairs on representative collections from Michoacan. A) G. cycloptera, Jessup 4033; B) 
G. synoptera, Jessup 4020 (type); C) G. andersonii, Jessup 4026 (type); D) G. symplecta, Jessup 4074 (type); E) G. 
diandra, Jessup 4034. 


Fic. 9. 


2002] 
to that found in Gaudichaudia sp. ined. “‘hirsuta,”’ 
and some of the collections tested share a cpDNA 
haplotype with members of that lineage. The type 
bears a cpDNA haplotype that, outside of the G. 
synoptera collections, is restricted to Oaxaca and 
Chiapas where it is most typically carried by Gau- 
dichaudia sp. ined. “hirsuta” and Gaudichaudia 
sp. ined. “‘velutina”’ (Fig. 1). 

Nuclear DNA analyses place the type closest to 
Gaudichaudia sp. ined. “‘hirsuta.”> Other members 
of the species carry a cpDNA haplotype that is 
most commonly carried by G. cycloptera. The 
name means “‘twining together,” reflecting the ten- 
dency of several twining branches, even from dif- 
ferent plants, and occasionally from different spe- 
cies, to form “‘limbs”’ by twining together. The ob- 
vious analogy is that the lineages themselves are 
intertwined in this species that apparently originat- 
ed through hybridization of plants from different 
sections of Gaudichaudia. The literal meaning of 
the name and the metaphorical meaning coincide in 
this plant to the extent that the habit of forming 
twined limbs complexed from shoots of different 
genets facilitates wide outcrossing and a reticulated 
ancestry. The pollinators are anthophorine bees 
specialized for collecting oil from the calyx glands 
present as part of the conservative malpighiacean 
floral syndrome (Vogel 1974; Anderson 1990), and 
are adapted for the family-level floral characters. 
The bees likely do not distinguish among species 
within a genus, and are thus more apt to cross pol- 
linate divergent lineages when the flowers are 
closely juxtaposed, as they often are in the tangled 
thickets inhabited by Gaudichaudia. Gaudichaudia 
synoptera has a rather narrow geographic distribu- 
tion (Fig. 8), ranging from southwestern Edo. Mex- 
ico, where it is especially common in the vicinity 
of Temascaltepec, eastward into Morelos, and in 
Edo. Mexico east of Distrit o Federal. 


Gaudichaudia symplecta S. L. Jessup, sp. nov. 
(Fig. 8) TYPE: Mexico. Michoacan, Mpio. Mo- 
relia, 23.5 km E of Morelia on rd from Cd Hi- 
dalgo, 2170 m, 1 Nov 1988, Jessup 4074 (Ho- 
lotype MICH; isotypes CHAP, IEB, UC). 


A Gaudichaudia cycloptera secedit incisura ap- 
icali insigni alae lateralis et apice acuto-acuminata 
marginis posticae loborum lateralium alae. A Gau- 
dichaudia synoptera et Gaudichaudia andersonii 
differt indumento caulis, pilis adpressis, numquam 
erecto-patentibus vel subaristatis, caulibus ita seri- 
ceis, non hispidis, brachiis trabeculae subaequali- 
bus in longitudine. A Gaudichaudia intermixteca 
differt lobis alae lateralis samarae plerumque sym- 
metricis et samaris latioribus ad vel sub medium. 

Separable from G. cycloptera by the pronounced 
apical notch in the lateral wing, and the acute-acu- 
minate apex of the postical margin of the lateral 
wing lobes. Distinct from G. synoptera and G. an- 
dersonii in the stem vesture, comprising hairs that 


JESSUP: GAUDICHAUDIA 29 


1oe) 


are subequal in length of the trabecula limbs, uni- 
formly appressed, never erect-spreading or subar- 
istate, the stems thus sericeous, not hispid. Differ- 
ing from G. intermixteca in the generally symmet- 
ric lobes of the lateral samara wing, and in having 
the samaras widest at or below the middle. 


Paratypes. Mexico. Estado Mexico: 0.5 km N of 
Amatepec on rd to Tejupilco, 1750 m, 11 Oct 1988, 
Jessup 4025 (CHAP, IEB, MICH, UC). Michoacan: 
Mpio. Zitacuaro, Puerto del Gato, 5 km N of Zi- 
tacuaro on hwy 15, 1800 m, 26 Oct 1983, Anderson 
13291 (MICH). 

The name means twisted and plaited together, re- 
ferring to the habit, as in the previous species, of 
forming branches that appear almost braided from 
separate twining branches. Both the type and one 
of the paratypes (Anderson 1329]) bear a cpDNA 
haplotype that is characteristic of section Tritomop- 
terys. The other paratype (Jessup 4025) bears a 
cpDNA haplotype that is otherwise confined to lin- 
eages within section Cyclopterys. Unlike its close 
relative, G. synoptera, this species has stem tri- 
chomes that are in all respects similar to those typ- 
ically found in G. albida sen. str. Nuclear DNA 
analyses also place members of this species close 
to lineages within section Cyclopterys. The conclu- 
sion that G. symplecta originated as an amphiploid 
cross between G. albida and G. cycloptera is thus 
well supported. 

Among the plethora of specific and infraspecific 
taxa treated by Niedenzu, none is a feasible can- 
didate for assignment of this species. Niedenzu 
moved G. arnottiana Juss. to subspecific rank un- 
der G. pentandra Juss. (=G. cycloptera (DC.) W. 
R. Anderson) and described three new varieties and 
two new forms of G. pentandra subsp. arnottiana 
(Juss.) Niedenzu. One or more of those infraspecific 
names might be construed as referring to this spe- 
cies, but the characters Niedenzu emphasized in his 
diagnoses make it difficult to reach a conclusion 
about circumscription of his taxa. Jussieu (1843) 
clearly expressed doubt that his G. arnottiana is 
distinct from G. cycloptera: ‘“‘admodum. affinis 
precedenti, cum qua staminibus quinque antheri- 
feris equalibus couvenit foliorumque forma (fide 
iconis); calyce, ut videtur, 10-glanduloso ut quibus- 
dam levioris momenti notis subdissimilis; forsan 
conjungenda. Species Candolleana forsan conspe- 
cifica, certe congener fide iconis Flor. Mexic. ined. 
in qua carpellum calyce eglanduloso stipatum pen- 
dere e filo videtur.’’ The “‘closely related preceding 
species” is G. cycloptera (DC.) W. Anderson, and 
nothing in the protologue clearly differentiates G. 
arnottiana from G. cycloptera or indicates charac- 
ters that would suggest the name should apply to 
the species that I am naming G. symplecta. Jussieu 
in fact expressed doubt that G. arnottiana is distinct 
and suggested that they are conspecific. Niedenzu’s 
infraspecific elaborations notwithstanding, G. ar- 


254 


nottiana Juss. can be considered a synonym of G. 
cycloptera (DC.) W. R. Anderson. 


Gaudichaudia (sect. Cyclopterys) andersonii S. L. 
Jessup, sp. nov. (Fig. 8)—TYPE: Mexico. Estado 
Mexico: 1 km S of Temascaltepec on rd to Te- 
jupilco, 1790 m, 13 Oct 1988, Jessup 4026 (Ho- 
lotype MICH). 


A speciebus alis differt samaris magnis incisura 
apicali insigni alae lateralis, lobos duos laterales 
symmetricos sinu lata v-forma separatos formanti- 
bus, marginibus alae lateralis rotundatis et ad con- 
strictionem sub centrum contractis, lobo postico ex- 
panso e constrictione sub nucem et interdum cauda 
brevi abrupta ornato. 

Distinct from other species in the large samaras 
with pronounced apical notch in the lateral wing, 
forming two symmetric lateral lobes separated by 
a broad v-shaped sinus, the lateral wing margins 
rounded to a constriction in the wing below center, 
the postical lobe flared from the constriction below 
the nut and sometimes abruptly appendaged by a 
short tail. 


Paratype. Mexico. Estado Mexico: 1.6 km S of 
Temascaltepec on Temascaltepec-Tejupilco-Amate- 
pec rd, 1600 m, 14 Oct 1966, Anderson & Las- 
kowski 3988 (MICH). 

This very distinctive species is named in honor 
of William R. Anderson who has devoted years of 
field and laboratory work to the study of Gaudi- 
chaudia, among other malpighs. I found the species 
during a prolonged foray in the tangled vegetation 
south of Temascaltepec, occasioned by mechanical 
failure of the VW microbus, belonging to S. D. 
Koch, in which Anderson and I were passengers 
during an expedition into northern Guerrero. An- 
derson had collected the species in 1966 in the 
same general area (Fig. 8). Nothing like it has been 
found anywhere else, and other than the type and 
paratype cited here, I am unaware of other collec- 
tions of this species. 

In vegetative characters and floral morphology 
G. andersonii resembles both G. cycloptera and G. 
synoptera. The long, rather narrow stem hairs with 
distinctly unequal limbs of the trabecula (Fig. 9) 
are a fairly close match to those found in G. cy- 
cloptera, but are somewhat smaller than those 
found in G. synoptera, and although the hairs are 
distinctly erect-spreading in G. andersonii, they do 
not form the distinctive subaristate stem vesture 
found in G. synoptera. The type of G. andersonii 
was found to carry a cpDNA haplotype otherwise 
known only from a topotype collection of G. chas- 
ei. In the nuclear DNA analysis G. andersonii is 
closely placed with G. cycloptera, well apart from 
G. chasei, which is placed closer to G. intermixteca 
and G. galeottiana in that analysis (Jessup 1994). 
The molecular data suggests that G. andersonii is 
a hybrid within section Cyclopterys between G. cy- 
cloptera and G. chasei, and the intermediacy of sa- 


MADRONO 


[Vol. 49 


mara wing morphology is consistent with that con- 
clusion. 


DISCUSSION 


Tribe Gaudichaudieae, as most recently defined 
(Anderson 1985) includes Gaudichaudia, Aspicar- 
pa, Janusia, Camarea, and Peregrina. According 
to Davis et al. (2001), the tribe is well supported 
as a monophyletic group nested within a well sup- 
ported monophyletic stigmaphylloid clade, which 
includes (in addition to Tribe Gaudichaudieae) Stig- 
maphyllon and Banisteriopsis, among other genera. 
A lineage resembling one of the stigmaphylloids is 
the most plausible ancestor for Tribe Gaudichau- 
dieae. Examining the distribution of chromosome 
numbers (Anderson 1993) across the entire stig- 
maphylloid clade we see in species representing 
basal lineages a high frequency of nm = 10 meiotic 
chromosome pairs, probably the base number for 
the clade. In Banisteriopsis 13 of 14 species re- 
ported have n = 10 pairs, and one has n = 20 pairs. 
In Stigmaphyllon 4 of 4 species reported have n = 
10 pairs. Within Tribe Gaudichaudieae, ploidal lev- 
els in Janusia are n = 10 (2 of 12 species reported), 
n = 20 or, in one case, aneuploid n = 19 (9 of 12 
species reported), and n = 40 (1 of 12 species re- 
ported). Chromosome numbers in Aspicarpa are n 
= 20 (1 of 6 species reported) and n = 40 (5 of 6 
species reported). Camarea all have n = 17, and 
Peregrina has n = 19 meiotic pairs. In Gaudichau- 
dia chromosome number is n = 40 or n = 80 (and 
n = 120 in one isolated collection). 

The chromosome numbers in combination with 
the molecular data strongly support the hypothesis 
that evolution in the stigmaphylloids has proceeded 
by a series of genomic doublings trending from n 
= 10 to n = 80, producing a polyploid series on 
the base number x = 10. In the stigmaphylloid 
clade, Banisteriopsis and Stigmaphyllon are ances- 
tral diploids, with a single tetraploid in Banister- 
iopsis. Janusia is primarily tetraploid with two dip- 
loid lineages remaining and a single octoploid lin- 
eage. Camarea and Peregrina are most likely an- 
ueploid reductions from the tetraploid state. 
Aspicarpa 1s now understood to be at least in part 
nested within Gaudichaudia and retains a single 
known tetraploid lineage, but is primarily octaploid. 
Gaudichaudia sensu stricto is fundamentally octo- 
ploid, but developed a series of wide crosses at the 
sextodecaploid level, including the species de- 
scribed in this paper. 

The genus Gaudichaudia is apparently built on 
an ancient polyploid complex. Regular meiotic 
pairing among homologous chromosomes resulted 
in essentially instantaneous diploidization of am- 
phiploid crosses. Since the trend toward higher 
ploidy is clearly demonstrated within the stigma- 
phylloid clade, the obvious hypothesis is that line- 
ages with lower ploidy will be phylogenetically 
more basal within Tribe Gaudichaudieae. If that hy- 


2002] 


pothesis is supported by molecular phylogenetic 
tests, I predict that relict lineages will be found to 
exist within Gaudichaudia, perhaps as narrow en- 
demics or isolated populations in central Mexico, 
that retain the primitive lower ploidal levels. The 
occurrence of such lineages is already a strong pos- 
sibility within section Oligopterys, since a single 
species of Aspicarpa (A. schinnii W. R. Anderson) 
is known to retain the n = 20 condition (Anderson 
1993). That species may or may not properly be- 
long within Gaudichaudia, and it remains to be 
seen whether molecular data will place that lineage 
as basal within the section. A thorough screening 
of isolated lineages within other sections of Gau- 
dichaudia is likely to reveal additional instances if 
indeed any are extant. 


ACKNOWLEDGMENTS 


The author thanks William R. Anderson, who gener- 
ously shared his knowledge and research collections and 
provided guidance and assistance with field work, and E. 
Pichersky and R. Fogel, who provided laboratory space 
and shared equipment. Stephen Koch at CHAPA provided 
hospitality and generous assistance with fieldwork. Many 
thanks to reviewers who provided helpful comments on 
the manuscript. This research was funded in part by NSF 
grant BSR-8700340 to W. R. Anderson, and by NSF Doc- 
toral Dissertation Improvement grant BSR-8823076 to W. 
R. Anderson for S. L. Jessup. 


LITERATURE CITED 


ANDERSON, W. R. 1979. Floral conservatism in neotropical 
Malpighiaceae. Biotropica 11:219—223. 

. 1980. Cryptic self-fertilization in the Malpighi- 

aceae. Science 207:892-893. 

. 1985. Peregrina, a new genus of Malpighiaceae 

from Brazil and Paraguay. Systematic Botany 10: 

303-307. 

. 1987. Notes on neotropical Malpighiaceae—II. 

Contributions from the University of Michigan Her- 

barium 16:55—108. 

. 1993. Chromosome numbers of neotropical Mal- 
pighiaceae. Contributions from the University of 
Michigan Herbarium 19:341—354. 

BUCHMANN, S. L. 1987. The ecology of oil flowers and 


JESSUP: GAUDICHAUDIA 


259 


their bees. Annual Review of Ecology and System- 
atics 18:343-—369. 

CAMERON, K. M., M. W. CHASE, W. R. ANDERSON, AND H. 
G. HILuis. 2001. Molecular systematics of Malpighi- 
aceae: evidence from plastid rbcL and matK sequenc- 
es. American Journal of Botany 88:1847—1862. 

CANDOLLE, A. P. DE. 1824. Malpighiaceae. Prodromus Sys- 
tematis Natualis Regni Vegetabilis [:577—592. 

Davis, C. C., W. R. ANDERSON, AND M. J. DONOGHUE. 
2001. Phylogeny of Malpighiaceae: evidence from 
chloroplast ndhF and trnL-F nucleotide sequences. 
American Journal of Botany 88:1830—1846. 

Dow LInG, T. E., C. Moritz, J. D. PALMER, AND L. H. RIE- 
SEBERG. 1996. Nucleic acids III: analysis of fragments 
and restriction sites. Pp. 249-320 in D. M. Hillis, C. 
Moritz, and B. K. Mable (eds.), Molecular system- 
atics, 2nd ed. Sinauer, Sunderland, MA. 

Farris, J. S. 1989. Hennig86. Port Jefferson Station, New 
York, NY. 

GREUTER, W., J. MCNEILL, EF R. BARRIE, H.-M. BURDET, 
V. DEMOuLIN, D. S. FILGUEIRAS, D. H. NICHOLSON, P. 
C. Si_vA, J. E. Skoc, T. TREHANE, N. J. TULAND, AND 
D. L. HAwKSworTH (eds.). 2000. International Code 
of Botanical Nomenclature (St. Louis Code). Regnum 
Vegetabile 131. Koeltz Scientific Books, K6nigstein, 
Germany. 

Jessup, S. L. 1994. Reticulate evolution in Gaudichaudia 
(Malpighiaceae). Ph.D. dissertation. University of 
Michigan, Ann Arbor, MI. 

. 2002. Reticulate ancestry in Mexican Gaudi- 
chaudia (Malpighiaceae) analyzed with RAPDs and 
southern hybridization. Madrono 49:256—273. 

Jussieu, ADR. 1843. Monographie de la famille des Mal- 
pighiacées. Archives du Museum d’ Histoire Naturel- 
le. Paris 3:5—151, 255-616, pl. 1—23. 

McVauGu, R. 1980. Botanical results of the Sessé and 
Mocino expedition (1787—1803) II. the Icones Florae 
Mexicanae. Contributions from the University of 
Michigan Herbarium 14:99—140. 

NIEDENZU, F. 1928. Malpighiaceae. Jn A. Engler (ed.), Das 
Pflanzenreich IV, 141:1—870. 

SWOFFORD, D. L. 1990. PAUP: Phylogenetic analysis us- 
ing parsimony, version 3.0. Illinois Natural History 
Survey, Champaign, IL. 

TAYLOR, D. W. AND W. L. CREPET. 1987. Fossil floral ev- 
idence of Malpighiaceae and an early plant-pollinator 
relationship. American Journal of Botany 74:274- 
286. 

VoGEL, S. 1990. History of the Malpighiaceae in the light 
of pollination ecolgy. Memoirs of the New York Bo- 
tanical Garden 55:130—142. 


MApRONO, Vol. 49, No. 4, pp. 256-273, 2002 


RETICULATE ANCESTRY IN MEXICAN GAUDICHAUDIA 
(MALPIGHIACEAE) ANALYZED WITH RAPD’s AND 
SOUTHERN HYBRIDIZATION 


STEVEN L. JESSUP 
Department of Biology, Southern Oregon University, Ashland, OR 97520 
Jessup @sou.edu 


ABSTRACT 


Evidence of relationships based on randomly amplified polymorphic DNA (RAPD) data combined with 
information about cpDNA haplotypes can be used to resolve details of reticulate ancestry in an otherwise 
intractable polyploid complex in Gaudichaudia (Malpighiaceae). Robust inference of genetic relationships 
among taxa, however, depends critically on two assumptions: (1) character states compared among taxa 
are homologous, and (2) characters scored as different are independent. Application of randomly amplified 
DNA methods, such as RAPD’s, have largely made these assumptions without testing them. In this study 
I use RAPD’s to elucidate relationships among lineages and to infer reticulate ancestry of amphiploid 
lineages in Gaudichaudia. | test the assumption that comigrating RAPD fragments are homologous using 
hybridization of radio-labeled RAPD fragments probed against blots of randomly amplified DNA as an 
indicator of homology. The probes bind strongly only to fragments on the blots having sequence homol- 
ogy. Results demonstrate that all gel fragments included in the analysis meet the assumption of homology. 
Gel fragments can therefore be reliably scored directly as characters. The assumption of independence of 
RAPD fragments is also explored. Although multiple fragments with strong sequence homology appear 
in most blots, gel-visualized fragments are generally independent. 


RESUMEN 


La evidencia de relaciones basadas en datos de DNA polimorfico aleatoriamente amplificado (RAPD) 
combinados con informacion sobre haplotipos cpDNA se puede utilizar para resolver detalles, de otra 
manera insuperables, de ascendencia reticulada en un complejo poliploide en Gaudichaudia (Malpighi- 
aceae). La solida inferencia de relaciones genéticas entre especies, sin embargo, depende criticamente de 
dos supuestos: (1) los estados del caracter comparados entre grupos taxonomicos son homdlogos, y (2) 
los caracteres anotados como diferentes son independientes. El uso de los métodos de DNA amplificado 
aleatoriamente, tales como los RAPD’s, ha hecho estas asunciones en gran parte sin probarlas. En este 
estudio utilizo RAPD’s para aclarar relaciones entre linajes y para deducir la ascendencia del reticulado 
de linajes de anfiploides en Gaudichaudia. Pongo a prueba la asuncion de que los fragmentos comigrantes 
de RAPD son homodlogos usando la hibridaci6n de fragmentos RAPD marcados radiactivamente como 
testigos contra manchas de DNA aleatoriamente amplificado como un indicador de la homologia. Los 
testigos se adhieren fuertemente solamente a aquellos fragmentos dentro de las manchas que tienen 
homologia de secuencia. Los resultados demuestran que todos los fragmentos del gel incluidos en el 
andlisis cumplen con la asuncion de la homologia. Los fragmentos de gel se pueden por lo tanto contar 
confiablemente directamente como caracteres. También se explora la asunci6n de independencia de los 
fragmentos de RAPD. Aunque los fragmentos multiples con fuerte homologia de secuencia aparecen en 
la mayoria de las manchas, los fragmentos gel-visualizados son generalmente independientes. 


Key Words: Polyploid, introgression, Malpighiaceae, RAPD, Southern hybridization. 


Gaudichaudia (Malpighiaceae) has been revised chromosome numbers, showed combinations of 


based in part on inference from patterns of variation 
in cpDNA restriction sites (Jessup 2002). Evidence 
of relationships among lineages within Gaudichau- 
dia from morphology, chromosome counts, and 
amplification of RAPD fragments corroborates the 
cpDNA evidence and helps to further resolve retic- 
ulate ancestry of some species in the genus (Jessup, 
1994). In particular, a minimum spanning tree anal- 
ysis of RAPD fragments (based on presence of am- 
plified fragments visible on agarose gels), con- 
structed using Jaccard similarity, shows broad con- 
gruence of cpDNA haplotypes and nuclear ge- 
nomes. Plants that were inferred to be intersectional 
amphiploids, based on morphological features and 


cpDNA haplotypes and nuclear genomes character- 
ized by RAPD profiles that supported the hybridity 
hypotheses. 

While the RAPD data helps define lineages and 
clearly demonstrates patterns of relationship and 
ancestry among the lineages, a test of the validity 
of the underlying assumptions about the RAPD data 
nevertheless remains worthwhile (Arnold and 
Emms 1998; Rieseberg 1996; Wolfe and Liston 
1998), especially since gel fragments alone might 
be useful for inferring relationships. Homology of 
character states and independence of characters are 
prerequisite features of characters used in many 
systematic analyses. Restriction fragments are 


2002] 


strongly correlated, which is why restriction frag- 
ments alone cannot be scored for presence/absence 
and used directly as characters. While randomly 
amplified DNA markers, such as RAPD’s, have not 
been shown to be correlated in the same way that 
restriction fragments are, estimates of relationship 
based on RAPD fragments would be similarly 
skewed if tightly linked markers were treated as 
independent estimators of relatedness (Lynch 
1988). Estimates of relatedness would also be in- 
accurate if fragments of the same electrophoretic 
mobility were commonly of heterologous origins. 
In this paper I present the results of the RAPD anal- 
ysis in Gaudichaudia and report experiments that 
test RAPD band homology and independence using 
hybridization of *?P labeled RAPD probes to south- 
ern blots of RAPD gels. I demonstrate the utility 
of the procedure in identifying the specific ancestry 
of amphiploids. When combined with knowledge 
of cpDNA haplotypes in the samples, this proce- 
dure further resolves reticulate ancestry by speci- 
fying which of the parental lineages contributed the 
maternal genome. 


Expected behavior of RAPD markers in amphi- 
ploids. In tetraploid Gaudichaudia where chromo- 
somes from divergent lineages may reside within 
the same nucleus, and where the lineages are re- 
producing primarily by selfing (Anderson, 1980), 
markers that occur within the same set of chro- 
mosomes can be tightly linked. When homeologous 
chromosome sets are divergent but still pair at mei- 
osis in wide amphiploid crosses, we expect a 
marked reduction in viable gametes and reduced 
fecundity in the F, progeny. Given that strongly in- 
breeding lineages quickly approach fixation of var- 
iable alleles (Li 1976), it seems likely that a pre- 
ponderance of markers will frequently be fixed 
within a set of chromosomes in strongly selfing lin- 
eages. With little or no recombination between 
homeologous chromosomes most variability occurs 
among rather than within homeologous sets of 
chromosomes. Lineages with size variants of the 
same marker fixed on different (homeologous) sets 
of chromosomes—synologous loci as defined by 
Mindell & Meyer (2001)—will exhibit fixed het- 
erozygosity. The variants that occurred in the an- 
cestral diploids as orthologous loci, as fixed differ- 
ences at the same locus in different lineages, are 
combined in the amphiploid as synologous loci, 
though they are less likely to be fixed in progeny 
resulting from outcrossing. Since RAPD sites are 
restricted to individual chromosomes, they neces- 
sarily obey all of the constraints associated with 
chromosomal inheritance. 

We expect that RAPD fragments of the same mo- 
lecular weight—fragments that appear at precisely 
the same location on a gel—are products of ho- 
mologous sites. There is, however, the possibility 
of heterologous fragments with the same mobili- 
ty—tfragments that are identical in state but not ho- 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 


257 


mologous. We also expect that fragments of differ- 
ent mobility are independent products of unlinked 
sites. The assumptions of homology and indepen- 
dence, then, can be reduced to two questions. (1) 
Is there sequence homology among bands of dif- 
ferent electrophoretic mobility within a sample? If 
we count multiple bands as distinct when they are 
in fact the result of a single locus then we violate 
the assumption of independence. (2) Is there se- 
quence homology among bands of the same elec- 
trophoretic mobility among samples? If we count 
bands of the same electrophoretic mobility as iden- 
tical when they are in fact distinct then we violate 
the assumption of homology. Testing those assump- 
tions is simple in principle. An estimate of se- 
quence homology can be obtained by observing the 
relative strength of hybridization signal between a 
labeled RAPD probe of known origin and samples 
from bands of the same molecular weight whose 
homology is in question. Bands that hybridize 
strongly to the probe are inferred to have a high 
level of sequence homology with the probe. By in- 
ference, a strong hybridization signal is an indicator 
of close relatedness. 


MATERIALS AND METHODS 


DNA extraction. Plants in the genus Gaudichau- 
dia were sampled from a wide geographic area in 
Mexico, representing the center of diversity in the 
genus. Detailed source data for the collections is 
reported elsewhere (Jessup, 1994). Total genomic 
DNA was extracted from leaves that were snap- 
frozen in liquid nitrogen and subsequently stored at 
—80°C. Total genomic DNA extraction was based 
on the CTAB procedure presented by Hillis et al. 
(1990) and Dowling et al. (1996). All DNA extrac- 
tions were purified by ultracentrifugation on cesium 
chloride gradients, dialysis to remove the cesium 
chloride, and ethanol precipitation following pro- 
cedure in Sambrook et al. (1989). Final yield of 
DNA was between 25 and 750 pg DNA per ex- 
traction (from 1—2 grams of leaf tissue). In all, 134 
different DNAs were successfully prepared for use 
in the molecular procedures. DNA concentrations 
and estimates of DNA purity were calculated from 
optical density measurements at \ = 260, A = 280, 
and A = 320 nm, on a Beckman DU-64 UV spec- 
trophotometer. 


RAPD reactions and Southern hybridization. Pu- 
rified template DNA was diluted to 1 ng/ml in a 
reaction elixir buffered to pH 8.3 with 10 mM Tris- 
HCl containing 50 mM KCl, 2 mM MgCl, dNTPs 
at a concentration of 100 M each, five arbitrary 
10-base oligonucleotide primers (Operon®) were 
used in separate reactions, each at a concentration 
of 0.2 ~M, and Taq DNA polymerase (Perkin E]- 
mer®) was used in all reactions at a concentration 
of 0.07 units/ng template DNA. The reaction was 
carried out in a total volume of 25 wl on an 
M.J.Research® thermal cycler programmed at max- 


258 


imum ramp speed for: 1 cycle of 30 seconds at 
94°C; 45 cycles of 1 minute at 94°C, 1 minute at 
35°C, and 2 minutes at 72°C; and 1 cycle of 5 min- 
utes at 72°C, followed by a hold at 4°C. 

The products of the PCR reactions were separat- 
ed by electrophoresis on 2% agarose gels with 200 
ng/ml ethidium bromide, at 25V constant voltage 
for approximately 14 hours at room temperature in 
1X TAE, pH 7.6. A 100-bp ladder (Pharmacia Bio- 
tech®) was loaded into three or four lanes on each 
gel at even intervals among samples as a high res- 
olution molecular weight marker. Each gel run con- 
tained from 120 to 134 sample lanes and 12—16 
molecular weight marker lanes over four gels. 
RAPD fragments were visualized by UV transillu- 
mination, then photographed on Polaroid-57 at f8, 
3’ 20”. Negatives were washed in NaSO, 18% for 
1 minute, then rinsed 30 minutes in H,O. RAPD 
fragments visible on the negatives were scored by 
careful measurement on a lightbox, and molecular 
weights were determined with the gel analysis pro- 
gram, Gel Match®® (UVP). All visually detectable 
fragments on either the films or the scanned images 
were scored as present. Computer image enhance- 
ment was used to intensify faint bands. Lanes not 
showing a fragment at the same position on the gel 
were scored as absent. 

RAPD fragments from the PCR reactions were 
collected from ethidium bromide stained gels under 
UV transillumination. Bands selected for **P label- 
ing were sampled by inserting the tip of a pasteur 
pipet into the center of the band and applying light 
vacuum pressure with a pipet pump as the tip was 
withdrawn from the gel. This produced a cylindri- 
cal gel section about 1 mm wide by about 7 mm 
long while preserving the source gel for blotting. 
The gel section was extruded into a microfuge tube 
and kept on ice. Collected fragments were diluted 
10:1, reamplified using the original PCR protocol, 
then electrophoresed on 3% low Ty, agarose gels to 
further purify the fragment. 

The whole reamplified fragment was collected 
after gel purification and 12 pl was radio-labeled 
with a-**P tagged ATP using a polymerase reaction. 
A few ng of molecular weight marker were also 
labeled at the same time as the RAPD probes. La- 
beling reactions were carried out using random 
priming with a mixture of hexadeoxyribonucleo- 
tides (6 bp oligonucleotides) according to the pro- 
tocol developed by Feinberg and Vogelstein (1983, 
1984). This technique resulted in probes labeled to 
high specific activity. Unincorporated nucleotides 
were separated from labeled RAPD fragments us- 
ing sephadex columns set up in 9” glass pasteur 
capillary pipettes. Purified labeled probe was de- 
natured by immersion in boiling water for 10 min- 
utes, then ‘quenched’ on ice for 3—5 minutes before 
beginning the membrane hybridization reaction. 

RAPD gels were blotted to nylon (Zetabind®) 
membranes following procedures in Maniatis et al. 
(1989). Membranes were allowed to dry after trans- 


MADRONO 


[Vol. 49 


fer of the amplified fragments then stored at room 
temperature until Southern hybridization. 

Prehybridization and Southern hybridization re- 
actions followed procedure outlined in Dowling et 
al. (1996). The prehybridization solution was 4X 
SSC, 1% SDS, and 0.5% nonfat dry milk. Blots 
which had not been previously probed were first 
pretreated by washing in 0.1X SSC, 0.5% SDS for 
1 hour at 65°C. Prehybridized blots were removed 
from the incubator and all but about 15 ml of the 
prehybridization solution was removed from the 
hybridization tray. The labeled probe was then add- 
ed to the tray and thoroughly mixed. Several blots 
were hybridized simultaneously. Care was taken 
not to introduce bubbles into the space between 
blots. Hybridization reactions were allowed to pro- 
ceed for 12—18 hours at 62°C. 

Hybridized blots were removed from the trays 
and washed in three or four changes of 2* SSC, 
0.5% SDS: two short washes at room temperature 
followed by one or two 30—45 minute washes at 
60°C. Blots and discarded wash were monitored 
with a Geiger counter during the wash procedure 
to assess when background radiation on the filters 
had been adequately reduced. Filters were removed 
from the final wash and blotted to remove excess 
wash, then wrapped in plastic and placed in x-ray 
film cassettes with intensifying screens. Kodak X- 
OMAT® AR film was loaded into the cassettes and 
they were exposed over night at —85°C. Films were 
removed and developed on an X-OMAT® auto- 
matic X-ray film developer. Following autoradiog- 
raphy, blots were stripped of probe in hot 0.5 SSC 
and monitored until radiation was reduced to low 
levels. Blots were then prepared for reprobing with 
a different RAPD fragment. 


RESULTS 


Relationships inferred from RAPD fragments. 
Seventy-five plants were scored for 79 RAPD sites. 
Table 1 records the molecular weights of RAPD 
fragments scored from the gels for each of the 
primers used in the study. Table 2 records the frag- 
ments scored from the gels for each collection. Us- 
able sites were those that could be consistently 
scored for all 75 collections included in the analy- 
sis. Figure 1 shows a typical RAPD gel, in this case 
with RAPD bands produced using primer sequence 
CAAACGTCGG (A-19). Collections are arrayed 
on the gels by taxon defined on morphological sim- 
ilarity, and within taxon by geographic region. 
There are four marker lanes per gel, each a 100 bp 
ladder with molecular weight indicated for the 800 
bp fragments. Blank lanes are collections that did 
not amplify. Blank lanes are excluded from the data 
tables. Notice the general pattern of shared bands. 
Bands of the same molecular weight tend to occur 
in adjacent lanes on the gel, i.e., among collections 
grouped by taxon and geographic proximity. 

The data were clustered with a minimum spanning 


2002] 


TABLE 1. RAPD FRAGMENTS AMPLIFIED FROM 76 COLLEC- 
TIONS. Size of each fragment (kb) is reported for six dif- 
ferent primers used in the analysis. Column numbers refer 
to the data matrix (Table 2). 


Column 1-16 17-31 32-34 45-49 50-62 63-79 
Primer A-9 A-10 A-15 A-16 A-18 A-19 
kos ~ 228) 2.60 3.50. — 1-95, 3.70 
ys 2et5 S90) 32:00: — 1.70-—'3.50 
Eos -- 2:00 12:80.> 1-90 — “1.607 2.50 
1.55 1.65 1:60" 1:40~— 1.53 °°'2.20 
1.45 60> 50) -" 30" ‘1.45 1.90 
£35 P50 3t55 L420 © 1270 
PaG, — E30\<- £25 138 1.50 
5 £20... £20 L395 1.48 
E20 2 Ris. ero 1.28 1.45 
Lal fe | 1.00 1.20 1.43 
ROS ~- 0:95-0:90 1.15 1.40 
95; 0:85-0:80 1.00 1.38 
ESS | O70 S “O75 O90 54230 
0.80 0.65 1.20 
0.70 0.60 1.10 
0.60 0.90 
0.85 


tree using the program Minspan (Podani 1993). A 
minimum spanning tree is the branching graph of 
OTU-wise association coefficients that minimizes 
the sum of all edges. Minimum spanning trees were 
computed for the RAPD data using the coefficient 
of Jaccard (Sneath and Sokal 1973), which does not 
include negative matches as a component of simi- 
larity (or dissimilarity). This is necessary when es- 
timating relationships from randomly amplified 
DNA data since absence of a site does not convey 
any useful information about relatedness. Many dis- 
tantly related OTU’s will have state 0 for a large 
number of sites. Most of those characters, identical 
in state, will not be identical by descent. The RAPD 
minimum spanning tree is plotted (Figs. 2—4) and 
mapped with representative fruits from plants bear- 
ing each of the cpDNA haplotype defined in Jessup 
(2002). Each plant included in the analysis is des- 
ignated with an OTU number in the diagrams cor- 
responding to a row in Table 1. 


Correspondence of cpDNA haplotypes and 
RAPD’s. The longest span in the tree falls between 
OTU 3 and OTU 5, and effectively defines a left 
and right half of the minimum spanning tree. 
cpDNA haplotypes from section Tritomopterys map 
in part to each side of the minimum spanning tree 
(Fig. 2). Most plants bearing cpDNA haplotypes 
from section Tritomopterys with asymmetric fruit 
wings map to the right half of the tree. All but one 
plant (OTU 5) bearing cpDNA haplotypes from 
section Tritomopterys that map to the left side of 
the tree have irregular or intermediate fruit wing 
symmetry and belong to G. implexa Jessup (notho- 
section Tritomochaudia). Plants bearing cpDNA 
haplotype D (OTU’s 64, 66, and 76) or cpDNA 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 


259 


haplotype B (OTU 59) having large symmetric fruit 
wings typical of G. cycloptera (DC.) W. R. Ander- 
son map in a cluster far to the right with plants 
from section Tritomopterys and are all members of 
nothosection Cyclotomopterys, either G. synoptera 
Jessup (OTU’s 59 and 66) or G. symplecta Jessup 
(OTU’s 64 and 76). Plants with asymmetric fruit 
wings bearing cpDNA haplotype D (OTU’s 2, 3, 6, 
8, 18) are all members of section Tritomopterys and 
map just to the right of the span separating the two 
halves of the spanning tree. 

Plants with cpDNA haplotypes from section Cy- 
clopterys (OTU’s 21, 59, 61) map to the same area 
on the minimum spanning tree as those with similar 
fruits bearing cpDNA haplotypes from section Tri- 
tomopterys (Fig. 4). Plants in G. mcvaughii W. R. 
Anderson, however, map to the far left (OTU’s 22 
and 23), even though they carry cpDNA haplotypes 
that clearly place them in section Cyclopterys (Fig. 
4). In particular, OTU 21 (in G. cycloptera), far 
right, and OTU 23 (in G. mcvaughii), far left, both 
bear cpDNA haplotype AB from section Cyclop- 
terys. One plant with morphology of G. cycloptera, 
and bearing a cpDNA haplotype from section Cy- 
clopterys (OTU 19), maps closer to the asymmet- 
ric-winged plants bearing section Tritomopterys 
cpDNA haplotypes. 

Plants bearing section Gaudichaudia cpDNA 
haplotypes all map on the left side of the RAPD 
minimum spanning tree (Fig. 3). One plant bearing 
a section Gaudichaudia cpDNA haplotype but with 
fruit wing morphology similar to G. cycloptera or 
G. mcvaughii, maps to the far left close to the po- 
sition of G. mcvaughii. Several plants having inter- 
mediate fruits and carrying section Gaudichaudia 
cpDNA haplotypes (Fig. 3) map just to the left of 
comer (OOS 52. 67, 5608;.09; 70. 71, 75). Plants 
with similarly intermediate fruit wing shapes but 
carrying section Tritomopterys cpDNA haplotypes 
map in the same region on the RAPD tree (OTU’s 
S49, 50, 56, 655° 72)2 All of ‘the plants 
with intermediate fruit wing morphology mapping 
to this region of the minimum spanning tree are G. 
implexa Jessup in nothosection Tritomochaudia. 
Plants with morphology typical of G. cynanchoides 
H. B. K. map near the center of the left half of the 
tree (OTU’s 40, 41, 42, 44, 45, 46, 47, 48). Three 
plants with intermediate fruits map to the far left, 
one bears section Tritomopterys cpDNA haplotype 
L (OTU 50), and the other two bear section Gau- 
dichaudia cpDNA haplotype S (OTU’s 49, 51). 
Those plants are also G. implexa. 

Plants in G. galeottiana (Nied.) Chodat, or with 
fruit morphology approaching that of G. galeotti- 
ana, and bearing section Zygopterys cpDNA hap- 
lotypes (Fig. 4) cluster together on the RAPD tree 
on one long branch just to the right of center 
(OTU’s 27, 28, 29). Plants in that cluster with 
slightly asymmetric wings are G. zygoptera Jessup 
(OTU’s 28 and 29) or G. intermixteca Jessup (OTU 
26) in nothosection Zygotomopterys. Only one 


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260 


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JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 261 


2002] 


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‘dHANILNOD ‘7 ATAV 


MADRONO 


[Vol. 49 


1400 bp = os 


SRS 


1400 bp = 


800 bps 


1400 bp- 
- ‘ :. Ais 


800 bp# 7 


Su 


102 104 106 108 110 112 113 115 117 119 121 123% 24 126 128 130. 132. 134 


Fic. 1. 


ie 


Example of RAPD products on agarose gel, in this case amplified with primer A-19. Gels were stained with 


ethidium bromide and visualized with UV transillumination. Molecular weight marker fragments are 100 bp apart. 
Lane numbers and the 800 and 1400 bp marker fragments are labeled. 


plant bearing a section Zygopterys cpDNA haplo- 
type (OTU 57) maps on the left side of the tree. 
That plant is notable in the morphological similarity 
of its fruit to other plants with intermediate samaras 
that carry section Gaudichaudia cpDNA haplotypes 
and map to the same region of the RAPD tree 
(OTU’s 52, 68, 75). It also falls close to OTU 58 
which has similar fruit shape but bears section Tri- 
tomopterys cpDNA haplotype L. OTU 57 is a geo- 
graphically isolated plant carrying a section Zyg- 
opterys cpDNA haplotype. A plant from the type 
locality of G. chasei W. R. Anderson (OTU 24), 


carrying cpDNA G. chasei cpDNA haplotype AE, 
maps close to the cluster of G. galeottiana on the 
RAPD tree (Fig. 4). A plant with intermediate fruit 
wing morphology (OTU 26), bearing section Tri- 
tomopterys cpDNA haplotype A, subtends G. chas- 
ei on the RAPD minimum spanning tree (Fig. 2). 
The only other plant collected that carries G. chasei 
cpDNA haplotype AE (OTU 63) is G. andersonii 
Jessup which maps with plants in G. cycloptera 
carrying section Cyclopterys cpDNA haplotypes 
(OTU’s 21, 61, 62). The narrowly endemic sub- 
shrubs, G. krusei W. R. Anderson (OTU 30) and G. 


2002] 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 


263 


RAPD minimum spanning tree mapped with fruits from plants bearing section Tritomopterys cpDNA 


23) Qs 


apy 


2060-49) 89) 88) 87474044 — 67) 68 


a) @2) 4348) G6) 70)69 ) 


haplotype OTU(s 42) Se GB. YA 

A 26 (G. intermixteca) Wy We tly 

B 4 (G. albida) 15, 16, 17 (G. ‘velutina’) e iy 

B 59 (G. synoptera) 

C 11 (G. ‘intermedia’) 

D 2,3 (G. albida); 6, 8 (G. diandra); 18 (G. ‘hirsuta’) 

D 66 (G. synoptera) 

D 64, 76 (G. symplecta) 

F 1 (G. albida) 

K 12, 13 (G. ‘intermedia’) 

L 5 (G. albida); 7, 9,73 (G. diandra); 14 (G. ‘intermedia’) 

L 25, 50, 53-56, 58, 65 (G. implexa) 

M 72 (G. implexa) 

N 10 (G. diandra) 28) B5) 
Fic. 2. Samaras from plants carrying cpDNA haplotypes from section Tritomopterys plotted on the RAPD minimum 


spanning tree. 


subverticillata Rose (OTU 31), map together on the 
extreme right side of the RAPD tree (Fig. 4). Those 
taxa are similar in the cpDNA haplotypes they car- 
ry as well. Gaudichaudia subverticillata and G. 
krusei form the paraphyletic section Archaeopterys 
comprising the internal outgroups to section Tvito- 
mopterys in the cpDNA haplotype phylogeny (Jes- 
sup 2002). OTU 38, which also carries a cpDNA 
haplotype positioned as an outgroup to section Tri- 
tomopterys in the phylogeny, has a fruit shape sim- 
ilar to G. cynanchoides and maps on the far left of 
the RAPD tree, close to G. mcvaughii. 


Southern hybridization results. Table 3 presents 
the results for lanes on the blots that showed hy- 
bridization to the probes. The number of fragments 
per lane hybridizing to the probe is given in the 
table for each probe used in the study. Chloroplast 
DNA haplotypes determined with restriction frag- 
ment analysis (Jessup 1994, 2002) are also indicat- 
ed for each lane in Table 3. In each of Figs. 5—10, 
the probe source is represented by an illustration of 
a samara at the top of the figure or to one side of 
the autoradiograph. A line points to the fragment 
that was labeled for the probe. Superimposed on 
the image of the autoradiograph are illustrations of 
Samaras from plants that showed strong hybridiza- 
tion to the probe. The results thus presented show 


the intersection of data from RAPD’s, cpDNA, and 
fruit wing morphology. 

When blots of the RAPD gels were probed with 
a labeled 1400 bp fragment from lane 12 (Anderson 
12937, G. diandra (Nied.) Chodat, Morelos, lane 
12) all (100%) of the plants expressing the 1400 bp 
fragment on the gel showed strong hybridization 
signal (Fig. 5). The surprising result was that all 
(100%) lanes showing hybridization to the probe 
hybridized to fragments of different molecular 
weights as well. Not surprisingly, the 1400 bp frag- 
ment (probe source) on the blot showed the stron- 
gest hybridization signal. Other fragments showing 
strong hybridization signal (secondary bands) on 
the blot showed up on the original gel as faint 
bands, but none of the secondary bands hybridizing 
the probes were visible on the gels as bright bands, 
and some of the secondary bands were not at all 
visible on the gel. Only rarely (one in twenty) did 
a lane without a 1400 bp fragment show hybridiza- 
tion to a fragment of a different size (middle frame, 
bottom panel), and that hybridizing fragment was 
the same molecular weight as a secondary fragment 
in lanes with multiple fragments hybridizing to the 
probe (Fig. 5). The same pattern of results is re- 
peated with the other probes. All bands of the same 
electrophoretic mobility hybridized to the probe 


264 


MADRONO 


[Vol. 49 


RAPD minimum spanning tree mapped with fruits from plants bearing section Gaudichaudia cpDNA 


23 


Oe 
ay 
SNS "Ds v 
4) 4S 


IS 
a 


S ? A 


OE) 


Ee 


Se WW. 
W 


Fic. 3. 


43.49 a8 COGN Ga 


40, 41, 44-48 (G. cynanchoides) 
49, 51, 67-71 (G. implexa) 

42 (G. cynanchoides) 

52 (G. implexa) 

75 (G. implexa) 


F 20 (G. cynanchoides x cycloptera) 


H8OGOSOL 


Samaras from plants carrying cpDNA haplotypes from section Gaudichaudia, plotted on RAPD minimum 


spanning tree. The right side of the tree is omitted since plants carrying section Gaudichaudia haplotypes are restricted 


to the left side of the tree. 


made from a band at that location. This clearly 
demonstrates that bands of the same molecular 
weight have sequence homology and can therefore 
be reliably considered identical by descent, at least 
in Gaudichaudia. 


Oligopterys 32 (G. hirtella) 


Fic. 4. 
chaeopterys, and section Oligopterys. 


RAPD minimum spanning tree mapped with fruits from plants bearing cpDNA 
haplotypes from sections Cyclopterys, Zygopterys, Archaeopterys, and Oligopterys 


The probe source in Fig. 6 is G. diandra. All of 
the plants hybridizing to the 1400 bp probe are 
found to cluster with the G. albida complex (sec- 
tion Tritomopterys and intersectional hybrids with 
section Tritomopterys) in the RAPD minimum 


AQ o@ 


CSS, 
cb AS 
66/30-31) 


O-O—O0-8'@- 
aye 


haplotype _ section OTU(s) 73) (9) 42) (1) 41) 
Y Cyclopterys 19 (G. cycloptera) 
AB Cyclopterys 60, 21 (G. cycloptera) 49 43 
AB Cyclopterys 61, 62 (G. synoptera, G. symplecta) 
AE Cyclopterys 24, 63 (G. chasei, G. andersonii) 
AB Cyclopterys 23 (G. mcvaughii) 
AC Cyclopterys 22 (G. mcvaughii) @7) 26 34 
AG Zygopterys 27 (G. galeottiana) 14 ip 
AD Zygopterys 57 (G. implexa x zygoptera) ata 4) 6 
Ww Zygopterys 28, 29 (G. zygoptera) NG & 
; bees HC 
G Archaeopterys 38 (indet. amphiploid) Spl | 
I Archaeopterys 31 (G. subverticillata) () 28 (is ee) 
J Archaeopterys 30(G. krusei) 


Samaras from plants carrying cpDNA haplotypes from section Cyclopterys, section Zygopterys, section Ar- 


2002] 


spanning tree (Figs. 2—4). The lanes showing hy- 
bridization to the G. diandra probe represent plants 
carrying cpDNA haplotypes from sections T7vito- 
mopterys, Cyclopterys, and Zygopterys. All of those 
plants bearing cpDNA from other than section 77i- 
tomopterys were identified as amphiploids on the 
RAPD minimum spanning tree. 

In the hybridization shown in Fig. 6, the probe 
(1150 bp) was taken from a member of the G. cy- 
nanchoides complex (Jessup 4112, Hidalgo, lane 
75). The probe was hybridized against the same set 
of blots shown in Fig. 5 (i.e., from primer A-19 
gels, Fig. 1). Again, all of the lanes showing hy- 
bridization signal except one are clustered together 
on the RAPD minimum spanning tree (Fig. 3), in- 
cluding plants identified as G. implexa, hybrids 
with maternal G. albida and paternal (pollen donor) 
G. cynanchoides. One lane, representing a plant 
with a samara morphology diagnostic of the G. al- 
bida complex (lane 6, Jessup 4052, Oaxaca), shows 
strong hybridization with the G. cynanchoides 
probe. That lane represents the same plant found 
clustering anomalously with the G. cynanchoides 
group on the RAPD minimum spanning tree (Fig. 
3, OTU 5). The only other lane represented by a 
G. albida samara (lane 119, Jessup 4032, Michoa- 
can) shows a relatively weak signal, suggesting 
some involvement with the G. cynanchoides group, 
perhaps via introgression. The two strong signals in 
the right panel of the top gel (Fig. 6) are bound 
probe from the first run (cf. Fig. 6) that did not 
melt from the blotted DNA during stripping of the 
probe. These “‘ghost band”’ appear on other blots 
as well. 

The hybridization run represented in Fig. 7 used 
the same blots discussed above. The probe for this 
hybridization experiment (1900 bp) is from the 
same source as that described for Fig. 6 (Jessup 
4112, Hidalgo, lane 75). Here again we see mainly 
plants from the G. cynanchoides group hybridizing 
to the probe (most of gel panel 3). Again we see 
the anomalous plant from the G. albida complex 
(lane 6, Jessup 4052, Oaxaca) hybridizing to the 
probe. A possible G. cycloptera X G. cynanchoides 
hybrid also shows up on this autoradiograph (gel 
panel 2). Aspicarpa, the putative sister lineage to 
Gaudichaudia, has recently been shown to nest, at 
least in part, within Gaudichaudia (Davis et al. 
2001). The Aspicarpa accession used in this study 
(G. hirtella (Rich.) Jessup, formerly in Aspicarpa), 
also hybridized to the 1900 bp probe. Three of the 
suspected amphiploids (G. implexa) hybridizing to 
the 1900 bp probe (gel panel 4) also hybridized to 
the 1150 bp probe (Fig. 6, panel 4). Two of the 
amphiploids (G. implexa) hybridize to the 1900 bp 
probe but not to the 1150 bp probe, and two (also 
G. implexa) hybridize to the 1150 bp probe but not 
to the 1900 bp probe (Fig. 8). 

Likewise, within the G. cynanchoides complex 
(Fig. 7, gel panel 3) some plants hybridize to one 
probe but not the other, while some hybridize to 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 


265 


both (Fig. 8). Plants that hybridized just to the 1900 
bp probe all have morphology typical of G. cynan- 
choides, and all bear the cpDNA haplotype S, 
which is the most common cpDNA haplotype with- 
in section Gaudichaudia, the G. cynanchoides com- 
plex (Fig. 3). Two of the three plants binding the 
1150 bp probe also bear cpDNA haplotype S. The 
other plant bears cpDNA haplotype L, the most 
common cpDNA haplotype the G. albida complex. 
The samara morphology of that plant is, however, 
more typical of G. implexa, the amphiploids be- 
tween G. albida and G. cynanchoides (section Tri- 
tomochaudia). Most of the plants hybridizing to 
both probes show the samara morphology typical 
G. implexa. Eight out of fifteen plants binding both 
probes have cpDNA haplotypes from section Tri- 
tomopterys, indicating their affinity with the G. al- 
bida complex, six have cpDNA haplotypes from 
section Gaudichaudia, indicating their affinity with 
the G. cynanchoides complex, and one has a cp- 
DNA haplotype identified as a member of section 
Archaeopterys, an outgroup to section Tritomopter- 
ys. 

Experiments using primer sequence CAGGCC- 
CTTC (primer A-1) yielded similar results (Fig. 9). 
Blots were probed with two labeled fragments from 
the gels used to produce the autoradiographs. The 
first probe was made with a 1400 bp fragment col- 
lected from lane 47 (Jessup 4039, G. galeottiana, 
Puebla). The second probe was made with a 1000 
bp fragment collected from lane 16 (Jessup 4055, 
G. albida, Chiapas). The first probe (from G. gal- 
eottiana) hybridized strongly to five collections 
from section Zygopterys, including the source lane 
(G. galeottiana) three lanes representing G. zyg- 
optera and one lane representing G. intermixteca. 
Lanes representing G. krusei (section Archaeopter- 
ys) and G. hirtella (section Oligopterys, formerly 
in Aspicarpa) were also hybridized by the probe. 
Hybridization signal from lanes representing G. 
krusei and G. hirtella were weak, but stronger than 
hybridization to lanes representing other elements 
of Gaudichaudia. The second probe (from G. al- 
bida) hybridized strongly to plants in the G. albida 
complex, and to several of the amphiploids. Two 
plants were hybridized by both probes. One of 
those plants (Jessup 4047, Oaxaca) carries a cp- 
DNA haplotype (A) from section 7Jritomopterys, 
the clade associated with the G. albida complex. 
The other plant (Anderson 12990, Oaxaca) carries 
a cpDNA haplotype (X) from section Zygopterys, 
the clade associated with the G. galeottiana com- 
plex. From this we can infer that Anderson 12990, 
carrying RAPD sites from both G. albida and G. 
galeottiana and a cpDNA haplotype from the G. 
galeottiana complex, must be derived from a cross 
between a plant in the G. albida complex, the pol- 
len donor, and a plant in the G. galeottiana com- 
plex, the maternal parent (Fig. 9). The hybrid, Jes- 
sup 4047, must have formed the other way around. 
The pollen donor was from the G. galeottiana com- 


266 MADRONO [Vol. 49 


TABLE 3. LABELED RAPD PROBES HyYBRIDIZED TO BLOTS. Numbers of bands hybridizing probes is recorded for all 
lanes with a strong hybridization to the probe. Lanes that were blank on the gels have been omitted. Collections are 
grouped by taxonomic assignment based on morphology. Asterisks indicate probe sources. 

Primer Probe 


Collec- Al Al A-2 A-3 A-7 A-19 Al9 A-19 A-19 


tion # cpDNA Lane # 16:1000 47:1400 8:1250 28:950 4:850 12:14000 12:1100 75:1150 75:1900 
Section Tritomoptetys 
Chi4056 cpDNA 1-F 1 4 2 
Gro4067 cpDNA 1-D 3 Zz 4 2) 4 3 
Mex13275 cpDNA 1-D 4 3 4 £2, 4 3 
Oax4041 cpDNA 1-B ) 3 4 D 
Oax4052 cpDNA 1-L 6 3 2) 2 D 3 
Gro4066 cpDNA 1-D 8 3 *A 2D 4 3 
Mic13309 cpDNA 1-L 10 3 2 
Mor12937 cpDNA 1-D IZ 4 *A #8) 
Nay4088 = cpDNA I-L 13 4 Dy) 3 
S$in4102 cpDNA 1-N 14 3 4 2 3 
Chi4055 cpDNA 1-C 16 oS) 4 2 2 
Chil3244 cpDNA 1-K 18 It 4 D) 
Chil3225 cpDNA 1-K 19 3) 4 2 D) 
Jal4084 cpDNA 1-L aM D) 
Jal4085 cpDNA 1-L 22 2) 
Oax405 1 cpDNA 1-L 24 3 4 2 
Oax13216 cpDNA 1-K 26 3 4 2 2 
Chi4058 cpDNA 1-B Di 2) 4 
Chi4060 cpDNA 1-B 28 4 oe 
Chi4061 cpDNA 1-B 29 4 2 2 4 
Oax4054 cpDNA I-K 30 4 2 
Oax4048 cpDNA 1-D 37) 4 4 
Natural and Artificial Hybrids 
Zac4 106 cpDNA 1- 84 4 
Zacl2624 cpDNA 2- 85 4 2 3 3} 4 
Gua4006 cpDNA I- 86 4 2 3 


1-L 
2-S 
1-L 
Gua4108 cpDNA 1-L 
Hid32522 cpDNA 1-L 88 
1-L 
1-L 
1-L 
1-L 


(ee) 
~ 
iW) 
OO 


4 2 2) 3 
Tam4001 cpDNA 1- 91 4 D 1 3 4 
Tam4113. cpDNA I- 92 4 2, 3 
Tam4114 cpDNA 1- 93 4 2 1 4 
Nue4115 cpDNA 1- 94 4 2) 1 3) 4 
SLP4002. cpDNA 1-L 95 1 3 4 
SLP4043, + =cpDNA 4-AD 97 1 4 2 1 3) 4 
SLP4056 cpDNA 1-L 98 4 2 1 3 4 
Mex4020 cpDNA-1B 102 4 2 3 
Mex4021 cpDNA 1-AB 103 4 3 
Mex4023 cpDNA 1-AB 104 4 2 4 
Mex4025 cpDNA 1-AB 106 4 
Mex4026 cpDNA 5-AE 107 4 3 
Mic4074 cpDNA 1-D 108 4 2 3 
SLP13316 cpDNA 1-L 109 4 D l 3 2 
Jal4076 cpDNA 2-S 11] 4 3 3 
Jal4079 cpDNA 2-S 112 4 2 3 5 4 
Jal4080 cpDNA 2-S 3 2, 2 3 
Jal408 1 cpDNA 2-S 114 2 
Jal4082 cpDNA 2-S 115 2 3 
Jal3707 cpDNA 1-M 116 2 I 3 + 
Mic403 1 cpDNA 1-L 118 3 
Mic4032 cpDNA 1-L tS 4 Ss) 
Mic4034 cpDNA 1-L I 2 2 | 
Mic4035 cpDNA 2-P 122 D 
Mic4037 cpDNA 2-Q 124 3 4 2 2 3 
Mic13291 cpDNA 1-D QS 4 2 2: 
Oax4043 cpDNA 4-AD 2 3 4 2 
Oax4044. cpDNA 4-V 128 3 4 4 Z 3 


2002] JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 267 
TABLE 3. CONTINUED. 
Primer Probe 

Collec- Al Al A-2 A-3 A-7 A-19 Al19 A-19 A-19 

tion # cpDNA Lane # 16:1000 47:1400 8:1250 28:950 4:850 12:14000 12:1100 75:1150 75:1900 
Oax4046 cpDNA 1-A 130 p) 
Oax4047 cpDNA 1-A 131 2 + 2 
Oax13031 cpDNA 1-A 132 4 p) 
H687 cpDNA 1-D 39 4 2 4 3 
H2187 cpDNA 1-L 36 4 2 
H2487 cpDNA 1-M 64 4 3 3 3 4 
H5487 cpDNA 1-L 65 4 2 2 l 3 4 
H289 cpDNA 1-L 66 4 j 3 4 
H489 cpDNA 1-L 67 4 l 3 4 
H4387 cpDNA 1-L 99 4 2 l 3 4 
H689 cpDNA 1-L 100 4 ] 3 
H4687 cpDNA 1-L 101 4 2 ] 3) 4 
H2586 cpDNA 1-L 133 3 + 5) 3 
H2687 cpDNA 1-M 134 3 2 3 4 

Section Guadichaudia cpDNA 
Zac4103 cpDNA IO-G 68 ] 3 4 
Zac4104 cpDNA 2-R 69 ] 3 
Gua4107 cpDNA 2-R 70 l 3 3 
Gua4007 cpDNA 2-R a 3 
Hid4109 cpDNA 2-S 72 4 1 3 4 
Hid4110 cpDNA 2-S a3 + ] 3 3 
Hid4111 cpDNA 2-S 74 + 2 3 3 
Hid4112 cpDNA 2-U iS + ] 23) *4 
SLP4004. cpDNA 2-S 76 4 3 
Jal4077 cpDNA 2-S 78 2 l 3 4 
Jal4078 cpDNA 2-S TS 3 
Mex82260 cpDNA 2-S 80 2, 3 
Mic4075 cpDNA 2-S 81 2 ] 3 4 
Mic13296 cpDNA 2-S 82 2 3 
Zac4105 cpDNA 2-S 83 4 2 3 3 3 
Sections Cyclopterys, Zygopterys, Oligopterys and outgroups 

Gro4510 cpDNA 3-AF 39 4 2 2 3 
Jal4083 cpDNA 3-AB 40 = 
Col12699 cpDNA 3-AC 4] 4 
Oax4293 =ocpDNA 1-L 44 3 4 2 l 3 4 
Pue4040 cpDNA 1-A 45 4 2 
Pue4039 cpDNA 4-AG 47 *4 
Oax4049, = =cpDNA 4-W 49 p 4 2 3 
Oax12990 cpDNA 4-X 50 3 4 4 2 
Oax13138 cpDNA 4-W 51 4 2 3 
Gro4069 cpDNA IO-J 54 2 
Asp525 cpDNA ASP 56 2 3 
Asp526 cpDNA ASP a7 2 
Jan1254 cpDNA JAN 59 2 
Jan3373 cpDNA JAN 61 2 


plex and the maternal lineage from the G. albida 
complex (Fig. 9). 

Blots made from gels of RAPD produced with 
primer sequence TGCCGAGCTG (A-2) were 
probed with a labeled 1250 bp fragment from G. 
diandra (lane 8, Jessup 4066, Guerrero) (Table 3). 
The probe bound strongly to RAPD fragments from 
plants throughout the genus. About 77% of the 
plants sampled in the G. albida complex hybridized 
the probe, but only 40% of the plants sampled in 
the G. cynanchoides complex hybridized the probe. 
About 76% of the plants sampled from among the 


amphiploids hybridized the probe. Representatives 
from G. cycloptera, G. mcvaughii, and G. galeot- 
tiana also bound the probe. 

Blots made from gels of RAPD fragments pro- 
duced with primer sequence AGTCAGCCAC (A- 
3) were probed with a labeled 950 bp fragment am- 
plified from an unpublished species in the G. albida 
complex (lane 28, Jessup 4060, Chiapas) (Table 3). 
About 86% of the plants sampled in the G. albida 
complex hybridized the probe. Only 13% of the 
plants sampled in the G. cynanchoides complex hy- 
bridized the probe. About 53% of plants sampled 


268 


MADRONO 


[Vol. 49 


RAPD A-19: Primer sequence: *>CAAACGTCGG?* 


Probe source: lane 12, ~1400 bp; Morelos 
r ton 2 


Fic. 5. 


~N 


Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1) probed with 1400 bp 


fragment from lane 12 (Anderson 12937, G. diandra, Morelos), indicated by arrow. Fruits are illustrated in lanes 
showing strong hybridization to the probe for all plants with fruits available. 


in the amphiploid complex hybridized the probe. 
The probe also hybridized to one plant in the G. 
cycloptera complex, and two plants from the G. 
galeottiana complex. 

Blots made from gels of RAPD fragments pro- 
duced with primer sequence GAAACGGGTG (A- 
7) were probed with a labeled 850 bp fragment 
from G. albida sensu stricto (lane 4, Anderson 
13275, Mexico) (Table 1). The probe bound strong- 
ly to diverse elements of Gaudichaudia and to both 
collections of Janusia, another closely related ge- 
nus, included in the sample, but did not hybridize 
to G. hirtella (formerly in Aspicarpa), and was not 
prevalent within any of the groups sampled. About 
22% of the plants sampled in the G. albida complex 
hybridized the probe, and about 27% of plants sam- 
pled in the G. cynanchoides complex hybridized the 
probe. Among members of the amphiploid com- 
plex, about 26% hybridized the probe. Two plants 
representing the G. galeottiana complex, and one 
member of the G. cycloptera complex also hybrid- 
ized the probe. 

Comparing the distribution of hybridization sig- 
nal across probes, two probes hybridized specimens 


predominantly in the G. cynanchoides complex (A- 
19 75:1150 and A-19 75:1900) but hybridized very 
few specimens in the G. albida complex. Three 
probes (A-1 16:1000, A-3 28:950 and A-19 12: 
1400) hybridized specimens predominantly in the 
G. albida complex but very little or not at all in 
the G. cynanchoides complex. All five of those 
probes hybridized samples prominently in the am- 
phiploid complex. Among the amphiploids, probes 
hybridizing predominantly in the G. albida com- 
plex were combined in some combination with 
probes hybridizing predominantly in the G. cynan- 
choides complex in about 26% of the plants sam- 
pled. 


DISCUSSION 


The assumption of homology. One of the key as- 
sumptions allowing the use of RAPD markers as 
characters is that bands occurring in different sam- 
ple lanes at the same position, i.e., bands having 
the same molecular weight, have DNA sequences 
sharing sequence homology, and are therefore re- 
lated by ancestry. One way to test that assumption 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 269 


2002] 
RAPD A-19: Primer sequence: *>CAAACGTCGG* 
Probe source: lane 75, ~1150bp; Hidalgo 

Fic. 6. 


fragment from lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits are illustrated in lanes 
showing strong hybridization to the probe for all plants with fruits available. 


would be to sequence several sites having the same 
molecular weight, but that approach is expensive 
and would be limited to relatively few specimens. 
Use of RAPD hybridization blots permits screening 
a large set of DNA’s, in this study representing 75 
plants in the genus. The procedure probes a radio- 
labeled RAPD fragment of known size and source 
against Southern blots made from the RAPD gels. 
RAPD fragments on the blot hybridizing to the la- 
beled probe must have substantial sequence ho- 
mology for strong hybridization. In the experiments 
reported here only lanes that showed a strong hy- 
bridization signal were included in the data matrix. 
Many lanes showed weak binding of the probe and 
might have some sequence homology, but diver- 
gence was sufficient to weaken the signal. Weak 
hybridization is expected from the primer sequence 
alone. 


The assumption of independence. In addition to 
the question of sequence homology of RAPD frag- 
ments of the same weight across samples, there is 
the question of sequence homology of RAPD frag- 
ments of different weight within a sample. Since 


both the theory of how RAPD markers behave in 
amphiploids and the empirical evidence presented 
here suggests that hybrids combine distinct RAPD 
sites of the parental lineages, we expect that some 
of those sites will be homologous, or more specif- 
ically, synologous (Mindell and Meyer 2001 )—di- 
vergent and descended from a common ancestor 
but residing in the same genome by virtue of retic- 
ulate ancestry. Synologous fragments would have 
sufficient sequence divergence (insertions, dele- 
tions, substitutions) to express different electropho- 
retic mobility, but would retain enough sequence 
homology to hybridize to a probe from the synol- 
ogous locus. Changes in size of a RAPD site caused 
by insertion/deletion events are likely to develop in 
reproductively isolated lineages. When the lineages 
bearing the divergent sites merge in an amphiploid 
each will be expressed, resulting in complementa- 
tion. In the absence of recombination between pa- 
rental genomes the hybrid lineage would then be a 
fixed heterozygote. Detection of fixed heterozygos- 
ity corroborates other evidence supporting an am- 
phiploid origin of the lineage. It is tempting to in- 


270 


MADRONO 


[Vol. 49 


RAPD A-19: Primer sequence: °>CAAACGTCGG* 


Probe source: lane 75, ~1900bp; Hidalgo 


126 128 130 132 1 


119 121 123%y 


Fic. 7. Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1) probed with 1900 bp 
fragment from lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits are illustrated in lanes 
showing strong hybridization to the probe for all plants with fruits available. 


terpret multiple bands with sequence homology as 
indicating successive layers of fixed heterozygosity 
built up in the tiered genomes of an ancient poly- 
ploid complex. 

The evidence presented here is consistent with 
synologous origins for fragments that bind the 
probe but which have different molecular weights 
from that of the probe. Competing hypotheses can 
not, however, be ruled out with the available data. 
For example, multiple fragments hybridizing the 
probe within a specimen could indicate multiple 
nested priming sites within the amplification win- 
dow of PCR conditions used. Some PCR products 
would encompass three pairs of priming sites, some 
two, and some only one, resulting in three frag- 
ments sharing overlapping sequence identity. Other 
scenarios explaining the appearance of different 
sized RAPD fragments with sequence homology 
can be envisioned. Without a detailed study of how 
the coamplifying fragments are arranged on the 
chromosomes it is not possible to support or reject 
the alternative hypotheses. 


Introgression. A plausible explanation for the 
observation that all plants binding both the A-19 
1900 bp and A-19 1150 bp probes show morphol- 
ogy typical of hybrids between G. cynanchoides 
and G. albida (Fig. 8) is that cryptic sibling species 
within G. cynanchoides are forming tetraploids and 
crossing, and those polyploids are capable of form- 
ing amphiploids with similar lineages from the G. 
albida complex. It is quite possible that some lin- 
eages in the G. cynanchoides complex (and in other 
groups as well) are geographically restricted where- 
as the polyploid lineages involved in most of the 
wide crosses are weedy and wide ranging and carry 
more of the genetic diversity as fixed heterozygos- 
ity. 
If we compare the autoradiograph of the A-19 
1150 bp probe (Jessup 4112, G. cynanchoides, Hi- 
dalgo, lane 75), and that of the A-19 1100 bp probe 
(Anderson 12937, G. diandra, Morelos, lane 12) 
for just eight plants from the G. cynanchoides com- 
plex (lanes 81—88) (Fig. 10), we observe one plant 
hybridizing only to the 1100 bp probe (probe from 


2002] 


Probe source: lane 75; Hidalgo 
mat68 70 72 #7 7% 7 79 8 


1900 bp 


= —- 2 a> ae we oo 


1150 bp Ea 


vie 2 eee 


1900 bp 
blot ii 


(cpDNA from 
section Tritomopterys) 


(cpDNa from 
section Gaudichaudia) 


os ‘s M3 é 3 

\ ; > 
cpDNA from id \ 
section Archeopterys 


G. cynanchoides 


Fic. 8. 


: 2 Plants hybridizing on 


cpDNA from section Gandichaudia 
| G. implexa 
Plants hybridizing both 1900 bp and 1150 bp probes displayed in order of appearance on blot. 


JESSUP: RAPD HYBRIDIZATION ANALYSIS OF RETICULATE ANCESTRY 271 


RAPD A-19: Primer sequence: °>CAAACGTCGG? & 


~ 


a. , 
“sy 


F 3 a 
, ly the 
i” 1900 bp probe; all section Gaudichaudia 


- 
s & @ 


7 (cpDNA from 
"section Gaudichaudis) 


Ap 


3 ne intermixteca 


vial saat 


\ 
cpDNA from section Tritomopterys 


Autoradiographs of RAPD products from primer A-19 (blotted from gels in Fig. 1), lanes 68-101. Gel, at top, 


is compared with autoradiographs from 1900 bp probe and 1150 bp probe. Both probes were prepared from fragments 
in lane 75 (Jessup 4112, G. cynanchoides, Hidalgo), indicated by arrow. Fruits illustrated top panel are from plants 
hybridizing only the 1900 bp probe; fruits illustrated middle panel are from plants hybridizing only the 1150 bp probe; 
fruits illustrated bottom panel are from plants hybridizing both the 1900 bp and 1150 bp probes. In bottom panel 
sectional affiliation of cpDNA haplotypes is indicated for each fruit illustrated, and dotted lines separate described 


species. 


G. albida sens. lat.), two plants hybridizing only to 
the 1150 bp probe (probe from G. cynanchoides), 
and three plants hybridizing to both probes. The 
plants hybridizing both probes all carry cpDNA 
haplotypes characteristic of section Gaudichaudia. 
Two of the three plants hybridizing both probes are 
G. implexa (nothosection Tritomochaudia) and ex- 
hibit the samara morphology of the amphiploids. 
The plant hybridizing only the 1100 bp probe car- 
ries a cpDNA haplotype from section Tritomopter- 
ys, the G. albida complex. Of the two plants hy- 
bridizing only the 1150 bp probe, one carries a 


cpDNA haplotype from section Tritomopterys, and 
one carries a section Gaudichaudia cpDNA hap- 
lotype. 

One explanation for sites shared in this way is 
that introgression is occurring between the G. dian- 
dra lineages and the G. cynanchoides lineages. The 
1100 bp probe (from G. diandra) hybridizes to sev- 
eral plants in the G. albida group, and to almost 
every plant in the G. cynanchoides group. Among 
the plants in the G. cynanchoides group, plants that 
exhibit the typical cynanchoid samaras hybridize 
only a single fragment, the 1100 bp fragment. In 


ie 


RAPD A-1: primer sequence: 
*CAGGCCCTTC* 
Probe source: lane 47, 1400 bp, Puebla 


G. galeottiana (section Zygopterys) 


G. intermixteca 
Oaxaca 


i 


G. zygoptera 


(nothosection se 


G. zygoptera 


(cpDNA from 
section Zygopterys) 
Pollen source = Pollen source = 
G. albida sens. lat. G. galeottiana 


Two plants bound both 1000 and 1400 bp probes. 


(_pDNA from 
section Tritomopterys) 


Fic. 9. Autoradiographs of RAPD products from primer 
A-1 probed (upper panel) with 1400 bp fragment from 
lane 47 (Jessup 4039, G. galeottiana, Puebla), indicated 
by arrow. Lower panel shows the two lanes with a strong 
hybridization to both the 1000 bp probe from lane 16 (Jes- 
sup 4055, G. albida sensu lato, Chiapas) and the 1400 bp 
probe from lane 47. 


the amphiploids the probe hybridizes to two or 
three fragments. The single G. cynanchoides frag- 
ment is evident in lane 81, Fig. 10 (lanes other than 
81—88 are not illustrated for the 1100 bp probe). 
That observation is consistent with the hypothesis 
that G. cynanchoides, which carries a single variant 
(1100 bp in length), is introgressing with G. dian- 
dra. It is possible, though, that the probe source 
(Anderson 12937, G. diandra, Morelos, lane 12) is 
itself a fixed heterozygote. The probe hybridizes to 
all three fragments in the probe source lane (not 
illustrated). Though the evidence is suggestive, it 


MADRONO 


[Vol. 49 


RAPD A-19: Primer sequence: 
*CAAACGTCGG* 


Probe source: lane 12: 1100bp fragment ~% 
section Tritomopterys) 


(cpDNA fom 
section Tritomopterys) 


"85 86 87 88 


Probe source: lane 75: 1150bp fae Gh 
(section Gaudichaudia) 


(cpDNA from 
section jPrajemliae) Co 


» (cpDNA from 
section Gaudichaudia) 


Plants binding only 1150 EB probs 


G. implexa 


G. cynanchoides 
Plants binding both 1100 and 1150 bp probes. 


Fig. 10. Autoradiographs of RAPD products from 
primer A-19 (blotted from gels in Fig. 1), lanes 81-88. 
Upper panel shows hybridization to the 1100 bp probe 
from lane 12 (Anderson 12937, G. diandra, Morelos). 
Middle panel shows hybridization to 1150 bp probe from 
lane 75 (Jessup 4112, G. cynanchoides, Hidalgo). Fruits 
illustrated in bottom panel for plants hybridizing both 
probes. 


may not be possible with the sample in hand to 
eliminate the hypothesis that what we are seeing 1s 
a shared polymorphism. Sequencing the RAPD 
fragments would show how length variants differ. 
Examining variation of RAPD fragment patterns 
within and among populations of each species 
would be a fruitful approach to the population level 
dynamics of introgression. 


Conclusion. What we can say for certain is that 
RAPD fragments of the same molecular weight can 
be reliably considered homologous sequences. We 
can also conclude that fragments of different mo- 
lecular weight within a lane, i.e., coming from the 
same nucleus, often represent size variants of a sin- 


2002] 


gle RAPD site. Hybrids exhibit complementation of 
sites from parental lineages that are presumably 
fixed for different size fragments. This allows us to 
use the presence of fragments on the gels as binary 
characters in phenetic analysis of relatedness 
among the collections, even though different size 
fragments have homologous sequences. 

We can get even more specific in identifying the 
parental lineages for a given hybrid by combining 
information from the Southern transfer hybridiza- 
tion experiment with information from analysis of 
cpDNA restriction site data. The experiments using 
primer sequence CAGGCCCTTC (primer A-1) 
demonstrate this application of RAPD hybridization 
(Fig. 9). From the forgoing evidence it seems likely 
that RAPD fragments of different molecular 
weights but similar sequence, as demonstrated by 
strength of probe hybridization, can be used to 
characterize genomes within polyploids. Markers 
thus developed can be used to resolve reticulate an- 
cestry in amphiploid complexes that are otherwise 
intractable. 


ACKNOWLEDGMENTS 


The author thanks William R. Anderson, who gener- 
ously shared his knowledge and research collections and 
provided guidance and assistance with field work, and E. 
Pichersky and R. Fogel, who provided laboratory space 
and equipment. This research was funded in part by NSF 
grant BSR-8700340 to W. R. Anderson, and by NSF Doc- 
toral Dissertation Improvement grant BSR-8823076 to W. 
R. Anderson for S. L. Jessup. 


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1994. Reticulate evolution in Gaudichaudia 
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. 2002. Six new species and taxonomic revisions 
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RIESEBERG, L. H. 1996. Homology among RAPD frag- 
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SAMBROOK, J., E. E FRITSCH, AND T. MANIATIS. 1989. Mo- 
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Spring Harbor Laboratory Press, Cold Spring Harbor, 
NY. 

WoLFE, A. D. AND A. LISTON. 1998. Contributions of 
PCR-based methods to plant systematics and evolu- 
tionary biology. Chapter 2 in D. E. Soltis, P. S. Soltis 
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MADRONO, Vol. 49, No. 4, pp. 274-284, 2002 


LONG-TERM POPULATION DYNAMICS OF NATIVE NASSELLA 
(POACEAE) BUNCHGRASSES IN CENTRAL CALIFORNIA 


JASON G. HAMILTON! 


Department of Ecology, Evolution and Marine Biology, University of California, 
Santa Barbara, CA 93106 


JAMES R. GRIFFIN AND MARK R. STROMBERG2 


Hastings Natural History Reservation, Museum of Vertebrate Zoology, University 
of California, 38601 E. Carmel Valley Road, Carmel Valley, CA 93924 


ABSTRACT 


California bunchgrass communities are one of the most endangered ecosystem types in the United 
States. In this study, we sought to determine long-term (52+ years) changes in populations of native 
bunchgrasses, Nassella pulchra (A. Hitchce.) Barkworth and Nassella cernua (Stebb. & Love) Barkworth, 
in unmanaged stands. At the landscape scale, Nassella has increased. However, population dynamics of 
individual stands appeared related to land-use history. Non-native annuals, by themselves, did not seem 
to cause decline of Nassella stands, but light grazing did cause reduction of Nassella basal cover. Areas 
that were historically cultivated supported Nassella stands with lower basal cover and size distributions 
qualitatively different from areas that were never cultivated. Mortality of Nassella was concentrated in 
small plants. Interspecific interference probably was important in limiting seedling recruitment in stands 
with low Nassella basal cover, and intraspecific interference appeared to become important as Nassella 
basal cover increased. Even in the presence of non-native annuals, Nassella stands in areas that have not 
been disturbed by cultivation do not appear to require management for maintenance. New individuals are 
recruiting into populations, and conservative a estimate of longevity of large individuals of Nassella is 
100 years. However, in areas that have been cultivated, active management may be required to increase 


the abundance of Nassella. 


Key Words: California, Nassella, grassland, survival, longevity, long-term. 


INTRODUCTION 


Perennial bunchgrass communities are one of the 
rarest plant communities in California (Keeley 
1989, 1993) and are considered to be one of the 
most endangered ecosystem types in the United 
States (Noss et al. 1995; Peters and Noss 1995). 
Since the founding of the Spanish missions in the 
mid-1700’s, massive invasions of annual grasses 
from the Mediterranean basin have altered native 
communities to such a degree that today the origi- 
nal extent and composition of these communities is 
unknown (Keeley 1989; Heady et al. 1992; Ham- 
ilton 1998). Today in California, an area of approx- 
imately 7,000,000 ha is dominated by non-native 
annual grasses (Huenneke 1989). In many cases, 
these non-natives comprise from 80% to 100% of 
the cover (Biswell 1956; Heady 1956; Macdonald 
et al. 1988; Heady et al. 1992), and the small patch- 
es of perennial bunchgrasses that still exist in Cal- 
ifornia (including Nassella (=Stipa) pulchra (A. 
Hitche.) Barkworth and closely related Nassella 
cernua (Stebb. & Love) Barkworth) always include 
non-native grasses. 


' Present address: Biology Department, 161 CNS, Ith- 
aca College, Ithaca, NY 14850-7278. 

* Author for correspondence, e-mail: 
socrates.berkeley.edu 


stromber @ 


In most of California, the original community 
composition of areas in which Nassella bunchgrass- 
es are found today is a matter of conjecture (Ham- 
ilton 1998). However, it is clear that over the past 
two hundred years, the biotic environment has 
changed dramatically for these bunchgrasses (Dyer 
and Rice 1999). There are no ‘pristine’ areas of 
California grassland left. Non-native annual grasses 
such as Bromus hordeaceous L., Bromus diandrus 
Roth, Avena fatua L., and Avena barbata Link have 
invaded every known bunchgrass stand. Further- 
more, due to land clearing, farming, and extreme 
over-grazing (Burcham 1957), even areas that are 
currently protected have been previously disturbed 
in some manner. Because of a lack of long-term 
studies, it is unknown whether a new steady-state 
situation has been achieved in the California grass- 
lands, or whether bunchgrass stands are still ad- 
justing to the altered conditions. 

In California, there is a growing interest in res- 
toration and conservation of Nassella bunchgrass 
communities (Knapp and Rice 1994; Stromberg 
and Kephart 1996; Carlsen et al. 2000; Kephart 
2001). Attempts at generalized management pre- 
scriptions that promote grazing and/or burning as a 
tool to reduce competition from annual grasses and 
enhance longevity of mature bunchgrasses have 
been proposed (e.g., Menke 1992) for inland sites. 
Substantial differences are evident between inland 


2002] 


and coastal native grasslands where Nassella is a 
co-dominant (Stromberg et al. 2001). On inland 
sites, recruitment of Nassella appears to be limited 
by competition by non-native annuals (Dyer and 
Rice 1999) and management strategies have been 
developed to improve establishment by reducing 
exotic seed banks (Stromberg et al. 2002). Studies 
of mortality and recruitment, along with restoration, 
are lacking in coastal environments of California 
and management strategies suggested or inland 
sites (Menke 1992) may require modification. We 
lack the fundamental information concerning long- 
term stability characteristics of Nassella stands in 
the face of competition from non-native annual 
grasses. Because of this, land managers have been 
forced to rely on hearsay to determine whether na- 
tive grasslands require management in order to per- 
sist, and if so, what kind. Results from short-term 
studies (one-two years) have tended to be unreli- 
able indicators of longer-term dynamics. For ex- 
ample, in one study, preliminary results after 16 
months indicated that burning and early-spring 
grazing were effective at increasing Nassella pul- 
chra seedling establishment and survival (Fossum 
1990). However, in the same study, after four years, 
it was concluded that burning and grazing were not 
effective at enhancing Nassella pulchra seedling re- 
cruitment (Dyer et al. 1996). 

In our study, we sought to determine long-term 
changes in populations of Nassella bunchgrasses in 
unmanaged stands. In particular, we asked: (1) In 
the absence of fire or grazing, has Nassella in- 
creased or decreased at the landscape scale? (2) At 
the scale of individual stands, are established pop- 
ulations of Nassella stable? (3) At the scale of sin- 
gle individuals, is there life-stage-related mortality 
that suggests interference from non-native grasses? 
(4) Do trends in multi-scale population dynamics 
suggest that Nassella requires management for per- 
sistence when there is interference from non-native 
annuals? 

There are very few sites in California where data 
exist that allow for analysis of long-term trends in 
Nassella bunchgrass populations. One such place is 
the Hastings Natural History Reservation in the 
foothills of the South Coast Range in central Cali- 
fornia. Here, in a study initiated in 1944 by G. L. 
Stebbins, White (1966) described old field succes- 
sion Over a 22-year period. Using unpublished data 
from studies by both Stebbins and White and more 
detailed data from a number of other bunchgrass 
stands at Hastings Reservation, we have been able 
to extend the original findings of White to encom- 
pass a period of 52 years, and to compare a number 
of sites around Hastings Reservation representing 
many ecologically distinct situations with different 
land-use histories. 


METHODS 


The 911-ha Hastings Natural History Reserva- 
tion (36°33'30"’N, 121°33'30”W) is located in the 


HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 


275 


interior foothills of the coastal Santa Lucia Moun- 
tains in central California, 33 km southeast of Mon- 
terey. The Mediterranean climate, characterized by 
hot, dry summers and cool wet winters, supports a 
number of plant communities including oak wood- 
land, chaparral, and grassland (Griffin 1971; 
MacRoberts and MacRoberts 1976; Williams and 
Koenig 1980). Hastings Reservation has been pro- 
tected from fire and grazing since its establishment 
in the fall of 1937, except for a 40 ha horse pasture 
that was lightly grazed until 1968. 

Between 1944 and 1977, several plots were es- 
tablished at sites with a variety of land use histories 
(all within four km of each other) around Hastings 
Reservation to monitor native Nassella (including 
both N. pulchra and N. cernua) bunchgrasses (Ta- 
ble 1; detailed maps showing stand locations are 
available at Hastings). Original plots were estab- 
lished to monitor small patches of Nassella that re- 
mained, for whatever reason, in what was often a 
much larger expanse of introduced, annual grasses 
on abandoned fields or in oak savanna. Because 
data collection for these plots was not coordinated, 
available data differed for each plot. Data collected 
ranged from multiple censuses of size and location 
of every Nassella individual in a given plot, to sin- 
gle censuses indicating only presence or absence in 
a plot. The two species of Nassella that occur at 
Hastings Reservation are extremely similar in veg- 
etative morphology and are known to hybridize 
(Stebbins and Love 1941; Love 1954). Because 
many of the data sets for plots did not differentiate 
between these closely related species, we did not 
differentiate these species in our data analysis. All 
nomenclature follows Hickman (1993). 

We used data from nine plots (Table 1) that were 
established in four areas at Hastings Reservation: 
White Prairie (one plot), South Sandstone (one 
plot), North Field (five plots), and Arnold Field 
(two plots). White Prairie, South Sandstone, and 
North Field are all within a few hundred meters of 
each other (elevation ca. 550 m), and Arnold Field 
(elevation ca. 730 m) is located approximately four 
km SW of the other three areas. Nassella bunch- 
grasses (also known as tussock grasses or tufted 
grasses) are perennial grasses that have a clumped 
or cespitose growth habit. For all plots, Nassella 
individuals were defined as any physically distinct 
tussock that was not clearly a clonal fragment from 
some larger tussock (see e.g., Wilhalm 1995). 

An analysis of size structure of the Nassella pop- 
ulations was based on historical data or 1996 mea- 
surements of basal diameter measurements of in- 
dividuals. For many plots, historical data were tak- 
en from detailed tracings or maps of individual 
plants. Plants were divided into size classes based 
on basal diameters: (1) less than or equal to one 
cm, (2) greater than one cm to 5 cm, (3) greater 
than five cm to 10 cm, and (4) greater than 10 cm. 
Plants that were not circular were assigned to di- 
ameter classes based on the corresponding basal ar- 


276 MADRONO 


[Vol. 49 


TABLE |. SUMMARY INFORMATION FOR PLOTS USED IN THIS STUDY. 


Notes 


Plot area Date 
Plot name (m7) established 

White Prairie 10 1977 
South Sandstone 10 1966 
North Field 427 149 1966 
North Field 428 56 1966 
North Field 429 84 1966 
North Field 409 9 1965 
North Field 412 9 1964 
Arnold 420 9 1964 
Arnold 449 10 1979 


Considered to be undisturbed relict of bunchgrass prairie; 
moderate gopher activity 

Never cultivated; in a 40 ha area lightly grazed by 2—5 horses 
1940-1968 

Originally woodland; lightly cultivated for barley ca. 1860— 
1937 

Originally woodland; lightly cultivated for barley ca. 1860— 
O37 

Originally blue oak woodland; lightly cultivated for barley ca. 
1860-1937 

Sub-plot of North Field 429 

Originally valley oak savanna; cleared; cultivated as vineyard 
ca. 1920-1937 

Originally valley oak savanna; cleared; cultivated barley ca. 
1860-1937; many gophers 

Probably lightly cultivated 1860—1937; burned in 1979, many 
gophers 


eas. To estimate minimum longevity of Nassella in- 
dividuals, we used direct tagging of plants. We es- 
timated the age of large individuals by determining 
average rates of increase in basal area and calcu- 
lating the number of years required for an individ- 
ual to attain a given size. 

The plot in White Prairie was established in 
1977. This plot is surrounded by oak woodland and, 
as there is no record of land clearing, the original 
vegetation was probably grassland. White Prairie is 
considered to be a relict of pre-European Nassella 
bunchgrass grassland because it has not been 
cleared, and was probably only occasionally 
grazed. The plot showed evidence of moderate go- 
pher (Thomomys bottae) activity. In 1977, individ- 
ual Nassella plants were tagged, and, in many cas- 
es, wire loops were placed around the base of the 
plants to ensure future identification of individuals. 
Historical data include scale maps showing location 
and shape of each Nassella individual, as well as 
basal diameter measurements. In most cases Nas- 
sella individuals were generally elliptical or circu- 
lar, and, because of the detail of the maps, it was 
possible to identify individuals that were more ir- 
regular in shape. A digital image analysis system 
(Decagon Devices, Pullman Washington) was used 
to calculate basal area of irregular-shaped clumps. 
In the 1993 census, many tags could not be re- 
located; however, carefully drawn maps from 1977 
allowed us to identify most individuals. 

The plot in South Sandstone was established in 
1976. This plot is located adjacent to oak wood- 
land, and, like White Prairie, the original vegetation 
was probably grassland. The South Sandstone plot 
was never cleared or cultivated, but is located in a 
40-ha pasture that was lightly grazed by two to five 
horses until 1968. In 1976, individual Nassella 
plants were tagged using the same methods as pre- 
viously described. Historical data include scale 


maps showing location and shape of each Nassella 
individual, as well as basal diameter measurements. 
Almost all original tags in the South Sandstone plot 
were re-located in the 1993 census and, thus, tem- 
poral comparisons are always on the same individ- 
uals. A separate study using the South Sandstone 
plot counted Nassella seedlings in 1976, 1977, 
1978, and 1979. 

The five plots in North Field were established 
between 1964 and 1966 (Table 1), and are all lo- 
cated within about 60 m of each other. Before 1900, 
North Field was probably oak woodland dominated 
by Quercus douglasii (White 1966). Around 1900 
the trees were cleared, and the relatively level plots 
427, 428, and 429 (including 409, a subplot of 429) 
were cultivated for barley. Plots were located on 
clay-sand soils with rock outcrops. Although the 
field was cultivated for small grain production, only 
mule-drawn implements were used. Isolated, rocky 
outcrops where Nassella persisted were relatively 
undisturbed by the light cultivation equipment used 
during farming. North Field plot 412 occurs on a 
slope and was a vineyard. Data for North Field 
plots 427, 428, 429, and 409 from 1966 (1964 for 
plot 409) were taken from maps on which number 
and approximate location (but not size) of Nassella 
individuals were recorded. In the 1996 re-census, 
in order to calculate basal cover, two orthogonal 
diameters were recorded for every individual and 
basal area calculated (assuming an ellipse). Nassel- 
la individuals in North Field plot 412 were mapped 
over a period from 1964 through 1996. In 1977, 
1993, and 1996, basal area of each individual was 
also measured. All five North Field plots are in- 
cluded in a larger area that had been monitored 
since 1944. Available data from 1944 through 1964 
indicate only whether or not Nassella bunchgrasses 
were present in the larger area. 

The two plots in Arnold Field were established 


2002] 


HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 


TABLE 2. TOTAL BASAL AREA OF NASSELLA IN PLOTS AT HASTINGS RESERVATION (cm2?/m72). 


White South North 
Year Prairie Sandstone Field 412 
1964 e @ 9 
1976 @ 740 ® 
1977 1140 770 370 
1993 1190 1180 0) 
1996 e e 19 


in 1964 and 1979 (Table 1). Before 1937, these two 
plots were probably dominated by chaparral, since 
they occur near stands of Adenostoma fasciculatum 
Hook. & Arn., and 20% of Arnold 449 was covered 
with this shrub in the 1996 re-census. Plots in Ar- 
nold field area also on relatively shallow soils with 
rocks near the surface, and may not have been 
deeply tilled with the mule-drawn discs used at the 
time. Both Arnold plots show evidence of very high 
gopher activity. Historical data include scale draw- 
ings of Nassella individuals in Arnold 420 (used to 
obtain information on density and basal diameter of 
Nassella in 1964) and rough drawings for Arnold 
449 (used to calculate density). In the 1996 re-cen- 
sus, basal areas were calculated as for the other 
areas. 


RESULTS 


Basal area of Nassella showed different patterns 
of change in different plots (Table 2). The relict 
White Prairie plot was essentially constant over the 
16-year period from 1977 to 1993. Nassella at 
South Sandstone increased in basal area over this 


TABLE 3. 


277 
North North North Arnold 

Field 427 Field 428 Field 429 420 
6 * @ 400 

@ e ry @ 

@ @ ® @ 

@ @ € € 
250 310 290 670 


time period, achieving in 1993 a value essentially 
the same as White Prairie. There were insufficient 
data to draw conclusions about changes in total bas- 
al cover of Nassella in North Field, except for plot 
412 (former vineyard). In this case, Nassella dis- 
appeared between 1977 and 1993; however, in 
1996, a very small amount of Nassella was again 
found in this plot. In Arnold 420, Nassella basal 
area increased between 1964 and 1996. 

Density of Nassella individuals also showed dif- 
ferent patterns of change in different plots (Table 
3). The relict White Prairie plot had the highest 
density of all measured plots in 1977 and exhibited 
a slight increase with time. Mature Nassella indi- 
viduals at South Sandstone decreased by about 50% 
between 1976 and 1993, even though Nassella 
seedling recruitment pulses temporarily increased 
total Nassella density in 1978 and 1979. In North 
Field 412 (former vineyard), density of Nassella 
increased from 1951 until 1970, and then declined 
to zero by 1993. In 1996, three individuals were 
again found in this plot. In North Field plots 427 
and 428, Nassella density increased between 1966 


DENSITY OF NASSELLA (PLANTS/m?) FOR PLOTS AT HASTINGS RESERVATION. Missing entries indicate that plots 


were not yet established or were not measured in that year. * Density of mature individuals only is 12.2. ° Density of 


mature individuals only is 11.7. 


South 

White Sand- North North 
Year Prairie stone Field 412 Field 427 
1944 6 @ none none 
1945 ® ® none none 
1946 6 e none none 
1947 e e none none 
1951 e ® present present 
1964 ® ® 1.8 present 
1965 e e 1.9 ® 
1966 é e ® OF 
1969 6 ® 6.1 e 
1970 e e 7.8 e 
1972 e e 327, e 
1974 * ® 4.4 ® 
1976 e 12.8 @ e 
1977 13.8 i sey 5.4 ® 
1978 | he 60.4 e e 
1979 ® 48.0° e @ 
1984 ® 6 | e 
1991 e e 1 e 
1993 14.7 6.9 0 ® 
1996 @ e 0.3 2.9 


North 
North North Field Arnold Arnold 

Field 428 Field 429 409 420 449 
none present present e e 
none increase present e e 
none increase present e e 
none stable present e & 
none increase present e ® 
present increase 2.8 33 ® 
e 2.9 6 ® 
0.8 1.76 e e e 
® ® 3.1 @ & 
6 ® 4 a « 
e 6 é @ @ 
@ e 0.89 @ e 
§ « ® ® * 
@ e « « é 
6 6 e e 4 

® ® e « [2:5 

* ® « ® 2 Jo 
6 e @ e ® 
e ® ® « « 

3 2 2.4 12.3 6.7 


278 
White Prairie CI) 1977 
60 1993 
CI 1977 
60 1993 
40 
20 
0 
> 
S) 
= 
o 
= 
roy 
= 
Le 
got North Field 412 
got North Field Composite 
1996 
40 
20 
$1.0 1.0-5.0 5.0-10.0 >10.0 
Diameter size class (cm) 
Fic. 1. Size structure of Nassella populations for differ- 


ent stands at Hastings Reservation. Plots in North Field 
were measured only once. Size structures of North Field 
plots 427, 428 and 429 did not differ and thus the com- 
posite of these three plots is shown. 


and 1996. In North Field plot 428, Nassella density 
increased between 1944 and 1966, and although 
density in 1966 was very similar to that in 1996, 
measurement of the subplot of 428 (North Field 
429) indicated that there were fluctuations in den- 
sity over this time period. Nassella density in the 
two plots in Arnold Field changed in opposite di- 
rections. Arnold 420 had greater density in 1996 
than in 1964. Density in Arnold 449 decreased 
from 1978 to 1996. 

Size structure of Nassella populations varied be- 
tween plots (Fig. 1). The White Prairie population 
showed little difference in size structure between 
1977 and 1993 (Fig. 1). Numbers of individuals in 
the three larger size classes were approximately 
equal to each other, and about three times more 
numerous than individuals in the smallest size 
class. At South Sandstone, in 1977, the smallest 
size class and the largest size class contained 
roughly equal numbers of individuals, and the two 
intermediate size classes were only slightly less nu- 


MADRONO 


[Vol. 49 


100 South Sandstone 


White Prairie 


Percent mortality of given size class 
S 
© 


1.0-5.0 5.0-10.0 
Diameter size class (cm) in 1977 


=120 >10.0 


Fic. 2. Mortality as a percent of given size class for two 
Nassella populations at Hastings Reservation. 


merous. However, in 1993, size structure of the 
stand had come to resemble White Prairie: the 
smallest size class had disappeared entirely, num- 
bers of plants in the one to five cm size class had 
decreased significantly relative to 1977, and the 
larger two size classes increased in number. For 
North Field plots, it was not possible to determine 
changes in population size structure because we 
had only one observation for each plot. In 1977, 
North Field 412 (former vineyard) had the most 
Nassella individuals in the five to 10 cm size class, 
with fewer larger and smaller plants. Size structure 
in North Field 427, 428, and 429 in 1996 was very 
similar to that of White Prairie in 1993. Because 
size distributions of the three North Field plots 
were very similar, only a composite of these plots 
is shown. In 1964, Arnold 420 had most plants in 
the larger two size classes, with no plants in the 
smallest size class and very few in the one to five 
cm class. In 1996, the largest two size classes had 
decreased significantly, the smallest size class was 
represented, and the one to five cm size class was 
the largest. 

Mortality patterns were very similar in the two 
plots (White Prairie and South Sandstone) for 
which individuals could be identified over time 
(Fig. 2). Most mortality between 1977 and 1993 
was in the smallest size class, with decreasing per- 
centage mortality for larger size classes. At South 
Sandstone, total mortality in the three larger size 
classes combined was only eight individuals. White 
Prairie showed only slightly greater mortality in the 
three larger size classes. 

Seedling recruitment was sporadic in space and 
time (Table 4). No general pattern of recruitment 
connected to either yearly average rainfall or 


2002] 


HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 


279 


TABLE 4. NASSELLA SEEDLING RECRUITMENT (SEEDLINGS/m*) FOR PLOTS AT HASTINGS RESERVATION. Missing entries 
indicate that plots were not yet established, or were not measured in that year. 


White South North Field 
Year Prairie Sandstone 412 
1964 e ® 0) 
1965 ® ® 0.2 
1969 ® 8 40 
1970 e e 1.3 
1972 e e 0.2 
1974 e ® 0.1 
1976 ® | Remi 8 
1977 0.7 0.6 0.5 
1978 0.8 49.8 @ 
1979 e 34.1 @ 
1984 e e 0) 
199] ® r) 0 
1993 0.7 0.1 0) 
1996 e e 0 


monthly average rainfall was found. In years with 
seedling recruitment, seedling mortality was very 
high the first spring, and declined with time (Fig. 
3). At South Sandstone, eight of 131 small individ- 
uals noted in 1976 survived to 1993. Of the six 
Nassella seedlings noted in 1977, one survived to 
1993. Although individuals recruited after 1977 
were not followed between 1978 and 1993, 24 new 
individuals recruited into the South Sandstone pop- 
ulation. At White Prairie, a comparison between 
maps made in 1977 and 1993 indicates 50 new in- 
dividuals in the population. However, some of these 
are likely clonal fragments of previously existing 
individuals. 

Minimum longevity measurements for Nassella 
were made using tagged individuals. At White Prai- 
rie, 38 plants tagged in 1977 could be unequivo- 
cally identified in 1993. Of these 38 individuals, 22 
had increased in basal area over this 16-year period, 


500 aq 
iy 1978 cohort 
“E 400 | 
ro) 1979 cohort 
= es 
< | a 
& 300 l 
= 
= | 
o 
® 599 |. 1976 | 
S cohort 
Fs} | 
= 
3 100 | 
| 1977 cohort 
a 
1975 1980 1985 1990 1995 
Year 
Fic. 3. Seedling mortality over time for four cohorts of 


seedlings at Hastings Reservation South Sandstone plot. 


North Field North Field Arnold 
427, 428, 429 409 420 
e 0) O 
8 @) ® 
ad 0.2 ® 
e 1.6 e 
e ® ® 
@ O @ 
® ® 
® ® € 
© ® @ 
® @ © 
® ® ® 
® 8 ® 
© co) ® 
0.03 0.1 Lee 


indicating that they were still vigorous after this 
amount of time. At South Sandstone, 69 plants 
were identified as being more than 17 years old, 
and of these, 51 had increased in basal area. At 
North Field 428, 12 plants were staked in 1966 and 
one of these was still alive thirty years later. At 
Arnold 449, 18 tags were re-located after 17 years, 
five of which still had living Nassella individuals. 

Actual age of large Nassella individuals was es- 
timated using the average rate of increase of basal 
area over a 16-year period (Table 5). Due to large 
variation, long-term average growth rates were not 
significantly different between size classes or be- 
tween sites. The largest individuals at White Prairie 
and South Sandstone had basal areas between 400 
and 700 cm’. Using the average long-term growth 
rate for both sites and all size classes (5.0 cm/?/ 
year), these individuals were calculated to be 80 to 
140 years old. 

Accumulation of dead material in individual 
Nassella tussocks does not predict individual mor- 
tality. In 1976, maps of the South Sandstone plot 


TABLE 5. INCREASE IN BASAL AREA BY SIZE CLASS (cm?/ 
YEAR) OF NASSELLA IN PLOTS AT HASTINGS RESERVATION. 
Only plants that increased in size over time period are 
included. Data are averages + | standard deviation. Num- 
bers in parentheses are numbers of plants. 


Beginning White Prairie South Sandstone 

size class 1977 to 1993 1976 to 1993 
=1 cm none S.2232 2:8 68) 
(<=0.78 cm?) 


1 cm to 5 cm 4.2 + 2.9 (9) Sos2s5,0°(04) 


(0.78 to 19.6 cm?) 


5 cm to 10 cm 2STEUL ST (S) 4.7 + 6.4 (14) 
(19.6 to 78.5 cm?) 

>10 cm 8.5 + 6.8 (4) Sb a.) 
(>78 cm?) 


280 


indicated 24 individuals that had large regions of 
dead material. In 1993, only two of these individ- 
uals had experienced large declines in living basal 
area. Between 1976 and 1993, many individuals 
that initially seemed to be senescing or fragmenting 
rebuilt their tussocks and increased in living basal 
area. 


DISCUSSION 


Landscape-scale Dynamics of Nassella Without 
Fire or Grazing 


In general, Nassella bunchgrasses have increased 
at Hastings Reservation in the absence of fire and 
grazing. Both our data and those of White (1966) 
indicate that Nassella colonized new areas from 
1944 to 1966 (Table 3). As Nassella has spread to 
new areas, total average basal cover in established 
stands has remained stable. In 1967, average basal 
cover of Nassella for 13 protected stands at Has- 
tings Reservation was 10% (White 1967). For a 
subsample of these areas, we also found a total av- 
erage basal cover of 10% in 1977 (White Prairie 
and South Sandstone) and again in the 1990’s 
(White Prairie, South Sandstone, and Arnold Field). 
At finer resolution, we found a difference in plots 
that were historically cultivated and those that were 
not. In the 1990’s, average basal cover in formerly 
cultivated plots was only about 4%, compared to 
12% for plots that were never cultivated. The only 
location where Nassella has not maintained itself is 
in the former vineyard. 


Stand-scale Dynamics of Nassella 


Non-native annuals are present in varying levels 
all these stands (Stromberg and Griffin 1996) and 
by themselves, did not seem to be sufficient to 
cause stand declines. At White Prairie, where the 
only known disturbance was the historical intro- 
duction of non-native annuals, (1971-1991 cover of 
non-native, annuals was 33.1%) there was essen- 
tially no change in basal cover, density, or size dis- 
tribution of Nassella over a 16-year period of this 
study. It is possible that one or more of these mea- 
sures showed a transient initial reaction to non-na- 
tive annuals when they first became important in 
the 1800’s. However, at another Nassella grassland 
site in California, micro-fossil evidence suggests 
that density, at least, has not been affected by non- 
natives (Bartolome et al. 1986). 

Although non-native annuals did not appear to 
impact Nassella, even light grazing did cause sig- 
nificant changes in Nassella stands. Over the same 
period that the ungrazed Nassella stand at White 
Prairie was stable, the stand at South Sandstone, 
which had been formerly grazed, increased both in 
total basal cover and in changes in size distribu- 
tions. By 1993, total Nassella basal cover at South 
Sandstone was indistinguishable from the relict 
White Prairie plot. This suggests that a basal cover 


MADRONO 


[Vol. 49 


of around 1000 cm?/m* may be the maximum that 
can be supported in these stands at Hastings Res- 
ervation. Spacing of large individuals probably re- 
flects long-term competition for water in limiting 
years, and for small individuals, competition with 
alien annual species (Dyer and Rice 1997, 1999). 

Areas that were disturbed by soil cultivation sup- 
ported stands with lower total cover of Nassella, 
and had size distributions that are qualitatively dif- 
ferent from areas that were never cultivated. It is 
not clear whether cultivation was a disturbance that 
permanently altered the ability of areas to support 
Nassella stands (Stromberg and Kephart 1996), or 
whether recovery is simply extremely slow. How- 
ever, there is little indication that stands in formerly 
cultivated areas are developing toward patterns 
similar to never-cultivated plots. Again, the only 
plot where Nassella disappeared was the site of the 
former vineyard. 

It is possible that differences between cultivated 
and non-cultivated plots are due simply to pre-ex- 
isting site differences from before 1900. However, 
both the White Prairie plot and the South Sandstone 
plot are associated with oak woodland, as was 
North Field originally. Furthermore, another study 
that included 80 sites from around Hastings Res- 
ervation and the Carmel Valley also concluded that 
Nassella and a number of other native plant species 
(e.g., Poa secunda J.S. Pres] and Chlorogalum 
pomeridianum (DC.) Kunth) are rare in sites that 
have been subjected to historic cultivation (Strom- 
berg and Griffin 1996). 


Life-stage Mortality of Nassella in the Presence 
of Non-native Annuals 


We found that mortality was concentrated in the 
smaller size classes, and that initial seedling mor- 
tality was very high. Similar results were found in 
a study of individually marked N. pulchra in the 
San Juaquin valley of California (Marty 2002). Dif- 
ferential mortality of young plants is very common 
(Sarukhan et al. 1984 and references cited therein), 
and studies have also noted very high seedling mor- 
tality for Nassella (Bartolome and Gemmill 1981; 
Dyer et al. 1996). However, it is unclear both the 
degree to which interference (sensu Harper 1961; 
Muller 1969) is involved, and the relative impor- 
tance of inter- versus intraspecific interference in 
this mortality. In plots where density of mature 
Nassella individuals was low, interspecific interfer- 
ence from non-native annuals is probably a primary 
factor leading to high seedling mortality and lim- 
iting seedling recruitment. Studies have shown ex- 
plicitly that interference from non-native annuals is 
detrimental to performance of Nassella individuals 
(Nelson and Allen 1993; Dyer and Rice 1997; 
Hamilton et al. 1999). However, N. pulchra seed- 
lings can recruit into areas dominated by non-native 
annuals, although competition for soil moisture 
greatly reduced their growth (Hamilton et al. 1999). 


2002] 


The ability of Nassella or other native, perennial 
grass seedlings to thrive in soil dominated by non- 
native annuals may be influenced by soil microbial 
communities. Robinson (1971) found that Nassella 
seeds planted in Hastings soils dominated by Avena 
had significantly lower survival and growth com- 
pared to seeds grown in soils from relict stands of 
Nassella. Steenwerth (2002) found that Hastings 
old fields, as well as nearby recently tilled fields of 
the similar soil, had dramatically different micro- 
bial communities compared to similar soils in un- 
disturbed, relict Nassella stands. Indeed, a simple 
innoculum of soil from non-native grasslands ap- 
pears to inhibit the growth of native California 
grasses (Subramaniam et al. 2001). 

When basal cover of mature Nassella individuals 
is high, intraspecific interference may be more im- 
portant than interspecific competition as a factor in 
Nassella seedling mortality. High seedling recruit- 
ment appeared to be associated with low basal cov- 
er of mature Nassella individuals (despite presence 
of non-native annuals) and declined as Nassella 
density increased. For South Sandstone in particu- 
lar, as total basal cover of Nassella became similar 
to White Prairie, seedling recruitment densities be- 
came similar. Intraspecific competition for soil re- 
sources in established bunchgrass stands in semi- 
arid grasslands often causes seedling recruitment to 
be very low in the absence of disturbance (Aguilera 
and Lauenroth 1993b; Hook et al. 1994; Aguilera 
and Lauenroth 1995). 

It has been suggested that the bunchgrass growth 
form may be inherently associated with an increas- 
ing risk of death as tussocks increase in size (Harp- 
er 1977). This could be due to accumulation of 
plant litter in the tussock in situations where grasses 
are not subjected to periodic fire or grazing. Thus, 
lack of fire or grazing could lead to senescence of 
adult Nassella individuals (Menke 1992). This does 
not appear to be the case for established stands at 
Hastings Reservation, an inland dry grassland co- 
dominated by Nassella. Mortality of large individ- 
uals in plots that were never cultivated was very 
low, despite the presence of non-native annuals. For 
example, South Sandstone showed no mortality in 
the largest size class and individuals that appeared 
to be fragmenting demonstrated the ability to re- 
build the tussock. Indeed, tussock fragmentation is 
very common in perennial bunchgrasses (Wilhalm 
1995), and individual tussocks would be expected 
to possess the ability to rebuild through activation 
of dormant meristems (Bell 1984; White 1984). 
There was a small amount of mortality among large 
individuals at White Prairie, but this was probably 
due, at least in part, to gopher activity (Stromberg 
and Griffin 1996) and did not impact total Nassella 
cover. In contrast, there was evidence of mortality 
of large individuals in formerly cultivated plots. It 
is possible that in these areas altered site conditions 
reduce longevity of Nassella individuals or that in- 
terference from non-native annuals is more detri- 


HAMILTON ET AL.: LONG-TERM NASSELLA SURVIVAL 281 


mental in these locations. Gopher activity could 
also increase mortality of Nassella, and, while this 
might be important in Arnold Field, we observed 
little evidence of gophers in North Field. Responses 
to grazing in wetter, coastal California grasslands 
may be dramatically different (Stromberg et al. 
2001) and longevity and responses to management 
may vary between inland and coastal California 
Nassella grasslands. 

Contrary to some predictions (e.g., Menke 1992), 
tagged plants in our plots indicate that mature Nas- 
sella individuals are vigorous after as much as 30 
years of protection from grazing and burning. Age 
of clonal plants, such as Nassella, is impossible to 
measure directly (Stebbins 1950), so use of rates of 
clonal spread to estimate age is commonly em- 
ployed (e.g., Harberd 1961; Harberd 1962). Esti- 
mates of longevity of Nassella based on average 
growth rates indicate that large individuals are like- 
ly more than 100 years old. Because Nassella in- 
dividuals break up into clonal fragments, as do 
many bunchgrasses (e.g., Lord 1993; Samuel and 
Hart 1995; Wilhalm 1995), individuals may persist 
in a series of clonal fragments much longer than 
this. In fact, clonal fragmentation is an alternative 
mechanism for recruitment of ‘new’ individuals 
into populations (Crampton 1974; Lord 1993). Al- 
though growth rates for individual clones can de- 
pend strongly on intraspecific competition (Aguil- 
era and Lauenroth 1993a; Dyer and Rice 1997), 
interspecific competition (Nelson and Allen 1993; 
Dyer and Rice 1997; Hamilton et al. 1999), and 
genotype (Samuel and Hart 1995; Skalova et al. 
1997), our age estimate is not unusually high. Stud- 
ies of other species of bunchgrasses have also 
found that individuals can be very long-lived, with 
estimates ranging from 450 or 500 years or even 
longer in undisturbed areas (Coffin and Lauenroth 
1988: Lord 1993; Lauenroth et al. 1994). 


The Requirement for Management in 
Nassella Stands 


Information on long-term population dynamics 
of native perennial grasses that have been left un- 
disturbed by humans, is necessary to develop ef- 
fective prescriptions for restoration and manage- 
ment of areas with native perennial grass stands. 
Large protected areas, such as Hastings Reserva- 
tion, provide important reference systems in which 
to gather such information (Bock et al. 1993). Be- 
cause there are no unaltered pre-European grass- 
land communities remaining in California, there is 
no naturally occurring large model by which to es- 
tablish goals or measure success of management or 
restoration efforts. Models for restoration of Cali- 
fornia native grasslands depend on observations of 
species dynamics in small patches. Such small 
patches may have been the common expression of 
native California grasslands and large homogenous 
stands of native grasses were rare. However, since 


282 


pre-European California grasslands probably con- 
tained many annual and perennial grasses and forbs 
(Heady 1977; Heady et al. 1992), one possible goal 
would be to promote overall diversity of native 
grassland species. This is complex because some 
management practices that could potentially benefit 
Nassella appear to be detrimental to other native 
bunchgrass species (Dennis 1989). Therefore, lack- 
ing detailed, species-specific information on the ef- 
fects of potential management practices, landscape- 
scale management should be conservative. Because 
Nassella bunchgrasses that have been protected 
from grazing and burning show no signs of disap- 
pearing at the landscape scale at Hastings Reser- 
vation (despite the introduction of non-native an- 
nuals), landscape-scale management to maintain 
relict Nassella in the landscape appears to be un- 
necessary. 

Stand-scale population dynamics are site depen- 
dent and appear to be related to land-use history; 
stand-scale management should take this into ac- 
count. At Hastings Reservation, some of the bunch- 
grass stands probably pre-date European arrival, 
while other stands occur in sites that have been 
converted from other vegetation types such as oak 
woodland or chaparral. This is true for other areas 
of California as well (Huenneke 1989; Keeley 
1993; Hamilton 1998). In areas that have never 
been cultivated, interference from non-native an- 
nuals does not seem to cause Nassella stand de- 
cline. Despite high seedling mortality and sporadic 
seedling recruitment, areas that were never culti- 
vated can have high basal cover of Nassella and 
some achieve replacement levels of recruitment. 
Our longevity estimates, and the demonstrated abil- 
ity of Nassella to repair dead portions of a tussock, 
suggest that many large Nassella individuals found 
in areas such as White Prairie and South Sandstone 
could have been present at Hastings Reservation 
before the area was first homesteaded in 1863 
(White 1967). For such long-lived species, even if 
conditions allowing for seedling recruitment are 
quite rare, stands can achieve replacement recruit- 
ment (Noble 1986; Lauenroth et al. 1994). Thus, 
lack of seedlings is not necessarily an indication of 
future stand decline. Nassella stands in areas that 
have not been disturbed by cultivation do not ap- 
pear to require management for maintenance. 

Even light grazing appears to greatly reduce Nas- 
sella basal cover. Although Nassella can sometimes 
persist in grazed areas (White 1967; Stromberg and 
Griffin 1996), intensive, year-round grazing on in- 
land stands seems to have the potential to be det- 
rimental to stand persistence. More studies are re- 
quired before decisions can be made concerning the 
compatibility of Nassella with grazers, or the use 
of grazing as a management tool as some have sug- 
gested (e.g., Menke 1992). 

Cultivation appears to be the most detrimental 
disturbance, and left alone, Nassella seems to re- 
cover extremely slowly (if at all) from a distur- 


MADRONO 


[Vol. 49 


bance of this type. Other studies have come to a 
similar conclusion (Stromberg and Griffin 1996). 
Nassella stands that were historically cultivated 
have low basal cover and do not appear to be de- 
veloping toward the condition of stands that were 
not subjected to cultivation (e.g., Fig. 2). Further- 
more, Nassella in at least one population in a his- 
torically cultivated stand has declined significantly. 
At Hastings Reservation, some of the Nassella 
stands that were historically cultivated occur in ar- 
eas that have been converted to grassland from oth- 
er vegetation types. In such cases, the appropriate 
management goal may be simply to promote Nas- 
sella. In formerly cultivated areas, there are few 
large individuals, and this suggests that Nassella 
may be shorter-lived in these areas than in areas 
that were never cultivated. Low basal cover in areas 
subjected to historic cultivation may be the result 
of seedling recruitment not being frequent enough 
for stand replacement or growth. Therefore, in areas 
with historical cultivation, active management to 
increase seedling survivorship, while not harming 
mature plants, may be required to increase abun- 
dance of Nassella. 


ACKNOWLEDGMENTS 


The UC Hastings Reservation made this long-term 
work possible. We thank Thomas Cate and Laura Rosen- 
feld for field assistance. We are grateful to J. R. Haller, 
Claus Holzapfel, Bruce Mahall, Elizabeth Painter, Laura 
Rosenfeld, Jochen Schenk, Ed Schneider, and Josh Schi- 
mel for providing helpful comments on the manuscript. 
We gratefully acknowledge the financial assistance pro- 
vided by the Andrew W. Mellon Foundation, and a UCSB 
General Affiliates Graduate Dissertation Fellowship. 


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MApRONO, Vol. 49, No. 4, pp. 285-288, 2002 


A NEW SPECIES OF PRUNUS (ROSACEAE) FROM THE MOJAVE DESERT 
OF CALIFORNIA 


BARRY A. PRIGGE 
Herbarium—Botanical Garden, University of California, 
Los Angeles, CA 90095-1606 
bprigge @ucla.edu 


ABSTRACT 


Prunus eremophila (subgenus Emplectocladus Torr.) is described and illustrated from the southern 
Mojave Desert. It is closely related to P. havardii (W. Wight) S. C. Mason but differs from it in pubescence 


of leaves and larger size of floral parts and fruit. 


Key Words: Prunus, Prunus eremophila, Rosaceae, Amygdaloideae. 


During vegetation surveys of the Mojave Desert 
by the Bureau of Land Management (BLM), some 
unusual specimens of Prunus were collected by 
BLM botanists. After reviewing specimens of other 
woolly fruited species of Prunus at LA, RSA, GH, 
and UC and borrowing specimens of P. havardii 
from SRSC, it is concluded that they merit recog- 
nition as a new species. 


Prunus eremophila Prigge, sp. nov. (Fig. 1).— 
TYPE: USA. California, San Bernardino Co. 
12.8 mi NNW of Goffs on Lanfair Road and 0.8 
mi E on road to True Blue Mine, 1145 m elev., 
19 March 1992, Prigge 9825 (Holotype: RSA; 
Isotypes: CAS, GH, LA, MO, NY, TEX, UC). 


Pruno havardii similis sed differt foliis villosis 
(non glabris vel subtiliter pubescentibus), hypanthio 
3—6 mm longo (vs. 2.0—2.5 mm), sepalis 1.0—1.6 
mm longis (vs. 0.5—1.0 mm), petalis 4—7 mm longis 
(non 2 mm longis vel carentibus), fructu lanato 
(non pubescenti vel canescenti) 10-16 mm long 
(vs. 7-10 mm) 9-11 mm lato (vs. 8—9 mm), et en- 
docarpio 9-14 mm longo (vs. 6—9 mm). 

Dioecious, + globose to widely spreading, intri- 
cately branched shrubs to 2.2 m tall; outer bark on 
older stems gray glaucous becoming reddish 
brown; inner bark orange; branchlets grayish pu- 
bescent, weakly thorny but without sharp, indurated 
tips; internodes 3—14 mm long; short-shoot spurs 
1.5—5 mm long. Leaves conduplicate in bud, spath- 
ulate to ovate, 5-20 mm long and 2—8 mm wide on 
short shoots, 13—30 mm long and 7-19 mm wide 
on long shoots, serrate (rarely entire or wavy) with 
a total of O—13 teeth that are often asymmetrically 
disposed, sparsely villous on both surfaces, cuneate 
at base, acute, round, truncate, or retuse, often mu- 
cronulate at apex, lacking stomata on adaxial sur- 
face. Petioles 0.5—3.0 mm long on short shoot 
leaves, 2.0-5.0 mm long on long shoot leaves. 
Flowers axillary with 2—4 flowers per leaf axil, pre- 
cocious, unisexual by abortion of either stamens or 
pistil. Hypanthium turbinate, pubescent externally, 


strigose internally in male flowers, glabrous inter- 
nally in female flowers. Sepals 5, 1.1—1.6 mm long, 
1.4-2.3 mm wide, deltate, externally pubescent. 
Petals 5, white, 2.7-5.8 mm long, 2.2—4.0 mm 
wide, round to spathulate, slightly narrower and 
more acute in male flowers. Stamens 10—15, in 3 
whorls of 5 stamens each or fewer by abortion, the 
uppermost whorl opposite the calyx lobes and orig- 
inating on the hypanthium rim, the middle whorl 
opposite petals, the lowermost whorl + opposite 
the calyx lobes, with the lower two whorls of sta- 
mens arising from wall of hypanthium; filaments 
1—2.6 mm long, white; anthers 0.7—1.2 mm long, 
0.7—1.0 mm wide, light yellow. Pistil one (but oc- 
casionally two and then connate at the ovary), 4.8— 
6.0 mm long, the ovary 1.6—2.5 mm long, 1.5—1.8 
mm in diameter, pubescent, the style 3.2—3.7 mm 
long. Fruit drupaceous, slightly laterally com- 
pressed ovoid to orbicular, (9—)11—16 mm long, 
(6—)8—11(—14) mm wide along major axis, (6—)7— 
10(—14) mm along minor axis, densely villous with 
grayish or rusty hairs; sarcocarp (exocarp + me- 
socarp) very slightly fleshy, ca 0.5 mm thick, easily 
separating from pit in fertile fruits but generally not 
split along the ventral suture, exocarp apricot color 
when ripe. Endocarp smooth, apiculate, and ven- 
trally ridged, 9-15 mm long, 6—10 mm wide. 


Paratypes. U.S.A. California. San Bernardino 
Co.: 12.8 road mi NNW of Goffs along Lanfair 
Road, then 0.8 mi E on trail-road toward Vontrigger 
Hills, 1160 m, 19 March 1992, Henrickson 22633 
(CSLA); southern Lanfair Valley E of Bobcat Hills, 
0.8 mi NE of True Blue Mine, 1145 m, 21 Mar 
1980, Prigge 3634 (LA); Vontrigger Hills, 12.8 mi 
NNW of Goffs on Lanfair Road and ca. 1.95 air 
mi E of junction of Lanfair Rd and road to True 
Blue Mine, 1155 m, 3 Apr 1985, Prigge 6235 (LA, 
RSA); (same area) 7 May 1985, Prigge, Thompson, 
and Davis 6374 (NY, TEX, UC, RSA, CAS, GH); 
(same area) 19 March 1992, Prigge 9826 (TEX, 
RSA); (same area and date), Henrickson 22632 
(CSLA); Lanfair Valley, 12.9 air mi N of Goffs 


286 MADRONO [Vol. 49 


WANs 
ZZ 
! 


MD 
WZ 


Fic. 1. Prunus eremophila: A) fruiting branch with short-shoot leaves (Prigge 6372); B) branch with long-shoot 
leaves and stipule; C) short-shoot leaves; D) male flower—early anthesis (left), dissected flower with rudimentary pistil 
(right), and petal (above); E) dissected male flower with two rudimentary pistils (abnormal); F) terminal branch with 
female flowers and rudimentary stamens; G) dissected female flower with rudimentary stamens; H) endocarp and fruit. 


where Old Government Road crosses Sacramento 
Wash, 1165 m, 13 May 1992, Prigge and Henrick- 
son 10220 (RSA). 

Prunus eremophila occurs primarily in washes 
but also on rocky slopes in the higher zones of the 
creosote bush series and the transition zone from 


creosote bush to the blackbush series where there 
are scattered Joshua trees (Sawyer and Keeler-Wolf 
1995). Substrates are derived from igneous rocks 
(granites and rhyolites). Common associates are Te- 
tradymia_ stenolepis, Ericameria cooperi, Yucca 
brevifolia, Y. schidigera, Acamptopappus sphaero- 


2002] 


115° 10' 


PRIGGE: PRUNUS EREMOPHILA 


287 


115° 10' 


LANFAIR VALLEY 


FENNER VALLEY 


| 
to Goffs 10 km 


Fic. 2. 


cephalus, Eriogonum fasciculatum, Larrea triden- 
tata, Salazaria mexicana, Yucca baccata, Lycium 
oligosperma, Krameria erecta, and Xylorhiza tor- 
tifolia (nomenclature follows Hickman 1993). Ele- 
vation ranges from 975-1175 m (3200-3850 ft). 
The species is known only from the vicinity of the 
Vontrigger Hills, Bob Cat Hills, southern Lanfair 
Valley, and one outlier in northeastern Fenner Val- 
ley of southeastern San Bernardino County (Fig. 2). 
Flowering time is from mid March to early April. 

Prunus eremophila is in the subgenus Emplec- 
tocladus Torr. (commonly included in subgenus 
Amygdalus L.) as envisioned by Mason (1913). The 
subgenus is characterized by dioecy through abor- 
tion of stamens or pistils, a fine pubescence on the 
inside of the hypanthium, stamen number reduced 
to 10 or 15 with some filaments inserted on the 
hypanthial wall, and with or without stomata on the 
upper leaf surface (Emplectocladus and Prunus are 
the only subgenera of Prunus that have species 
without stomata on the upper leaf sufaces). All 
these traits, including the absence of adaxial leaf 
stomata, occur in P. eremophila. When Mason 
studied this subgenus, only four species (all North 
American) were known: Prunus fasciculata (Torr.) 
Gray, P. microphylla (H.B.K.) Hemsl., P. minuti- 
flora Englem., and P. havardii. With the description 
of P. cercocarpifolia from the Chihuahuan Desert 
region by Villarreal (1989) and now P. eremophila, 
the subgenus now consists of six species. 

A comparison of vegetative, floral, and fruit 
characters among the species of Emplectocladus 
point to a possible close relationship between Pru- 


Known distribution of Prunus eremophila (@) in eastern San Bernardino County, California. 


nus eremophila and P. havardii of the Chihuahuan 
Desert of southwest Texas and northern Mexico 
and, to a lesser degree, with P. microphylla of cen- 
tral Mexico (states of San Luis Potosi, Hidalgo, 
México, and District Federal). Prunus havardii is 
thorny and has glabrous or sparsely pubescent 
leaves, and P. eremophila is only rigidly branched, 
not truly thorny and has villous leaves. The larger 
flower size (longer hypanthium length, longer pet- 
als, stamens, and pistils) and slightly larger fruit 
size of P. eremophila are distinctive within the sub- 
genus Emplectocladus indicating that P. eremophi- 
la is distinctive and recognizable as a distinct spe- 
cles. 

Prunus eremophila is sympatric with P. fasci- 
culata but flowering time overlaps only partially. 
The former begins flowering about two to three 
weeks earlier than the latter, so that P. eremophila 
has almost completed flowering by the time P. fas- 
ciculata begins. There is a slight overlap in flow- 
ering (perhaps as much as a week) when cross pol- 
lination could occur between these two sympatric 
species, but no hybrids were found suggesting that 
additional isolating mechanisms other than seasonal 
ones are present. 

At the present, the species does not appear to be 
threatened, but any species with such a limited dis- 
tribution and small size (perhaps ca. 2000 individ- 
uals) could become threatened fairly quickly from 
off-road vehicle (ORV) use, grazing pressure, and 
mining activities or global warming. All known 
populations are within the recently formed East 
Mojave National Preserve, and the Preserve should 


288 


minimize or eliminate most threats from human ac- 
tivities. Future ecological and physiological studies 
to determine the population size and structure, seed 
germination requirements, seedling survival, re- 
cruitment, and threats would provide valuable data 
for evaluating the long term survivability of this 
species. 

Prunus eremophila can be identified by making 
the following modification to the key in The Jepson 
Manual (Hickman 1993): 


5’ Twigs rigid; ovary and fruit generally densely pu- 
berulent. 

9. Flowers perfect; leaf blades glabrous (some- 
times minutely puberulent on petiole, midrib, 
and basal margin in P. fremontii) and more 
than 3 mm wide. 

10. Leaf blades ovate to round, 7—22 mm 
wide, base obtuse to + cordate; petals 
WnGer sae. et eye a ee oe P. fremontii 

10’ Leaf blades elliptic to oblanceolate, base 
tapered; petals reddish .... P. andersonii 

9’ Flowers imperfect; leaf blades glabrous to vil- 
lous, if glabrous the leaf blades less than 3 
mm wide. 

11. Leaf blades spathulate, 1-3 mm wide; 
puberulent or glabrous .... P. fasciculata 
12. Leaf surface puberulent; inland ... 

var. fasciculata 


eh Ge) jem ce uleh se) lel se” jel *s¥ fen ce] fe; lelurstiae 


MADRONO 


[Vol. 49 


12’ Leaf surface glabrous to low-papil- 
latesycoastall ea ae var. punctata 

11’ Leaf blades ovate, 2-19 mm wide, 
sparsely villous P. eremophila 


ACKNOWLEDGMENTS 


I thank Hyrum Johnson for bringing this plant to my 
attention, Henry J. Thompson, Stan Davis, and James 
Henrickson for their help and advise on this paper and 
their company on field trips, Guy Nesom for the Latin 
discription, Bobbi Angell for the illustration, the curator 
of the Sul Ross Herbarium for the loan of Prunus havar- 
dii, the curators of GH and UC/JEPS for permitting me to 
examine the specimens in their herbarium, and the review- 
ers (J. Henrickson and Richard Lis) for constructive and 
helpful comments. 


LITERATURE CITED 


HICKMAN, J. C. (ed.). 1993. The Jepson manual: higher 
plants of California. University California Press, 
Berkeley, CA. 

Mason, S. C. 1913. The pubescent-fruited species of Pru- 
nus of the southwestern states. Journal of Agricultural 
Research 1:147—177. 

SAWYER, J. O. AND T. KEELER-WoLFr. 1995. A manual of 
California vegetation. California Native Plant Socie- 
ty, Sacramento, CA. 

VILLARREAL Q., J. A. 1989. A new species of Prunus sub- 
genus Amygdalus (Rosaceae) from Coahuila, México. 
Sida 13:273—275. 


Maprono, Vol. 49, No. 4, pp. 289-294, 2002 


A NEW CEANOTAUS (RHAMNACEAE) SPECIES FROM NORTHERN 
BAJA CALIFORNIA, MEXICO 


STEVE BOYD 
Herbarium, Rancho Santa Ana Botanic Garden, 1500 N. College Ave., 
Claremont, CA 91711 
Steve.Boyd@cgu.edu 


JON E. KEELEY 
Herbarium, Rancho Santa Ana Botanic Garden, 1500 N. College Ave., 
Claremont, CA 91711 
U.S. Geological Survey, Western Ecological Research Center, 
Sequoia Field Station, Three Rivers, CA 93271 
Department of Organismic Biology, Ecology and Evolution, 
University of California, Los Angeles, CA 90095 


ABSTRACT 


Ceanothus bolensis S. Boyd & J. Keeley is a new species in the subgenus Cerastes from northwestern 
Baja California, Mexico. It is well represented at elevations above 1000 m on Cerro Bola, a basaltic peak 
approximately 35 km south of the U.S./Mexican border. It is characterized by small, obovate to oblan- 
ceolate, cupped, essentially glabrous leaves with sparsely toothed margins, pale blue flowers, and globose 
fruits lacking horns. Principal components analysis on morphological traits shows it to be distinct from 
other members of Cerastes which are distributed away from the coast in southern California and Baja 
California, Mexico. These phenetic comparisons also suggest that Ceanothus otayensis should not be 
subsumed under C. crassifolius, as treated in the Jepson Manual, but rather should be retained at specific 
rank as well. 


RESUMEN 


Ceanothus bolensis S. Boyd & J. Keeley es una nueva especie del subgénero Cerastes en el noroeste 
de Baja California, México. Esta bien representada en altitudes por arriba de los 1000 m en el Cerro 
Bola, un pico basdltico, apr6ximadamente a 35 km de la frontera de USA/México. Esta se caracteriza 
por hojas pequenas, obovadas a oblanceoladas, convexas, esencialmente glabras y con margenes espar- 
cidamente dentados, flores azul palido y frutos globosos sin corniculos. El andlisis de componentes prin- 
cipales con caracteres morfoldgicos la presenta como distinta de otros miembros de Cerastes distribuidos 
lejos de la costa en el sur de California y de Baja California, México. La comparaci6on fenética también 
sugiere que Ceanothus otayensis no debe ser incluida dentro de C. crassifolius como en el Manual de 
Jepson, sino mas bien deberia ser retenida a nivel especie. 


Key Words: Ceanothus, subgenus Cerastes, Baja California, Mexico, phenetic analysis, endemic. 


Ceanothus is a diverse genus of shrubs with the 
center of species diversity in the Mediterranean-cli- 
mate California Floristic Province (Schmitt 1993: 
source for nomenclature, except where indicated). 
It comprises approximately 60 species (van Rens- 
salaer and McMinn 1942) more or less equally di- 
vided into two clades that have long been system- 
atically recognized as subgenera Ceanothus and 
Cerastes (Harding et al. 2000). 

Recent collections of a Ceanothus from north- 
ern Baja California, Mexico suggest a new taxon 
that is worthy of recognition. These collections 
are from Cerro Bola, a basaltic mountain, ap- 
proximately 35 km south of the Tecate border 
crossing. At elevations between 1000 m and the 
peak at 1290 m, a small-leaved, erect, divaricate- 
ly branched Ceanothus species in the subgenus 
Cerastes is frequent in chaparral dominated by 


Arctostaphylos glandulosa Eastw. ssp. adamsii 
(Munz) Munz and Chamaebatia australis (Bran- 
degee) Abrams. It is less frequent on lower slopes 
from at least 500 m elevation. This Ceanothus 
shares features with several other Cerastes spe- 
cies distributed in southern California and adja- 
cent Baja California, Mexico. The lack of fruiting 
horns and deeply concave, toothed leaves sug- 
gests an affinity with Ceanothus greggii A. Gray 
var. perplexans (Trel.) Jepson and with the rela- 
tively recently described C. ophiochilus Boyd, 
Ross, & Arnseth (Boyd et al. 1991), a rare en- 
demic in southwestern Riverside County. The 
small leaf size of the Cerro Bola plants is similar 
to C. ophiochilus and C. otayensis McMinn (van 
Renssalaer and McMinn 1942), a localized en- 
demic on two mountain peaks in southern San 
Diego County, California and immediately adja- 


290 MADRONO 


cent Baja California, ca. 30—35 km northeast of 
Cerro Bola. In the most recent floristic treatment 
C. otayensis is treated as a hybrid derivative of 
C. crassifolius Torr., a widespread species away 
from the coast in southern California and Baja 
California, and C. greggii var. perplexans, the 
principle Cerastes species from the interior 
slopes of the Peninsular Ranges (Schmitt 1993). 


METHODS 


Herbarium specimens of the Cerro Bola taxon 
and other Ceanothus species in subgenus Cerastes 
from southern California were used for phenetic 
comparisons. Because fruit morphology has more 
distinguishing characters than flowers, only fruiting 
specimens were selected. Species used for compar- 
ison were Ceanothus greggii var. perplexans, C. 
greggil var. vestitus (Greene) McMinn, C. ophioch- 
ilus, C. otayensis, and C. crassifolius. 

We selected 14 characters for analysis; 6 con- 
tinuous quantitative, 7 qualitative and 1 calculated 
ratio (Table 1). For quantitative characters, two 
samples were measured for each specimen and the 
mean was used in the analysis. For qualitative 
characters, characteristics were given a relative 
score from 1 to 5. All character states were stan- 
dardized as z-scores by subtracting each observa- 
tion from the mean of all individuals, and dividing 
by the standard deviation. A species matrix of 
these scores was used for ordination with principal 
components analysis using SYSTAT 5.05 (Evans- 
ton, IL). 


RESULTS 


Means values for phenetic characters for all taxa 
discussed above are shown in Table 1. The Cerro 
Bola specimens have the smallest leaves but they 
are not significantly different from the other two 
small-leaved taxa, C. ophiochilus and C. otayensis. 
Cerro Bola plants are similar to C. ophiochilus in 
their very deeply concave leaves, limited pubes- 
cence, and smaller fruits that lack horns, however, 
these two taxa differ in their leaf shape, reflected 
in the leaf length/width ratio. The broad leaves of 
Cerro Bola plants are quite unlike the nearly linear 
leaves of C. ophiochilus, which generates a length/ 
width ratio roughly double that of all other taxa. 
The low apical angle on leaves of Cerro Bola plants 
is quite unlike C. ophiochilus but similar to C. otay- 
ensis and C. greggii var. perplexans. Ceanothus 
otayensis separates from all other taxa, except C. 
crassifolius, in having revolute leaves, well devel- 
oped, often brownish pubescence on branchlets and 
undersides of leaves. 

The principal components analysis for all taxa 
(Fig. 1), explained 50% of the total variance on the 
factor 1 axis and 20% on the factor 2 axis. Cea- 
nothus crassifolius was widely separated from the 
other taxa on the factor 1 axis; the most important 
components being convex leaves, revolute margins, 


LEAF AND FRUIT COMPARISON OF CEANOTHUS BOLENSIS AND OTHER SPECIES IN SUBG. CERASTES (X + SD). 


TABLE 1. 


C. greggii 
var. vestitus 


C. greggii vat. 


C. otayensis C. crassifolius 


C. ophiochilus 


perplexans 


C. bolensis 


Character 


10 


23.0 + 4.4 
WAS 2 2 


12 
Ply ee eg 
Sid) ee wile? 


12 
Bo) 25 OL 


26 
49+ 0.8 
39) ae (0),7/ 


n 


1.9 + 0.3 


1.4 + 0.2 
32 2 (0,5) 
3.8 + 0.8 
3,5) ae (04! 
522i SG 


Of aes 


to 22 OLS) 
32) Be il 


10.1 + 2.4 
Seo) 28) Wnts 
1.8 + 0.4 
358) SE (08) 
10+0 
le} Be iol 

Sia 22 (ils 

54.6 + 17.3 


14.7 + 2.6 
2 22 OJ 
2.9 + 0.6 


WA, = 9) 


1.3 2 O2 
4.3 + 0.7 
10+ 0 
3.4 + 0.7 
48 + 5.1 
50.1 + 13.6 


Leaf length (mm) 
Leaf width (mm) 
Length/width ratio 


1.0 + 0.0 

4.6 + 0.7 

Pigs oe Ilsil 
2S)S) 22 115).3 
SOON 


4.5 + 0.5 


extreme) 
extreme) 


Leaves concave (1 


1.0 + 0.0 


Io 26 @2 


3.8 + 0.6 
6.7 + 4.3 
ilo) 22 AO 


no, 5 = 
none, 5 


Leaves revolute (1 = 
Teeth on leaves (1 


53) 22 (0)07/ 
DY) BET 3} 


many) 


Wifes) 2a) Ahn 


Pubescence on branchlets 


Apical angle of leaves (°) 
Basal angle of leaves (°) 


Boo} 26 (0),2) 3.6 + 0.5 2M 28 (0).3) 44+ 0.5 D0) 22 OO 


2.6 + 0.5 


dense) 


glabrous, 5 = 


él 


Pubescence on leaf abaxial side 


[Vol. 49 


Le 2e 10) 2s] 0,5 1.0 + 0.0 4.3 + 0.5 49+ 0.3 


1.0 + 0.0 


glabrous, 5 = dense) 


ad 


2002] BOYD AND KEELEY: 


Factor 2 


-1.5 -1 -.5 0 y 1 
Factor 1 


CEANOTHUS BOLENSIS 


291 


C. bolensis 

C. crassifolius 

C. greggii perplexans 
C. g. vestitus 

C. ophiochilus 

C. otayensis 


BOS ae © 


co 2 


Fic. 1. Principal components analysis of Ceanothus crassifolius, C. greggii var. perplexans, C. greggii var. vestitus, 
C. otayensis, C. ophiochilus, and C. bolensis. Factor loading scores are in Table 2. 


leaf length, pubescence characters and fruit horns 
(Table 2). Ceanothus otayensis was clearly sepa- 
rated from C. crassifolius on both the factor 1 and 
factor 2 axes but overlapped with C. greggii var. 
perplexans (Fig. 1). 

Taking C. crassifolius out of the analysis gave 
greater resolution to the separation of C. otayensis 
and C. greggii var. perplexans (Fig. 2). In this 
analysis, factors 1 and 2 explained 36% and 21%, 
respectively of the total variance. Ceanothus otay- 
ensis WaS most prominently separated along the 
factor 1 axis, where pubescence characters and de- 
gree of leaf concavity were among the important 
components. On the factor 1 axis the Cerro Bola 
plants were distinctly intermediate to C. otayensis 
and C. ophiochilus. Cerro Bola plants were dis- 
tinctly separated from C. greggii varieties on the 
factor 2 axis, where the important components 
were pedicel length, fruit horns, leaf length and 
apical angle (Table 2). Clearly there is a sound 
morphological basis for the recognition of the Cer- 


TABLE 2. 
FIGURES 1 AND 2. 


ro Bola taxon (Figs. 1, 2), treated here as a new 
species. 


SPECIES TREATMENT 


Ceanothus bolensis S. Boyd and J. Keeley, sp. 
nov. (Fig. 3)—-TYPE: MEXICO, Baja California, 
chaparral with Chamaebatia australis on NE side 
of Cerro Bola, S of Tecate, elevation 1000 m, 26 
Apr 1996, Jon E. Keeley 27233 (RSA). 


Differt a C. ophiochilus foliis late obovatis vel 
late oblanceolatis; a C. otayensis foliis glabris, non 
concavis, marginibus revolutis; a C. greggii var. 
perplexans foliis glabris, parvulis (sub 6 mm lon- 
gis). 

Erect, divaricately branched shrub, 1—1.5 m tall, 
lacking basal burl and not resprouting after top- 
killed. Older stems ashy gray, intricately branched 
with rigid diverging branches. Younger branches 
reddish gray and lightly puberulent. Stipules thick- 


CHARACTERS USED IN PHENETIC ANALYSIS AND FACTOR LOADINGS FOR PRINCIPAL COMPONENTS ANALYSIS FOR 


Figure | Figure 2 

Character Factor 1 Factor 2 Factor 1 Factor 2 
Leaf length 0.88 0.04 0.64 0.60 
Leaf width 0.78 = | Pee 0.71 0.30 
Length/width ratio —0.08 0.86 —0.504 0.460 
Leaves revolute 0.83 —0.06 0.66 —0.39 
Leaves concave Sos 0.04 —0.78 —0.24 
Teeth on leaves 0.11 —0.84 0.52 ae 
Apical angle of leaves 0.35 OTF ==) Wes, 0.73 
Basal angle of leaves =0.22 0.70 —0.44 0.17 
Pubescence on branchlets 0.88 —0.17 0.84 —0.10 
Pubescence on bottom of leaves 0.88 —0.03 0.77 —0.11 
Pubescence includes brown hairs 0.78 tS 0.66 —0.41 
Pedicel length 0.67 0.31 0.36 0.81 
Fruit width 0.76 0.13 0.42 0.17 
Presence of fruit horns 0.82 0.27 0.49 0.63 


292 


Factor 2 


-3 -2.5 -2 -1.5 -1 -5 O .5 
Factor 1 


RiGs 2: 
1. Factor loading scores are in Table 2. 


ened and persistent, waxy or corky, dark reddish 
brown to purplish black. Leaves evergreen, thick, 
coriaceous, yellowish green, opposite, often clus- 
tered on short axillary spur branches; petioles 0.5— 
1 mm long, + 0.4 mm wide, minutely puberulent; 
blades broadly obovate to oblanceolate, deeply con- 
cave, (2.5)4—6(10) mm long and (3)4—7(9) mm 
wide, surfaces sparsely puberulent when young, 
glabrate in age, margins not revolute, sharply den- 
tate distally with (1)2—3 pairs of lateral teeth and 
an apical tooth; midvein prominent abaxially, lat- 
eral veins obscure, 3—5 pairs. Inflorescence a sub- 
umbellate axillary raceme, peduncle + 2 mm long, 
densely short puberulent, bearing 6—8 flowers. Ca- 
lyx, including receptacular disk + 5 mm wide at 
anthesis, lobes 5, pale blue, fading cream-white, 
1.8—2 mm long, deltoid to ovate, apex acute. Petals 
5, pale blue, fading cream-white, 1.8—2 mm long, 
ladle-shaped, + equally divided into deeply saccate 
distal blade and filiform proximal claw. Stamens 5, 
filament 1.8—2 mm long, anther + 0.4 mm. Ovary 
with style 1.8—2 mm long, 3-lobed, the lobes + 0.4 
mm; fruit a globose to depressed globose capsule, 
dark green to reddish, 3—4 mm diameter, smooth, 
lacking apical horns, lateral valve crests absent or 
vestigial. 


Distribution. At present, Ceanothus bolensis is 
known only from the mid- to upper slopes of Cer- 
ro Bola (500 m), where it is locally common in 
the chaparral vegetation. To our knowledge, no 
other member of subgenus Cerastes is found on 
the mountain, and plants are uniform in overall 
gross morphology. Cerro Bola is noteworthy as a 
station for several other phytogeographically in- 
teresting taxa, such as the near-endemic Arcto- 
staphylos bolensis P. V. Wells, as well as Ceano- 
thus papillosus Torr. var. roweanus McMinn and 
Lepechinia cardiophylla Epling, two taxa consid- 
erably disjunct from their previously know occur- 
rences in the Santa Ana Mountains of Orange and 


MADRONO 


[Vol. 49 


C. bolensis 

C. greggii perplexans 
C. g. vestitus 

C. ophiochilus 


IX <I I> ar © 


C. otayensis 


Principal components analysis without Ceanothus crassifolius, but including all other taxa considered in Fig. 


Riverside counties, California (Boyd et al. in 
prep). 


Paratypes. MEXICO, Baja California, chaparral 
with Chamaebatia australis on NE side of Cerro 
Bola, S of Tecate, elevation 1000 m, 26 Apr 1996, 
Jon E. Keeley 27232 (BCMEX); ibid, Jon E. Keeley 
27227 (CAS); ibid, Jon E. Keeley 27234 (SD); ibid, 
Jon E. Keeley 27238 (US); ibid, Jon E. Keeley 
27236 (MEXU); occasional in chaparral on north 
slope of Cerro Bola, seen to summit (1275 m), near 
31°19.5'N, 116°40’W, elevation ca. 550 m, 6 Jun 
1970, Reid Moran 17780 (RSA, SD). 


RELATIONSHIPS 


Ceanothus bolensis shows a marked morphologi- 
cal similarity to several other members of subgenus 
Cerastes in the southern California region. It is most 
distinct from the widespread C. greggii var. perplex- 
ans by having very small leaves that are deeply con- 
cave. These two leaf traits bear a strong resemblance 
to C. ophiochilus, however, leaf shape is markedly 
different, with the latter species being more similar 
in leaf shape to C. greggii var. vestitus. 

Ceanothus bolensis is quite distinct from another 
local endemic, C. otayensis, a taxon restricted to a 
few mountain peaks about 30-35 km northwest of 
Cerro Bola. Ceanothus otayensis has been sub- 
sumed under C. crassifolius (Schmitt 1993) due to 
the presence of several shared morphological sim- 
ilarities (revolute leaves and dense pubescence that 
includes brown hairs). Munz (1959) likewise treat- 
ed C. otayensis as C. xotayensis McMinn, and sug- 
gested it was probably a hybrid between C. cras- 
sifolius and C. greggii var. perplexans. However, 
principal component analysis on all 14 traits shows 
C. otayensis to be quite distinct from C. crassifolius 
(Fig. 1) and distinct from C. greggii (Fig. 2). Ex- 
tensive exploration of both known southern Cali- 
fornia localities of C. otayensis, the upper slopes of 
Otay Mtn. (1090 m) and San Miguel Mtn. (780 m), 


2002] 


BOYD AND KEELEY: CEANOTHUS BOLENSIS 293 


Fic. 3. 


Ceanothus bolensis. A. Fruiting branch showing characteristic small, toothed leaves, and hornless fruits on 


short axillary peduncles. B. Detail of individual flower. C. Detail of mature capsule showing absence of apical or lateral 


horns. (Illustrations by Melanie Baer-Keeley.) 


failed to uncover populations of either C. greggii 
or C. crassifolius. Ceanothus otayensis forms ho- 
mogenous populations that appear to be breeding 
true and lack any indication they are unstable hy- 
brid swarms (J. Keeley unpublished observations). 
In addition to the naming of C. bolensis, we suggest 
C. otayensis be recognized at the specific rank as 
described by McMinn (van Rensselaer and Mc- 
Minn 1942). 


ACKNOWLEDGMENTS 


Fieldwork and collections by Keeley were conducted in 
collaboration with Dr. Jose Delgadillo, Universidad Au- 
tonoma de Baja California, Ensenada, Baja California, 
Mexico (BCMEX). We wish to extend our thanks to Me- 
lanie Baer Keeley for preparing the illustration of C. bol- 
ensis, to Rosa Cerros Tlatilpa for kindly providing the 
Spanish resumen, and to Elizabeth Friar for assistance in 
producing figures | and 2. 


294 


LITERATURE CITED 


Boypb, S., T. Ross, AND L. ARNSETH. 1991. Ceanothus 
ophiochilus (Rhamnaceae): a distinctive, narrowly 
endemic species from Riverside County, California. 
Phytologia 70:28—41. 

Haropic, T. M., P. S. SOLTIS, AND D. E. Sottis. 2000. Di- 
versification of the North American shrub genus Ce- 
anothus (Rhamnaceae): conflicting phylogenies from 


MADRONO 


[Vol. 49 


nuclear ribosomal DNA and chloroplast DNA. Amer- 
ican Journal of Botany 87:108—123. 

ScHMIpT, C. L. 1993. Ceanothus. Pp. 932-938, 939, 941, 
943 in J. C. Hickman (eds.), The Jepson manual: 
higher plants of California. University of California 
Press, Berkeley, CA. 

VAN RENSSALAER, M. AND H. E. McMInn. 1942. Ceano- 
thus. Santa Barbara Botanic Garden, Santa Barbara, 
CA. 


MApRONO, Vol. 49, No. 4, pp. 295-297, 2002 


NOTE 


COLLINSIA ANTONINA IS EVOLUTIONARILY DISTINCT FROM C. PARRYI 
(SCROPHULARIACEAE SENSU LATO) 


BRUCE G. BALDWIN 
Jepson Herbarium and Department of Integrative Biology, 
1001 Valley Life Sciences Building #2465, University of California, 
Berkeley, CA 94720-2465 
bbaldwin @uclink4.berkeley.edu 


W. ScotrT ARMBRUSTER 
Department of Biology, Norwegian University of Science and Technology, N-7491 
Trondheim, Norway and Institute of Arctic Biology, University of Alaska, 
Fairbanks, AK 99775-7000 


A diminutive blue-eyed Mary from shale expo- 
sures in the San Antonio Hills (Santa Lucia Range) 
of southern Monterey County, California, was de- 
scribed by Hardham (1964) as Collinsia antonina. 
Plants assigned to C. antonina resemble members 
of C. parryi A. Gray, from the Transverse Ranges 
of southern California, and have been treated as 
either close relatives (Munz 1968) or, most recent- 
ly, as members (Neese 1993) of C. parryi. The con- 
siderable disjunction between populations in the 
Santa Lucia and Transverse ranges, although not 
unique [e.g., Syntrichopappus lemmonii (A. Gray) 
A. Gray (Compositae)], warrants closer scrutiny. 

In our evolutionary investigations of Collinsieae, 
we have examined phylogenetic relationships 
among taxa throughout the tribe using sequences of 
the internal transcribed spacer (ITS) region of nu- 
clear ribosomal DNA (see Armbruster et al. 2002). 
Upon including plants referable to C. antonina [E. 
C. Neese 21500 (JEPS)] in our analyses, we were 
surprised to find that members of C. parryi sensu 
stricto (s.s.) [E. C. Neese 21530 (JEPS)] appear to 
be more closely related to members of C. concolor 
Greene [E. C. Neese 21539A (JEPS)] than to mem- 
bers of C. antonina. Collinsia concolor and C. par- 
ryi S.S. constitute a strongly supported (99% boot- 
strap) clade, to the exclusion of C. antonina (Fig. 
1). Collinsia concolor and C. parryi s.s. are both 
endemic to the southern California Floristic Prov- 
ince and differ greatly in inflorescence architecture 
and flower size (Table 1). Based on the tree topol- 
ogy presented in Fig. 1, small flower size may be 
a shared, ancestral characteristic of C. antonina and 
C. parryi, with large flowers being a derived feature 
of C. concolor. Such major shifts in flower size 
have occurred repeatedly throughout the evolution- 
ary history of Collinsia (Armbruster et al. 2002). 


Evolutionary distinctiveness of C. antonina and 
C. parryi s.s. 1s also reflected by differences in 
morphological characteristics (Table 1). In keys to 
species of Collinsia (e.g., Newsom 1929), C. an- 
tonina might be confused with the distantly related 
C. childii Parry ex A. Gray because both taxa have 
glandular inflorescences and small flowers and 
both occur in the Santa Lucia Range. The two taxa 
can be readily distinguished by differences in seed 
number per capsule (Table 1). Hardham (1964) 
recognized two sympatric subspecies of C. anton- 
ina (C. a. subsp. antonina and C. a. subsp. pur- 
purea Hardham), which await further systematic 
evaluation. 

Collinsia antonina warrants recognition as a dis- 
tinct species based on the above molecular and 
morphological evidence. This narrowly endemic 
taxon was earlier considered rare and endangered 
by the California Native Plant Society (CNPS) 
(Smith and Berg 1988); since 1993, CNPS has fol- 
lowed Neese’s (1993) treatment of Californian 
members of Collinsia by including C. antonina 
within the circumscription of the common species, 
C. parryi (Skinner and Pavlik 1994; California Na- 
tive Plant Society 2001). In light of our findings, 
C. antonina deserves renewed attention by plant 
conservationists. 


ACKNOWLEDGMENTS 


We thank Elizabeth Chase Neese for her invaluable 
field collecting, taxonomic insights, and review of the 
manuscript, Bridget L. Wessa for extensive laboratory as- 
sistance, and reviewers Noel H. Holmgren, David J. Keil, 
and John L. Strother for helpful suggestions. This research 
was supported by grants from the Lawrence R. Heckard 
Endowment Fund of the Jepson Herbarium (to BGB), the 
Norwegian Research Council (to WSA), and NSF DEB- 
9708333 (to WSA). 


296 MADRONO 


100 
dg 
71 
50 
d2 
59 
d3 
97 
d6 

100 
ds 

97 

d7 
76 
d3 

65 
a3 100 
d1i3 
65 
d2 


Tonella tenella 


T. floribunda 


[Vol. 49 


Collinsia heterophylla 


C. greenei 

C. sparsiflora 
C. bartsiifolia 
C. corymbosa 


C. tinctoria 


C. parryi 


C. concolor 


C. antonina 


C. multicolor 
C. callosa 

C. childii 

C. parviflora 
C. grandiflora 
C. verna 

C. violacea 
C. rattanii 


C. linearis 


lA 


C. torreyi var. torreyi 


C. torreyi var. wrightii 


Fic. 1. The most parsimonious tree from phylogenetic analysis of nuclear ribosomal DNA sequences of the internal 
transcribed spacer region in tribe Collinsieae (Baldwin et al. unpublished; see Armbruster et al. 2002). The tree was 
rooted with sequences from outgroup taxa in tribe Cheloneae (Chelone, Keckiella, and Penstemon). Numbers above 
branches are bootstrap values (only values = 50% are shown); numbers below branches, preceded by “‘d’’, are decay 
values (only values > d1 are shown). The clade including Collinsia antonina, C. concolor, and C. parryi is highlighted. 


TABLE |. SOME MORPHOLOGICAL DIFFERENCES BETWEEN COLLINSIA ANTONINA AND SIMILAR OR CLOSELY RELATED SPECIES. 


C. antonina C. parryi 
Inflorescence glandularity glandular eglandular 
Longest-leaf length 5-10 mm (5—)10—45 mm 
Pedicel length <10 mm (5—)10—45 mm 
Corolla length 6—7 mm (5—)7—10 mm 
Corolla color white or purple lavender-blue 


Seeds/fruit 6-8 8—12 


C. concolor 


glandular or 
eglandular 

10—45 mm 

1-5 mm 

10-16 mm 

blue/lavender— 
blue and white 

11-12 


C. childii 


glandular 


10—50 mm 

5—25 mm 

6-8 mm 

pale lavender 
or white 

2 


2002] 


LITERATURE CITED 


ARMBRUSTER, W. S., C. P. H. MULDER, B. G. BALDwWIn, S. 
KALISZ, B. WESSA, AND H. Nute. 2002. Comparative 
analysis of late floral development and mating-system 
evolution in tribe Collinsieae (Scrophulariaceae s. 1.). 
American Journal of Botany 89:37—49. 

CALIFORNIA NATIVE PLANT SoctETy. 2001. Inventory of 
rare and endangered vascular plants of California, 6th 
ed. Rare Plant Scientific Advisory Committee, D. Ti- 
bor, Convening Editor. California Native Plant Soci- 
ety, Sacramento, CA. 

HARDHAM, C. B. 1964. A new Collinsia from Monterey 
County, California. Leaflets of Western Botany 10: 
133-135. 


BALDWIN AND ARMBRUSTER: COLLINSIA ANTONINA 


297 


Munz, P. A. 1968. Supplement to a California flora. Uni- 
versity of California Press, Berkeley. 

NEESE, E. C. 1993. Collinsia. Pp. 1024—1027 in J. C. 
Hickman (ed.), The Jepson manual: higher plants of 
California. University of California Press, Berkeley, 
CA. 

Newsom, V. M. 1929. A revision of the genus Collinsia. 
Botanical Gazette 87: 260-301. 

SKINNER, M. K., JR. AND B. M. PAVLIK (eds.). 1994. In- 
ventory of rare and endangered vascular plants of 
California, 5th ed. California Native Plant Society, 
Sacramento, CA. 

SmiTH, J. P., JR. AND K. BERG (eds.). 1988. Inventory of 
rare and endangered vascular plants of California, 4th 
ed. California Native Plant Society, Sacramento, CA. 


MADRONO, Vol. 49, No. 4, p. 298, 2002 


REVIEW 


Field guide to liverwort genera of Pacific North 
America, by W.B. Schofield. 2002. Global Forest So- 
ciety in association with the University of Washington 
Press, San Francisco, CA. 232 pp., 93 line drawings, 
glossary, and index. 7” X 10”. $25.00. ISBN 0-295- 
98194-6. Available at www.washington.edu/press 


Most of the liverwort field guides and other tax- 
onomic treatments covering this group of organ- 
isms in North America have focused primarily east 
of the 100th Meridian. It is therefore most welcome 
that this field guide covers much of the geograph- 
ical area of interest to Madrofo readers. This pa- 
perback book, while not quite a size to readily carry 
in the field, 1s nonetheless not heavy or excessively 
bulky. What we do have here is a well-designed 
volume. Bryophytes, the first land plants, are ar- 
ranged into three lineages. The liverworts (and 
hornworts) are clearly less well known to most vas- 
cular-trained botanists than are the mosses. This 
volume is easy to use and will expand one’s knowl- 
edge about this group of land plants. In some ways, 
this is a companion work to Schofield’s field guide 
of Some common mosses of British Columbia pub- 
lished back in 1969. The Introduction to the liver- 
wort genera field guide covers the first 22 pages 
and is concise yet very informative. The Introduc- 
tion addresses how to collect liverworts and horn- 
worts, followed by a short overview of collecting 
history of liverworts in Alaska, British Columbia, 
Washington, Oregon and California. Habitats, sea- 
sonality of finding liverworts, and distribution pat- 
terns in the region follow. Implications of liver- 
worts to people end the Introduction section. 

In most field guides, it is the illustrations and 
keys that determine how useful the book will ac- 
tually be, especially to someone approaching the 
group for the first time. In this regard Schofield’s 
book shines. While all keys have caveats about 
their use, I personally found the keys to be rather 
easy to use. The keys keep jargon and bryological 
terms to a basic minimum, and the couplets are 
succinct. The couplet choice is designed without a 
lot of overlapping characters. In many cases, genera 
key out in multiple places to accommodate the di- 
versity of species within larger liverwort genera. 
The key itself is divided into eight sub-key sections 
to further expedite the identification process. Each 
genus is arranged alphabetically so it is easy to lo- 
cate, and each genus has a full-page illustration. 
The illustrations are visually attractive and de- 
signed to represent a genus without being too tech- 
nical. In some ways, they are more “‘artistic”’ than 
a detailed illustration, such as would be used in 
describing a new taxon. Actually, I think the illus- 
trations provided are more effective as a represen- 


tation at the genus level. Since liverworts are gen- 
erally small plants, the bar scale provided with each 
illustration is important to review so one has a bet- 
ter idea of the organism’s actual size. 

A template is provided for each genus so com- 
parisons can be made readily among closely related 
genera. Each genus has a short explanation about 
the meaning of its name, a statement of the number 
of species in the genus, its habit, its habitat, repro- 
duction, local distribution, world distribution, dis- 
tinguishing characteristics and similar genera. 

The genera of hornworts within the Pacific North 
America are also included, adding further to the 
value of this book since one could confuse a horn- 
wort for a thallose liverwort. However, hornworts 
upon closer examination can readily be distin- 
guished from the liverworts, especially when spo- 
rophytes are present. The determination of the two 
hornwort genera covered in the book, cccurs at 
couplet 28 of key Hl. Another interesting feature, 
and I think a great decision, is adding Takakia 
(Takakiaceae) to the book. The placement of this 
genus within the bryophytes has been a consider- 
able puzzle, with some specialists insisting it is a 
liverwort while others opted to say it was more 
closely aligned with the mosses. Although it has 
been finally determined to be basal to the moss lin- 
eage (based on the sporophyte), it clearly looks 
more like a liverwort when only the gametophyte 
is present. I hope that additional occurrences of this 
relatively rare genus can be located in the Pacific 
Northwest. Having the genus illustrated here should 
be a good first step for others to seek it out. 

I strongly recommend this new field guide to all 
botanists with an interest in learning more about 
our non-flowering plants. As Schofield’s book 
states, liverworts are a remarkable group of plants, 
and upon closer observation, they make a consid- 
erable contribution to forest ecosystems, especially 
in the temperate rain forests along the coast of 
western North America. At $25.00 this guide is a 
good investment. Will you be able to key every 
liverwort to species with this field guide? Abso- 
lutely not. But, having the genus properly identified 
will speed up considerably the time to do so in 
more detailed bryofloras. This field guide is a won- 
derful introduction to recognizing genera of liver- 
worts in Pacific North America. In short order you 
will be able to recognize the common liverwort 
genera on rotten wood and logs, soil and humus, 
tree trunks, rock outcrops and even those liverworts 
submerged in springs and streams. I recommend it 
to botanists, ecologists, foresters, and conservation 
biologists without any reservations. 


—JAMES R. SHEVOCK, Research Associate, Department 
of Botany, California Academy of Sciences, Golden Gate 
Park, San Francisco, CA 94118-4599. 


MADRONO, Vol. 49, No. 4, p. 299, 2002 


PRESIDENT’S REPORT FOR VOLUME 49 


I am pleased to report that 2002 was a year of accom- 
plishment for the California Botanical Society, thanks to 
the dedicated efforts of the Editors and Council. Editor 
John Callaway skillfully concluded his first year in charge 
of Madrono, with four excellent issues completed. The 
quality, breadth, and depth of articles published in Ma- 
drono, Volume 49, testifies to the vibrancy of our journal 
and the effectiveness of John’s editorship. Thank you, 
John! I also thank our Book Editor Jon Keeley, Notewor- 
thy Collection Editors Dieter Wilken and Margriet Weth- 
erwax, and the Board of Editors for their important con- 
tributions to Volume 49. 

Visibility of articles in Madrono was enhanced consid- 
erably in 2002 by internet access to titles and abstracts for 
new issues of the journal through the California Botanical 
Society web-site (www.calbotsoc.org). John Callaway and 
Society web-master Curtis Clark worked with Allen Press 
to provide direct links to contents of Madrono (beginning 
with Vol. 49, issue 1) via Allen Press’s APT Online. The 
results of their efforts will help to increase citation and 
impact of articles in Madrono across the international bo- 
tanical community. 

Over the last year, web-masters Curtis Clark and John 
LaDuke have made other major improvements to the So- 
ciety web-site (www.calbotsoc.org), which has become 
our main clearinghouse for information about publishing 
in Madrono and about society activities, events, and mem- 
bership. Curtis and John developed a new, streamlined 
organization for the site, which is easily negotiated with 
well-placed links. The site also contains links to other so- 
cieties, institutions, and resources of importance to west- 
ern North American botany. Many thanks, Curtis and 
John! 

Thanks to efforts by Corresponding Secretary Sue 
Bainbridge and Treasurer Roy Buck, the Society can now 
accept credit-card payment for membership dues, using 
either the web-site membership forms (for new and re- 
newing members) or renewal envelopes sent with the third 
issue of Madronio each year. We trust that the convenience 
of using a credit card will encourage timely renewal of 
memberships, which remains an urgent need for the So- 
ciety. Thanks very much to those members who responded 
quickly to their renewal notices. For those who have not 
yet responded, please send your renewals in time for re- 
ceipt on 28 February 2003. Timely renewal of member- 
ship saves the Society considerable expenditure of effort 
(although late renewal is far better than a lapse in mem- 
bership). 

Significant growth of our membership base would allow 
the Society to do more to promote botanical research and 
education. Toward that end, please continue to encourage 
your colleagues to join us and to publish in Madrono. 
Also, you can help to support botanical research in eco- 
nomically depressed, developing countries by giving a 


sponsoring membership or subscription to a foreign sci- 
entist or scientific institution or by donating unused copies 
of Madrono. For more information on making such a gift, 
please contact Corresponding Secretary Sue Bainbridge 
(suebain @sscl.berkeley.edu). 

Fall 2002 saw outstanding lectures at our monthly 
meetings in Berkeley by Martin Bidartondo, Truman 
Young, and Randy Jackson, and we look forward to pre- 
sentations by distinguished scientists David Ackerly, Kim 
Steiner, Todd Dawson, and Marcel Rejmanek in winter 
and spring of 2003. I am indebted to outgoing First Vice- 
President Rod Myatt for his successful efforts, once again, 
at assembling such a fine lecture series for the Society’s 
program year. Attendance at our monthly meetings has 
been strong again this year and I encourage those of you 
who have an opportunity to participate to please join us, 
bring a friend, and inform your colleagues about upcom- 
ing lectures. 

I also strongly encourage our members to attend the 
Society’s biennial graduate student meeting and annual 
banquet at the University of San Diego on Saturday, 15 
February 2003. At the graduate student meeting, held dur- 
ing the day, students from different institutions will be 
presenting their research proposals, research-in-progress, 
or completed research in botany in a standard scientific- 
meeting format. We look forward to a stimulating day of 
research ideas and results from the promising, next gen- 
eration of botanists. In the evening, at the annual banquet, 
we will have the great pleasure of hearing from renowned 
Baja floristician and cactus expert Dr. Jon Rebman, who 
will be presenting an after-dinner lecture on recent floristic 
discoveries in Baja California, Mexico. Jon’s expeditions 
to remote, beautiful, and under-explored regions of Baja 
California, in collaboration with Mexican scientists, are 
fine examples of American/Mexican cooperation and ac- 
complishment in science. Thanks very much to Second 
Vice-President Michael Mayer for planning and organiz- 
ing the upcoming banquet and to Graduate Student Rep- 
resentatives Robert Lauri and Elizabeth Zacharias for co- 
ordinating and conducting the graduate student meeting. 

Last, but by no means least, I thank Recording Secre- 
tary Staci Markos and Council Members Jim Shevock, 
Dean Kelch, and Anne Bradley for their dedicated com- 
mitment and contributions to furthering the goals of the 
Society, and to all of our members for your continuing 
support and participation in the Society’s activities and 
events and for choosing to submit your botanical manu- 
scripts to Madrono. This is my last year as President of 
the Society and I especially thank all of you for making 
this experience so enjoyable and rewarding for me. Have 
a great year in 2003! 


—BRUCE G. BALDWIN 
December 2002 


MADRONO, Vol. 49, No. 4, p. 300, 2002 


EDITOR’S REPORT FOR VOLUME 49 


This report serves to inform the members of the Cali- 
fornia Botanical Society of the status of Madrono, from 
the number of manuscripts submitted to papers published. 
Since the previous editor’s report (see Madrono 49[4}), 
the journal has received 61 manuscripts for review, in- 
cluding Articles, Notes, and Noteworthy Collections; 31 
of these manuscripts have been accepted for publication 
in that same time period. The average time for article sub- 
mission to publication remains at approximately six 
months. Accepted manuscripts are typically published 
within approximately three to four months. Few manu- 
scripts were rejected after review; authors of Madrono ar- 
ticles did a fine job of responding to reviewers’ sugges- 
tions. 

Over the past five years, there has been a substantial 
effort to get Madrono back on its regular publication 
schedule, and we are now very close to being on schedule, 
with six issues published in 2002 (three from volume 48 
and three from volume 49). Kristina Schierenbeck (editor 
of volumes 45—48) deserves most of the credit for this, as 
she put in an enormous effort to get the journal back on 
schedule. In addition to catching up on our publication 
schedule, other improvements have been made with the 
journal. As noted by Bruce Baldwin in his President’s Re- 
port, abstracts of Madrono are now available on-line via 
the California Botanical Society’s web-site (www. 
calbotsoc.org). In addition the web-site will soon have 
more detailed Instructions for Authors for Madrono man- 


uscripts. With volume 49 we initiated a new policy, en- 
couraging authors to submit names of two to four potential 
reviewers for manuscripts. If readers have suggestions for 
other improvements for the journal please let me know. 

I want to thank the many people who make Madrono 
possible and who have been incredibly helpful in editing 
the journal: Kristina Schierenbeck, who gave me lots of 
help in taking over the editorship and has always been 
there to answer my questions about the journal; Bruce 
Baldwin and other members of the CBS Executive Coun- 
cil, who are always extremely helpful and supportive of 
everything related to Madrono; Dieter Wilken and Mar- 
griet Wetherwax, who handle all of the reviews for Note- 
worthy Collections (and without whom the Noteworthy 
Collections would not be possible); Jon Keeley, who does 
all of the book reviews; Steve Timbrook, who provides 
the annual index of Madrofo articles and the annual table 
of contents; the Board of Editors, who have provided in- 
put and advice whenever I’ve asked (especially Norm 
Ellstrand and Carla D’ Antonio, who are finishing their 
terms this year); Annielaurie Seifert at Allen Press, who 
has been extremely helpful when any editorial question 
arises; the College of Arts and Sciences at the University 
of San Francisco, who support my efforts on Madrono; 
and most of all the Madrono authors, who continue to 
submit outstanding manuscripts, and the reviewers (see 
the accompanying list of reviewers), who put in a sub- 
stantial effort to improve the quality of manuscripts that 
Madrono publishes. 


MADRONO, Vol. 49, No. 4, pp. 301, 2002 


Frank Almeda 
George Argus 
Jayne Belnap 
Jere Boudell 
Thomas Boyle 
Steven Broich 
Matt Brooks 
Mark Brunell 
Steve Caicco 
Robert Callihan 
Tina Carlsen 
Raymund Chan 
Anita Cholewa 
Ranessa Cooper 
Daniel Crawford 
Curtis Daehler 
Carla D’ Antonio 
Gerrit Davidse 
Chuck Davis 
Frank Davis 
Jose Delgadillo 
Rebecca Dolan 
Andrew Dyer 
Erin Espeland 
Ray Evert 


Phyllis Faber 
Richard Felger 
Wayne Ferren 
Peggy Fiedler 
Jack Fisher 

Ted Fleming 
Peter Goldblatt 
Leslie Gottlieb 
Jason Hamilton 
Héctor Hernandez Macias 
Noel Holmgren 
Kent Holsinger 
Bryan Jennings 
Steven Jessup 
Michael Josselyn 
Jon Keeley 
David Keil 
Seung-Chul Kim 
Robert Leidy 
Celi 

Tim Lowrey 
Carol Mallory-Smith 
Maria Mandujano 
Joe McAuliffe 
Kimberlie McCue 


REVIEWERS OF MADRONO MANUSCRIPTS 2002 


Dale McNeal 
Kyle Merriam 
Norton Miller 
Timothy Miller 
Richard Minnich 
Margaret Moore 
Lorraine Parsons 
Bob Patterson 
Eric Peterson 
Jeanne Ponzetti 
Jon Rebman 
John Reeder 
Marcel Rejmanek 
Rhonda Riggins 
James Shevock 
Stanley Smith 
Humberto Suzan 
Anthony Swinehart 
Teresa Terrazas 
Dale Vitt 

Dieter Wilken 
Paul Wilson 
Diana Wolf 
Lidia Yoshida 
Peter Zika 


MApRONO, Vol. 49, No. 4, pp. 302-304, 2002 


INDEX TO VOLUME 49 


Classified entries: major subjects, key words, and results; botanical names (new names are in boldface); geographical 
areas; reviews, commentaries. Incidental references to taxa (including most lists and tables) are not indexed separately. 
Species appearing in Noteworthy Collections are indexed under name, family, and state or country. Authors and titles 
are listed alphabetically by author in the Table of Contents to the volume. 


Aceana novae-zelandiae, noteworthy collection from OR, 
194. 

Agavaceae: Agave palmeri, effects of fire on reproductive 
biology, 1; nomenclatural history of Hesperoyucca 
whipplei and Yucca whipplei, 20. 

Agave palmeri, effects of fire on reproductive biology, 1. 

Agrostis Howellii, noteworthy collection from OR, 194. 

Alnus rubra, noteworthy collection from MT, 55. 

Ambrosia: A. includes Hymenoclea, 143. 

New combinations: A. monogyra, A. <Xplatyspina, A. 
salsola, A. salsola var. fasciculata, A. salsola var. 
pentalepis, 143. 

Arceuthobium gillii, adult sex ratio, 12. 

Arizona (see Mancoa and Mammillaria) 

Arundo donax, noteworthy collection from French Poly- 
nesia, 132. 

Asteraceae: Ambrosia includes Hymenoclea, 143; Crupina 
distribution in the Iberian Peninsula and Balearic Is- 
lands, 137; Senecio layneae, genetic structure, 150. 
New combinations: A. monogyra, A. <Xplatyspina, A. 

salsola, A. salsola var. fasciculata, A. salsola var. 
pentalepis, 143. 

Noteworthy collections: CA: Baccharis malibuensis, 
54. CO: Erigeron ochroleucus var. scriberni, 54. OR: 
Hieraceum caespitosum, 58; Petasites fragrans, 194. 
MT: Senecio congestus, 37. WA: Baccharis pilularis, 
132; Crepis setosa, Hieracium lachenalii, H. muro- 
rum, H. sabaudum, 196. 

Atriplex tridentata var. robusta, new var. from UT, cor- 
rection, 200. 

Azolla mexicana, noteworthy collection from MT, 55. 

Azollaceae (see Azolla) 


Baccharis: Noteworthy collections: B. malibuensis from 
CA, 54; B. pilularis from WA, 132. 

Balearic Islands (see Crupina) 

Bats (see Agave) 

Betulaceae (see Alnus) 

Biogenic hydrocarbons (see Gap Analysis Program) 

Biomass (see lichens) 

Botrychium pedunculosusm, noteworthy collection from 
6 eee 

Brassicaceae: Noteworthy collections: Cardamine flexu- 
osa from WA, 195; Lesquerella douglasii from MT, 57; 
Mancoa pubens from AZ, 132. 

Bryophyte records from the Mojave Desert, CA, 49. 

Bryophyta (see Bryophyte) 


Cactaceae: Hylocereus undatus and Opuntia ficus-indica, 
temperature limitations for cultivation, 228; Mammil- 
laria grahamii, flowering patterns and reproductive 
ecology, 201. 

Calamagrostis muiriana, new sp., 169. 

California: Bryophyte records from the Mojave Desert, 
49; canopy macrolichens from Sierra Nevada mixed co- 
nifer forests, 123; catalogue of non-native taxa not in 
The Jepson Manual, 61; Collinsia antonina evolution- 
arily distinct from C. parryi, 295; Cytisus scoparius and 


Genista monspessulana pollination, 25; Eremalche ex- 

ilis and E. kernensis, molecular and morphological per- 

spectives of sympatry, 22; Eschscholzia californica, fire 

and cold treatments effect on seed germination, 207; 

field assessment of the Gap Analysis Program GIS da- 

tabase, 99; Hylocereus undatus and Opuntia ficus- 

indica, temperature limitations for cultivation, 228; 

Nassella, population dynamics, 274; Pinus balfouriana 

importance in Klamath and southern Sierra Nevada 

mountains, 33; Quercus kelloggii seedling density fac- 

tors, 115; Senecio layneae, genetic structure, 150. 

New taxa: Eriogonum ovalifolium var. monarchense, 
16; Prunus eremophila, 285. 

Noteworthy collections: Baccharis malibuensis, 54; 
Drosera aliciae, 193; D. capensis, Utricularia sub- 
ulata, 194. 

Calystegia silvatica in western No. Am., 130; C. silvatica 
subsp. disjuncta, new subsp., 131. 

Cardamine flexuosa, noteworthy collection from WA, 
195%. 

Carex: Noteworthy collections: MT: C. chalciolepis, C. 
deflexa, C. lacustris, C. pallescens, C. prairea, C. va- 
ginata, 55. OR: Carex scirpoidea subsp. stenochlaena, 
194. 

Caryophyllaceae (see Cerastium and Moenchia) 

Caryophyllales (see Simmondsia) 

Ceanothus bolensis, new sp. from Baja Calif., Mexico, 
289. | 

Centaurium erythraea, noteworthy collection from MT, 
a1. 

Cerastium pumilum, noteworthy collections from OR and 
WA, 195. 

Chaparral (see Senecio layneae) 

Chenopodiaceae (see Atriplex) 

Chloris barbata, noteworthy collection from French Pol- 
ynesia, 132. 

Clusiaceae (see Hypericum) 

Collinsia antonina evolutionarily distinct from C. parryi, 
LS). 

Colorado: Sphagnum balticum in Rocky Mountain iron 
fen, 186. 

Noteworthy collections: Erigeron ochroleucus vat. scri- 
berni, Salix arizonica, §. discolor, 54. 

Columbia Plateau (see Navarretia) 

Compositae (see Asteraceae) 

Conifer diversity in Klamath and southern Sierra Nevada 
mountains, CA, 33. 

Convolvulaceae (see Calystegia) 

Cotoneaster: Noteworthy collections: OR: C. divaricatus, 
C. induratus, 195. WA: C. divaricatus, C. lucidus, C. 
nitens, C. salicifolius, C. tengyuehensis, 195. 

Crassulacean acid metabolism (see Hylocereus) 

Crepis setosa, noteworthy collection from WA, 196. 

Crupina distribution in the Iberian Peninsula and Balearic 
Islands, 137. 

Cryptogramma stelleri, noteworthy collection from ID, 
54. 

Cruciferae (see Brassicaceae) 


—— ye ee 


2002] 


Cyperaceae (see Carex and Cyperus) 
Cyperus difformis, noteworthy collection from WA, 196. 
Cytisus scoparius and Genista monspessulana pollination, 


Daphne laureola, noteworthy collection from OR, 194. 

Drosera aliciae and D. capensis, noteworthy collections 
from CA, 193. 

Droseraceae (see Drosera) 


Editor’s report, 300. 

Endemism (see Ceanothus, Guadalupe Island and Pucci- 
nellia) 

Eremalche exilis and E. kernensis, molecular and mor- 
phological perspectives of sympatry, 22. 

Erigeron ochroleucus var. scriberni from CO, 54. 

Eriogonum ovalifolium var. monarchense, new variety 
from CA, 16; E. visheri, noteworthy collection from 
MT, 57. 

Eschscholzia californica, fire and cold treatments effect 
on seed germination, 207. 


Fabaceae (see Cytisus) 

Fagaceae (see Quercus) 

Fire: Effect on reproductive biology of Agave, 1; Es- 
chscholzia californica, effect on seed germination, 207. 

Floral biology (see Mammillaria) 

Fraxinus pennsylvanica, noteworthy collection from WA, 
196. 

French Polynesia: Society Islands: Moorea: Noteworthy 
collections: Arundo donax, Chloris barbata, Hypar- 
rhenia rufa, Setaria sphacelata, 132. 


Galium pedmontanum, noteworthy collection from OR, 

194, and WA, 196. 

GAP (see Gap Analysis Program) 
Gap Analysis Program, field assessment of GIS database 

in central CA, 99. 

Gaudichaudia: Reticulate ancestry, 256, and revision of 

Mexican taxa, 237. 

New taxa: G. andersonii, 254; G. implexa, 247; G. 
intermixteca, 251; G. symplecta, 253: G. synop- 
tera, 251; G. zygoptera, 249; Nothosect. Cycloto- 
mopterys, 251; Nothosect. Tritomochaudia, 247; 
Nothosect. Zygomopterys, 249; Sect. Archaeopter- 
ys, 245; Sect. Cyclopterys, 240. 

Genetic structure (see Senecio layneae) 

Genista (see Cytisus) 

Gentianaceae see Centaurium) 

Geraniaceae (see Geranium) 

Geranium pyrenaicum, noteworthy collection from WA, 

196. 

Germination treatments (see Eschscholzia) 
Gramineae (see Poaceae) 

Grassland (see Nassella) 

Grossulariaceae (see Ribes) 

Guadalupe Island, Mexico, flora of Toro Islet, 145. 


Halophyte (see Pucinellia) 

Hepaticophyta (see Bryophyte) 

Hesperoyucca whipplei and Yucca whipplei, nomenclatur- 
al history, 20. 

Hieraceum caespitosum, noteworthy collection from OR, 
58; H. lachenalii, H. murorum, H. sabaudum, notewor- 
thy collections from WA, 196. 

Hylocereus undatus, temperature limitations for cultiva- 
tion, 228. 

Hymenoclea (see Ambrosia) 


INDEX TO VOLUME 49 303 


Hyparrhenia rufa, noteworthy collection from French Pol- 
ynesia, 132. 

Hypericum maculatum subsp. obtusiusculum, noteworthy 
collection from WA, 196. 


Iberian Peninsula (see Crupina) 

Idaho: Noteworthy collections: Cryptogramma stelleri, 
54; Viola selkirkti, 55. 

Introgression (see Gaudichaudia) 

Invasive plants: Catalog of non-native taxa occurring in 
CA not included in Jepson Manual, 61; Cytisus and 
Genista in CA, 25; Crupina in Iberian Peninsula and 
Balearic Islands, 137. 

ISSR (see Senecio layneae) 


Jepson Manual, catalog of non-native taxa occurring in 
CA not included in, 61. 

Juncaceae (see Juncus) 

Juncus patens, noteworthy collection from WA, 196. 


Klamath Mountains, CA (see Pinus) 


Labiatae (see Lamiaceae) 

Lamiaceae (see Stachys) 

Leguminosae (see Fabaceae) 

Lentibulariaceae (see Utricularia) 

Lesquerella douglasii, noteworthy collection from MT, 57. 

Lichens: Canopy macrolichens from Sierra Nevada, CA, 
mixed conifer forests, 123. 

Litterfall (see lichens) 


Malpighiaceae (see Gaudichaudia) 
Malvaceae (see Eremalche) 
Mammillaria grahamii, flowering patterns and reproduc- 

tive ecology, 201. 

Mancoa pubens, noteworthy collection from AZ, 132. 
Mexico: Gaudichaudia, reticulate ancestry, 256, and re- 
vision of Mexican taxa, 237; Guadalupe Island, flora of 

Toro Islet, 145. 

New taxa: Ceanothus bolensis, 289; Gaudichaudia 
andersonii, 254; G. implexa, 247; G. intermixteca, 
251; G. symplecta, 253; G. synoptera, 251; G. zyg- 
optera, 249; Nothosect. Cyclotomopterys, 251; No- 
thosect. Tritomochaudia, 247; Nothosect. Zygo- 
mopterys, 249; Sect. Archaeopterys, 245; Sect. Cy- 
clopterys, 240. 

Mimulus ringens, noteworthy collection from MT, 57. 
Moenchia erecta, noteworthy collection from WA, 196. 
Mojave Desert, CA (see Bryophyte and Prunus) 
Montana: Noteworthy collections: Alnus rubra, Azolla 
mexicana, Botrychium pedunculosusm, Carex chalci- 
olepis, C. deflexa, C. lacustris, C. pallescens, C. prai- 
rea, C. vaginata, Centaurium erythraea, Eriogonum 
visheri, Lesquerella douglasii, Mimulus ringens, Ribes 
laxiflorum, Senecio congestus, Ventenata dubia, Viola 


selkirkii, 55. 


Nassella, population dynamics in CA, 274. 

Navarretia leucophylla subsp. diffusa from eastern WA 
vernal pools, 165. 

New Mexico: Noteworthy collection of Salix wolfii var. 
wolfii, 54. 


Ophiogloassaceae (see Botrychium) 

Oleaceae see (Fraxinus) 

Opuntia ficus-indica, temperature limitations for cultiva- 
tion, 228. 

Oregon: Noteworthy collections: Aceana novae-zelandiae, 


304 


Agrostis Howellii, Carex scirpoidea subsp. stenochlae- 
na, 194; Cerastium pumilum, Cotoneaster divaricatus, 
C. induratus, 195; Daphne laureola, Galium pedmon- 
tanum, 194; Hieraceum caespitosum, 58; Petasites fra- 
grans, 194; Sorbus californica, Veronica verna, 195. 


Pest plants (see invasive plants) 

Petasites fragrans, noteworthy collection from OR, 194. 

Photinia davidiana, P. villosa, noteworthy collection from 
WA, 196. 

Pinaceae (see Pinus) 

Pinus balfouriana importance in Klamath and southern 
Sierra Nevada mountains, CA, 33; P. chihuahana in- 
fection by Arceuthobium, 12; P. subsect. Ponderosae 
novel lineage, 189. 

Poaceae: Nassella, population dynamics in CA, 274; Puc- 
cinellia howellii, salt spring zonation, 178. 

New taxon: Calamagrostis muiriana, 169. 

Noteworthy collections: Agrostis Howellii, 194; Arundo 
donax, Chloris barbata, Hyparrhenia rufa, Setaria 
sphacelata from French Polynesia, 132; Ventenata 
dubia from MT, 57. 

Polemoniaceae (see Navarretia) 

Polygonaceae (see Eriogonum) 

Polyploidy (see Calamagrostis and Gaudichaudia) 

President’s report, 299. 

Prunus eremophila, new sp. from CA, 285. 

Pteridaceae (see Cryptogramma) 

Puccinellia howellii, salt spring zonation, 178. 


Quercus kelloggii seedling density factors, 115. 


RAPD (see Gaudichaudia and Senecio layneae) 

Recruitment (see Quercus) 

Reviews: A Cactus Odyssey; Journeys in the Wilds of Bo- 
livia, Argentina, and Peru by James D. Mauseth, Rob- 
erto Kiesling and Carlos Ostolaza, 198; Field Guide to 
Liverwort Genera of Pacific North America by W. B. 
Schofield, 298; Illustrated Field Guide to Selected Rare 
Plants of Northern California, eds. Gary Nakamua and 
Julie Kiersteand Nelson, 48; Inventory of Rare and En- 
dangered Plants of California, 6th ed. California Native 
Plant Society, 135; The Manzanitas of California, also 
of Mexico and the World by Phillip V. Wells, 46; Seeing 
Things Whole: The Essential John Wesley Powell ed. 
by William deBuys, 143. 

Rhamnaceae (see Ceanothus) 

Ribes laxiflorum, noteworthy collection from MT, 57. 

Rosaceae: Prunus eremophila, new sp. from CA, 285. 
Noteworthy collections: OR: Aceana novae-zelandiae, 

194; Cotoneaster divaricatus, C. induratus, Sorbus 
californica, 195. WA: Cotoneaster divaricatus, C. lu- 
cidus, C. nitens, C. salicifolius, C. tengyuehensis, 
Photinia davidiana, 195; P. villosa, 196. 

Rubiaceae (see Galium) 


MADRONO 


[Vol. 49 


Salicaceae (see Salix) 

Salix: Noteworthy collections: S. arizonica, S. discolor, 
from CO, S. wolfii var. wolfii from NM, 54. 

Salt springs (see Puccinellia) 

Scrophulariaceae: Collinsia antonina evolutionarily dis- 
tinct from C. parryi, 295. 

Noteworthy collections: Mimulus ringens from MT, 57; 
Veronica verna from OR, 195, and WA, 197. 

Seed dormancy (see Eschscholzia) 

Seedlings (see Quercus) 

Senecio: S. congestus, noteworthy collection from MT, 57; 
S. layneae, genetic structure, 150. 

Setaria sphacelata noteworthy collection from French 
Polynesia, 132. 

Sex ratio (see Arceuthobium) 

Sierra Nevada, CA: Canopy macrolichens from mixed co- 
nifer forests, 123; Pinus balfouriana importance, 33; 
Quercus kelloggii seedling density factors, 115. 

Simmondsia anatomical evidence for inclusion in Cary- 
ophyllales s.l., 158. 

Simmondsiaceae (see Simmondsia) 

Sorbus californica, noteworthy collection from OR, 195. 

Sphagnum balticum in Rocky Mountain iron fen, 186. 

Stachys arvensis, noteworthy collection from WA, 195. 


Thymelaeaceae (see Daphne) 


Utah (see Atriplex) 
Utricularia subulata, noteworthy collection from CA, 
194. 


Vegetation survey (see Gap Analysis Program) 

Vegetation zonation (see Puccinellia) 

Ventenata dubia, noteworthy collection from MT, 57. 

Vernal pools (see Navarretia) 

Veronica verna, noteworthy collections from OR, 195, 
and WA, 197. 

Viola selkirkii, noteworthy collections from ID, 55, and 
NY, 6 oe 

Violaceae (see Viola) 

Viscaceae (see Arceuthobium) 


Washington: Navarretia from vernal pools, 165. 
Noteworthy collections: Baccharis pilularis, 132; Car- 

damine flexuosa, Cerastium pumilum, Cotoneaster 
divaricatus, C. lucidus, C. nitens, C. salicifolius, 195; 
C. tengyuehensis, Crepis setosa, Cyperus difformis, 
Fraxinus pennsylvanica, Galium pedmontanum, Ge- 
ranium pyrenaicum, Hieracium lachenalii, H. mu- 
rorum, H. sabaudum, Hypericum maculatum subsp. 
obtusiusculum, Juncus patens, Moenchia erecta, 
Photinia davidiana, 196; P. villosa, Stachys arvensis, 
Veronica verna, 197. 

Weeds (see invasive plants) 


Yucca whipplei (see Hesperoyucca) 


Mapbrono, Vol. 49, No. 4, pp. 305-306, 2002 


DEDICATION 


Donald R. Kaplan 


Donald R. Kaplan, Professor in the Department 
of Plant and Microbial Biology at UC Berkeley, has 
made exceptional contributions to research and ed- 
ucation in the field of plant morphology over the 
last four decades. He received his B.A. in Biology 
in 1960 from Northwestern University and his 
Ph.D. in Botany from the University of California, 
Berkeley in 1965. His dissertation research, on 
‘Floral morphology and reproduction in certain 
members of the genus Downingia (Campanulaceae; 
Lobelioideae)’’ marked the beginning of an illus- 
trious career that continues to this day. Don joined 
the UC Berkeley faculty in 1968, where he estab- 
lished himself as an outstanding researcher, mentor, 
and educator. He has always tried to blur the 
boundary between research and teaching. Accord- 
ing to Don, “I am a better teacher because I do 
research . . . similarly, being a teacher gives my re- 
search a better perspective and helps me to place 
the more detailed aspects of what I do in the broad- 
er context of knowledge in my field.”’ 

Don’s focus has been on the developmental basis 


that underlies the diversity of plant form. He has 
sought to move away from the traditional approach 
of plant morphology as a systematic survey of ma- 
jor groups in the plant kingdom. Instead, he has 
promoted the approach of integration and analysis 
of key concepts in morphology. His successful ef- 
forts have been well recognized. In 1998, he re- 
ceived the Jeannette Siron Pelton Award from the 
Botanical Society of America (BSA) for his sus- 
tained and creative contributions in experimental 
plant morphology. In addition, Don also received 
the Alexander von Humboldt Distinguished Senior 
U.S. Scientist Award. 

In citing Don for the Jeannette Siron Pelton 
Award, the BSA noted that “‘he has reached out 
both to traditional plant biologists as well as to 
those who have come to plant biology through an 
interest in molecular aspects of biology. He is the 
author of numerous substantive publications on 
leaf development that are recognized as classic pa- 
pers on the subject.’’ He also has completed influ- 
ential work evaluating the relationship of cells to 


306 


organisms in plants. This work has forced bota- 
nists and geneticists to “‘reevaluate their thinking 
about the underlying mechanisms responsible for 
the origin of plant form.”’ His review of the sci- 
ence and history of plant morphology, going back 
to its origins with Goethe in the late 1700s, which 
is based on his acceptance speech for the Jeannette 
Siron Pelton Award, was published in the October 
2001 issue of the American Journal of Botany (88: 
1711-1741). 

Don has trained a large number of graduate stu- 
dents, many who have gone on to very productive 
careers in plant morphology. He has reached an 
even larger number of aspiring young scientists 
through his courses, which have drawn high praise 
from generations of Berkeley’s undergraduates and 
graduate students. Don brings to the classroom an 
enthusiastic and innovative approach to the subject 
of plant form and inspires his students with his in- 
satiable curiosity about the biological world. As 


MADRONO 


[Vol. 49 


one student remarked about Don’s class on plant 
morphology, “the course is a must-have for any 
serious student of botany. One should not pass up 
the opportunity to learn from Dr. Kaplan.”’ Another 
student noted that “‘Dr. Kaplan is one of the people 
I’ve met in academia that has influenced and in- 
spired me the most, both as a researcher and as a 
teacher.’ His outstanding teaching was recognized 
in 1976 when he received the campus-wide Distin- 
guished Teaching Award at UC Berkeley. 

His teaching has also led to the development of 
a multi-volume text, entitled “Principles of Plant 
Morphology.”’ This book has been the primary fo- 
cus of Don’s recent work. We look forward to its 
completion and the insight that it will bring to cur- 
rent and future students of plant morphology. In the 
spirit of his true devotion to plant biology through 
outstanding research and inspiring teaching, we 
dedicate volume 49 of Madrono to Donald R. Kap- 
lan. 


MADRONO 


A WEST AMERICAN JOURNAL OF BOTANY 


VOLUME XLIx 
2002 


BOARD OF EDITORS 


Class of: 


2002—-NORMAN ELLSTRAND, University of California, Riverside, CA 
CARLA M. D’ ANTONIO, University of California, Berkeley, CA 


2003—-FREDERICK ZECHMAN, California State University, Fresno, CA 
JON E. KEELEY, U.S. Geological Service, Biological Resources Division, 
Three Rivers, CA 


2004—Davip Woop, California State University, Chico, CA 
INGRID PARKER, University of California, Santa Cruz, CA 


2005—J. MARK PorRTER, Rancho Santa Ana Botanic Gardon, Claremont, CA 
JON. P. REBMAN, San Diego Natural History Museum, San Deigo, CA 


Editor—JOHN CALLAWAY 
Department of Environmental Sciences 
University of San Francisco 
2130 Fulton Street 
San Francisco, CA 94117-1080 
callaway @usfca.edu 


Published quarterly by the 
California Botanical Society, Inc. 
Life Sciences Building, University of California, Berkeley 94720 


Printed by Allen Press, Inc., Lawrence, KS 66044 


MADRONO, Vol. 49, No. 4, pp. ii-iv, 2002 


TABLE OF CONTENTS 


Acker, Steven A. (see Shaw, David C.) 

Albert, Wallace E. (see Spribille, Toby, et al.) 

Albertson, Eugene D. (see Karlik, John F, et al.) 

Andreasen, Katarina, Ellen A. Cypher and Bruce G. Baldwin, Sympatry between desert mallow, Eremalche 
exilis and Kern mallow, E. kernensis (Malvaceae): Molecular and morphological perspectives 

Andrus, Richard E. (see Cooper, David J.) 

Armbruster, W. Scott (see Baldwin, Bruce G., and W. Scott Armbruster) 

Arp, Christopher D. (see Cooper, David J.) 

Arvidson, G. Michael (see Spribille, Toby, et al.) 

Ayres, Debra R. (see Marsh, Glenda D.) 

Bacca, Mary (see Levine, Larry) 

Baer-Keeley, Melanie, Review of Inventory of Rare and Endangered Plants of California, 6th edition, convening 
ed. Davad P. TibOtie2: es ch se LI Bb Sates! AM ae Bae RR A Nios kn lr od ee ae, ee 

Baldwin, Bruce G., President’s report for Volume 49 

Baldwin, Bruce G. (see also Andreasen, Katarina) 

Baldwin, Bruce G. (see also Strother, John L., and Bruce G. Baldwin) 

Baldwin, Bruce G., and W. Scott Armbruster, Collinsia antonina is evolutionarily distinct from C. parryi (Scro- 
phulariaceaessemsulato)y sca Fhe Sas ee Ne ce a a 

Bartolome, James W. (see McClaran, Mitchel P.) 

Beilman, David W. (see Nobel, Park S., et al.) 

Bjork, Curtis R., A new subspecies of Navarretia leucocephala (Polemoniaceae) from vernal pools in eastern 
Washimg@toms sxc. 4S yh oes a re oes Mea 2 a RT sae aI, A ee 

Bowers, Janice E., Flowering patterns and reproductive ecology of Mammillaria grahamii (Cactaceae), a com- 
mon, smiall.cactusim the Sonoram Desert’ 2... | ee 

Boyd) Steve, Noteworthy (collection) frome @ alist ori cgeeee eee see ee ea Nn 

Boyd, Steve, and Jon E. Keeley, A new Ceanothus (Rhamnaceae) species from northern Baja California, Mexico 


Brainerd, Richard (see Newhouse, Bruce) 

Brummitt, R. K., Calystegia silvatica (Convolvulaceae) in western North America 

Brunsfeld, Steven J. (see Patten, Ann M.) 

Carlquist, Sherwin, Wood anatomy and successive cambia in Simmondsia (Simmondsiaceae): Evidence for 
inclusion im’ Caryophyllales*s.l. ....2. 2. ne ee eee 

Chung, Y. Jae (see Karlik, John F, et al.) 

Cooper, David J., Richard E. Andrus and Christopher D. Arp, Sphagnum balticum in a southern Rocky Mountain 
ifOn: fem: 220.04 es OE, BE a UNE Se 

Cypher, Ellen A. (see Andreasen, Katarina) 

Daugherty, Carolyn M. (see Mathiasen, Robert L.) 

De la Barrera, Erick (see Nobel, Park S., et al.) 

Dean, Ellen (see Hrusa, Fred, et al.) 

Doherty, Jennifer H. (see Nobel, Park S., et al.) 

Dorn, Robert D., Noteworthy collections from Colorado and New Mexico 

Dwire; KathleenvA Noteworthy, (collection trong @ reo meee 

Eckert, Andrew J., and John O. Sawyer, Foxtail pine importance and conifer diversity in the Klamath Mountains 
and southern Sierra Nevada, California 

Engel, Alexandra (see Parker, Ingrid M.) 

Ertter, Barbara (see Hrusa, Fred, et al.) 

Fiest-Alvey, Laura J. (see Montalvo, Arlee M.) 

Fulgham, K. O. (see Levine, Larry) 

Gamarra, Roberto, and Cindy Talbott Roché, Distribution of the genus Crupina in the Iberian Peninsula and the 
Balearic Psland’s:xcc... 8 i ee Me A ert Er ree GE we a nC 

Garrison, Barrett A., Robin L. Wachs, James S. Jones and Matthew L. Triggs, Some factors influencing seedling 
density of California black oak (Quercus kelloggii) in the central Sierra Nevada, California 

Goodell, Karen (see Parker, Ingrid M.) 

Gray, Sami (see Wilson, Barbara L.) 

Greenhouse, Jeffrey A., and John L. Strother, Hesperoyucca whipplei and Yucca whipplei (Agavaceae) _______- 

Griffin, James R. (see Hamilton, Jason G.) 

Griffith, M. Patrick, Review of A Cactus Oddyssey: Journeys in the Wilds of Bolivia, Argentina, and Peru by 
James DF Mauseth Robesto kueslinieeand’ €anrlosi@ soles eee aa 

Hamilton, Jason G., James R. Griffin and Mark R. Stromberg, Long-term population dynamics of native Nassella 
(Poaceae) bunchgrasses in central California : 

Haubensak, Karen A. (see Parker, Ingrid M.) 

Heidel, Bonnie (see Spribille, Toby, et al.) 


DE 


135 
2g 


2S)5) 


165 


201 
54 


289 


130 


158 


186 


54 
58 


35 


198 


2002] TABLE OF CONTENTS 


Hrusa, Fred. Barbara Ertter, Andrew Sanders, Gordon Leppig and Ellen Dean, Catalogue of non-native vascular 
plants occurring spontaneously in California beyond those addressed in The Jepson Manual—Part I 

Jessup, Steven L., Reticulate ancestry in Mexican Gaudichaudia (Malpighiaceae) 

Jessup, Steven L., Six new species and taxonomic revisions in Mexican Gaudichaudia (Malpighiaceae) 

Johnson, Phillip (see Rice, Barry A., and Phillip Johnson) 

Jones, James S. (see Garrison, Barrett A., et al.) 

Kapliaas Douala de edicaony ar VONNNG 45 0) ee Ee ee 

Karlik, John F, et al., Eugene D. Albertson, Y. Jae Chung, Alistair H. McKay and Arthur M. Winer, Field 
assessment of the California GAP analysis program GIS database in central California 

Keeley, Jon E. (see Boyd, Steve, and Jon E. Keeley) 

Koehler, Catherine E. (see Montalvo, Arlee M.) 

Leon de la Luz, José Luis (see Rebman, Jon) 

Leppig. Gordon (see Hrusa, Fred, et al.) 

Levine, Larry, Mary Bacca and K. O. Fulgham, Plant zonation in a Shasta County salt spring supporting the 
Gnlgsknowe popllaiom Of Paccmciia howell (Poaceae): 22 

Maksnes blsdbor: NOlowotay Colleciiog from Arizona. 2-2 

Marsh, Glenda D., and Debra R. Ayres, Genetic structure of Senecio layneae (Compositae): A rare plant of the 
ic re ee Fe ea Ee wee EE 2 ee ee a et 

Mathiasen, Robert L., and Carolyn M. Daugherty, Adult sex ratio of Arceuthobium gillii (Viscaceae) _..._. 

McClaran, Mitchel P., and James W. Bartolome, Noteworthy collections from Moorea, Society Islands, French 
SESS loners et cee EI eee ee ac 

McKay, Alistair H. (see Karlik, John F, et al.) 

Mishler, Brent D. (see Stark, Lloyd R.) 

Montalvo, Arlee M., Laura J. Feist-Alvey and Catherine E. Koehler, The effect of fire and cold treatments on 
seed germination of annual and perennial populations of Eschscholzia californica (Papaveraceae) in south- 
EEE SPC es lee Se eee ee oa ae Re a A eth ep ae ae ee ee 

Newhouse, Bruce, Richard Brainerd and Peter F Zika, Noteworthy collections from Oregon — 

Nobel, Park S., Erick De la Barrera, David W. Beilman, Jennifer H. Doherty and Brian R. Zutta, Temperature 
BEseATOUS 20h Cll liVvaAlOn: OF CHibic cacunimec alilomiia «ea es ee oS 

Oberbauer, Thomas A. (see Rebman, Jon) 

Parker, Ingrid M., Alexandra Engel, Karen A. Haubensak and Karen Goodell, Pollination of Cytisus scoparius 
(Fabaceae) and Genista monspessulana (Fabaceae), two invasive shrubs in California 

Parker, V. Thomas (see Vasey, Michael C.) 

Patten, Ann M., and Steven J. Brunsfeld, Evidence of a novel lineage within the Ponderosae _. 

Pivorunas, David, Veva Stansell and Peter F Zika, Noteworthy collection from Oregon — 

Prigge, Barry A., A new species of Prunus (Rosaceae) from the Mojave Desert of California 

Rebman, Jon, Thomas A. Oberbauer and José Luis Ledén de la Luz, The flora of Toro Islet and notes on 
ees amt ty apa. Calin hilt VIC KACO) ee ee et 

ice TEA NeW COMECCHO MS MIN © ALIMENT A, o os ee ee 

nice. Samy A. and Phillip Johnson, Noteworuhy collection from Alaska <= 2s 

Roché, Cindy Talbott (see Gamarra, Roberto) 

Sanders, Andrew (see Hrusa, Fred, et al.) 

Sawyer, John O., Review of Jllustrated Field Guide to Selected Rare Plants of Northern California edited by 
Gary Nakamura and Julie Kiersteand Nelson 

Sawyer, John O. (see also Eckert, Andrew J.) 

Beale Ce Ole were COUECHOn 1100M WASHINOFOM | 08 

Shaw, David C., and Steven A. Acker, Canopy macrolichens from four forest stands in the southern Sierra mixed 
PURI CES Se Gigs S 2 Te iT CE (Gime hil) SEL gM oct dace ccs ee eee ce see ee 

Shevock, James R., Review of Field Guide to Liverwort Genera of Pacific Northwest America by W. B. Schofield 


Slauson, Liz A., Effects of fire on the reproductive biology of Agave palmeri (Agavaceae) — 

aC eeageRINGtWOrHy CONCCHOUS 1fOnT WanOn. so" ee ee 

pene slOuy cL a. Nelowonmny collections from Montana 

Stansell, Veva (see Pivorunas, David) 

Stark, Lloyd R., Alan T. Whittemore and Brent D. Mishler, Noteworthy bryophyte records from the Mojave 
eS ECE Lo ese Gee SI RE ee ee a <a Rede ed ee 

Stromberg, Mark R. (see Hamilton, Jason G.) 

Strother, John L. (see Greenhouse, Jeffrey A.) 

Strother, John L., and Bruce G. Baldwin, Hymenocleas are ambrosias (Compositae) — 

Triepke, F Jack (see Spribille, Toby, et al.) 

Triggs, Matthew L. (see Garrison, Barrett A., et al.) 

Tweed, William, Review of Seeing Things Whole: The Essential John Wesley Powell ed. William deBuys __ 

Vanderhorst, Jim (see Spribille, Toby, et al.) 

Vasey, Michael C., and V. Thomas Parker, Review of The Manzanitas of California, also of Mexico and the 
Scat ed Ie BTINLATS RES, nn eae seeemeeees See TPE) Ne ee 

Wachs, Robin L. (see Garrison, Barrett A., et al.) 

Wilson, Barbara L., and Sami Gray, Resurrection of a century-old species distinction in Calamagrostis 

Winer, Arthur M. (see Karlik, John F, et al.) 


iil 


MApRONO, Vol. 49, No. 4, pp. ii-iv, 2002 


TABLE OF CONTENTS 


Acker, Steven A. (see Shaw, David C.) 

Albert, Wallace E. (see Spribille, Toby, et al.) 

Albertson, Eugene D. (see Karlik, John FE, et al.) 

Andreasen, Katarina, Ellen A. Cypher and Bruce G. Baldwin, Sympatry between desert mallow, Eremalche 
exilis and Kern mallow, E. kernensis (Malvaceae): Molecular and morphological perspectives 

Andrus, Richard E. (see Cooper, David J.) 

Armbruster, W. Scott (see Baldwin, Bruce G., and W. Scott Armbruster) 

Arp, Christopher D. (see Cooper, David J.) 

Arvidson, G. Michael (see Spribille, Toby, et al.) 

Ayres, Debra R. (see Marsh, Glenda D.) 

Bacca, Mary (see Levine, Larry) 

Baer-Keeley, Melanie, Review of Inventory of Rare and Endangered Plants of California, 6th edition, convening 
ed. David -P "Tb iiiss. . bP aie a Cl ky tos Be i A Sses Sande ae) Lae ee 

Baldwin, Bruce.G:, Presidents reportsion Volume<49 2a ee 

Baldwin, Bruce G. (see also Andreasen, Katarina) 

Baldwin, Bruce G. (see also Strother, John L., and Bruce G. Baldwin) 

Baldwin, Bruce G., and W. Scott Armbruster, Collinsia antonina is evolutionarily distinct from C. parryi (Scro- 
phulaniaceae ‘sensw“lat), 22 2a 2 Ae ee eee 

Bartolome, James W. (see McClaran, Mitchel P._) 

Beilman, David W. (see Nobel, Park S., et al.) 

Bjork, Curtis R., A new subspecies of Navarretia leucocephala (Polemoniaceae) from vernal pools in eastern 
‘Weagshairn pits 5st 9 ap gn te ea ne oe ou as Peg 

Bowers, Janice E., Flowering patterns and reproductive ecology of Mammillaria grahamii (Cactaceae), a com- 
mon, small cactus: inthe Sonoran Desert = eee 2 ee eee 

Boyd) Steve, Noteworthy collecuon from Caliiomia = a ee eee 

Boyd, Steve, and Jon E. Keeley, A new Ceanothus (Rhamnaceae) species from northern Baja California, Mexico 


Brainerd, Richard (see Newhouse, Bruce) 

Brummitt, R. K., Calystegia silvatica (Convolvulaceae) in western North America 

Brunsfeld, Steven J. (see Patten, Ann M.) 

Carlquist, Sherwin, Wood anatomy and successive cambia in Simmondsia (Simmondsiaceae): Evidence for 
inclusion: imsCaryophivillales:s.l 23 a ee ee 

Chung, Y. Jae (see Karlik, John FE, et al.) 

Cooper, David J., Richard E. Andrus and Christopher D. Arp, Sphagnum balticum in a southern Rocky Mountain 
ION FEM, 228 ies ogee see ee el EE, SI yaa Se Seas Pi ee art er 

Cypher, Ellen A. (see Andreasen, Katarina) 

Daugherty, Carolyn M. (see Mathiasen, Robert L.) 

De la Barrera, Erick (see Nobel, Park S., et al.) 

Dean, Ellen (see Hrusa, Fred, et al.) 

Doherty, Jennifer H. (see Nobel, Park S., et al.) 

Dorn, Robert D., Noteworthy collections from Colorado and New Mexico 

Dywane, KathleenvA-, Noteworthyscollectromlir oma © re cone eee rt 

Eckert, Andrew J., and John O. Sawyer, Foxtail pine importance and conifer diversity in the Klamath Mountains 
and southern Sierra Nevada, California 

Engel, Alexandra (see Parker, Ingrid M.) 

Ertter, Barbara (see Hrusa, Fred, et al.) 

Fiest-Alvey, Laura J. (see Montalvo, Arlee M.) 

Fulgham, K. O. (see Levine, Larry) 

Gamarra, Roberto, and Cindy Talbott Roché, Distribution of the genus Crupina in the Iberian Peninsula and the 
Balkearic: Esler: co ces oe a os bo A eR oh UO SPOR a NE 

Garrison, Barrett A., Robin L. Wachs, James S. Jones and Matthew L. Triggs, Some factors influencing seedling 
density of California black oak (Quercus kelloggii) in the central Sierra Nevada, California 

Goodell, Karen (see Parker, Ingrid M.) 

Gray, Sami (see Wilson, Barbara L.) 

Greenhouse, Jeffrey A., and John L. Strother, Hesperoyucca whipplei and Yucca whipplei (Agavaceae) 

Griffin, James R. (see Hamilton, Jason G.) 

Griffith, M. Patrick, Review of A Cactus Oddyssey: Journeys in the Wilds of Bolivia, Argentina, and Peru by 
James D> Mauseth; Roberto: Kiesins and! Carlosy © sto eizay ee eee 

Hamilton, Jason G., James R. Griffin and Mark R. Stromberg, Long-term population dynamics of native Nassella 
(Poaceae) bunchgrasses in central California 

Haubensak, Karen A. (see Parker, Ingrid M.) 

Heidel, Bonnie (see Spribille, Toby, et al.) 


77) 


135 
299 


eS) 


165 


201 
54 


289 


130 


158 


186 


54 
58 


33 


198 


2002] TABLE OF CONTENTS 


Hrusa, Fred, Barbara Ertter, Andrew Sanders, Gordon Leppig and Ellen Dean, Catalogue of non-native vascular 
plants occurring spontaneously in California beyond those addressed in The Jepson Manual—Part I 

Jessup, Steven L., Reticulate ancestry in Mexican Gaudichaudia (Malpighiaceae) ______.-_-----_------ 

Jessup, Steven L., Six new species and taxonomic revisions in Mexican Gaudichaudia (Malpighiaceae) 

Johnson, Phillip (see Rice, Barry A., and Phillip Johnson) 

Jones, James S. (see Garrison, Barrett A., et al.) 

Kapiane Divhaldsice. Micticalomoray Ollie tO” oe ee Ee ee 

Karlik, John F, et al., Eugene D. Albertson, Y. Jae Chung, Alistair H. McKay and Arthur M. Winer, Field 
assessment of the California GAP analysis program GIS database in central California 

Keeley, Jon E. (see Boyd, Steve, and Jon E. Keeley) 

Koehler, Catherine E. (see Montalvo, Arlee M.) 

Leon de la Luz, José Luis (see Rebman, Jon) 

Leppig, Gordon (see Hrusa, Fred, et al.) 

Levine, Larry, Mary Bacca and K. O. Fulgham, Plant zonation in a Shasta County salt spring supporting the 
nly ows populatouvor Puccmellia howellit (POaCcac) a 

Makines sblizabeth. Notewormuy COUechHon trom ATIZONA, = 2 

Marsh, Glenda D., and Debra R. Ayres, Genetic structure of Senecio layneae (Compositae): A rare plant of the 
SUELO RINT I Ss es ce OE a = 58 Ne 0) Se eR ek ee he ee ee 

Mathiasen, Robert L., and Carolyn M. Daugherty, Adult sex ratio of Arceuthobium gillii (Viscaceae) 

McClaran, Mitchel P., and James W. Bartolome, Noteworthy collections from Moorea, Society Islands, French 
POLSTREST EL a2 sees eee a ad oa a ee NS eee nO ee 

McKay, Alistair H. (see Karlik, John F, et al.) 

Mishler, Brent D. (see Stark, Lloyd R.) 

Montalvo, Arlee M., Laura J. Feist-Alvey and Catherine E. Koehler, The effect of fire and cold treatments on 
seed germination of annual and perennial populations of Eschscholzia californica (Papaveraceae) in south- 
BOE Ge OES oe ee a a a Ee ee pe aera oa ta ie  ae 

Newhouse, Bruce, Richard Brainerd and Peter EK Zika, Noteworthy collections from Oregon _ 

Nobel, Park S., Erick De la Barrera, David W. Beilman, Jennifer H. Doherty and Brian R. Zutta, Temperature 
lintraviogs tor Cultivation, of edible cachiim: Califomia«.. 2 ae Se ee 

Oberbauer, Thomas A. (see Rebman, Jon) 

Parker, Ingrid M., Alexandra Engel, Karen A. Haubensak and Karen Goodell, Pollination of Cytisus scoparius 
(Fabaceae) and Genista monspessulana (Fabaceae), two invasive shrubs in California 

Parker, V. Thomas (see Vasey, Michael C.) 

Patten, Ann M., and Steven J. Brunsfeld, Evidence of a novel lineage within the Ponderosae 

Pivorunas, David, Veva Stansell and Peter E Zika, Noteworthy collection from Oregon 

Prigge, Barry A., A new species of Prunus (Rosaceae) from the Mojave Desert of California 

Rebman, Jon, Thomas A. Oberbauer and José Luis Leén de la Luz, The flora of Toro Islet and notes on 
Gradaimpodsiand. Baja Calera Meee er ne ee 

Paice inv ee INOLewortiny COMCciOnS 1oml) Calitonita,. 0... 0 ete ee ee Oe 

Rice, Barry A., and Phillip Johnson, Noteworthy collection from Alaska 

Roché, Cindy Talbott (see Gamarra, Roberto) 

Sanders, Andrew (see Hrusa, Fred, et al.) 

Sawyer, John O., Review of J/lustrated Field Guide to Selected Rare Plants of Northern California edited by 
Gary Nakamura and Julie Kiersteand Nelson 

Sawyer, John O. (see also Eckert, Andrew J.) 

SAVE Ie atMCeH NOL WOE COM ec itl: REmiy, VV AS CINTA LOIN a Sp ei 

Shaw, David C., and Steven A. Acker, Canopy macrolichens from four forest stands in the southern Sierra mixed 
SOARES CIEE SES Oe PILED cd) CEO SCO EN UNIT AS AEC a ALR 2 cect etter ser once 

Shevock, James R., Review of Field Guide to Liverwort Genera of Pacific Northwest America by W. B. Schofield 


Slauson, Liz A., Effects of fire on the reproductive biology of Agave palmeri (Agavaceae) 

SOLO keaEIN Ole WOOLEY CONCCHONS TtOnk Isao, nes. tr ee ee ce 

Sproulc, Lou, ct al. Noleworuny collections trom Montana 2 

Stansell, Veva (see Pivorunas, David) 

Stark, Lloyd R., Alan T. Whittemore and Brent D. Mishler, Noteworthy bryophyte records from the Mojave 
DSSERS oe po Reo a 2S es rr! 2 ee 

Stromberg, Mark R. (see Hamilton, Jason G.) 

Strother, John L. (see Greenhouse, Jeffrey A.) 

Strother, John L., and Bruce G. Baldwin, Hymenocleas are ambrosias (Compositae) 

Triepke, FE Jack (see Spribille, Toby, et al.) 

Triggs, Matthew L. (see Garrison, Barrett A., et al.) 

Tweed, William, Review of Seeing Things Whole: The Essential John Wesley Powell ed. William deBuys ___.. 

Vanderhorst, Jim (see Spribille, Toby, et al.) 

Vasey, Michael C., and V. Thomas Parker, Review of The Manzanitas of California, also of Mexico and the 
Oe PES EU Saas USE OS Dt ied BD ae ie D8 Pe ae 0 see ce 

Wachs, Robin L. (see Garrison, Barrett A., et al.) 

Wilson, Barbara L., and Sami Gray, Resurrection of a century-old species distinction in Calamagrostis —_... 

Winer, Arthur M. (see Karlik, John F, et al.) 


ill 


178 
132 


150 
12 


132 


207 
194 


228 


189 


285 


145 


193 


Whittemore, Alan T. (see Stark, Lloyd R.) 


York, Dana A., Eriogonum ovalifolium var. monarchense (Polygonaceae), a new variety from the southern Sierra 


Nevada, California 


Zika, Peter E, Noteworthy collections from Oregon and Washington 


Zika, Peter E (see also Newhouse, Bruce) 
Zika, Peter E (see also Pivorunas, David) 
Zutta, Brian R. (see Nobel, Park S., et al.) 


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